American Geographical Society

Oceanic Islands and Biogeographical Theory: A Review

Author(s): Jonathan D. Sauer
Source: Geographical Review, Vol. 59, No. 4 (Oct., 1969), pp. 582-593
Published by: American Geographical Society
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 OCEANIC ISLANDS AND BIOGEOGRAPHICAL
THEORY: A REVIEW
JONATHAN D. SAUER

T HE branchofbiogeographyconcernedwith life-formsandphysiognomicformations
was founded on the study of altitudinalzonation on mountains. These sharp local
patterns were soon joined with broad regional patterns under a simple theory of
climaticand, secondarily,edaphicdeterminism.Sincephysiognomy is supposedto show con-
vergence in similarenvironments,regardlessof origin, processesof migration and evolution
have been of slight interest, and island biotas have been given no special attention.
The modern development of the branch of biogeography concerned with species or
other taxa and with regional associationsof taxa began with the study of oceanic islands,
where climaticgradientsareweak and where the sharpest,most provocativepatternsdemand
ecological and historical explanations.The neatly delimited and relatively comprehensible
island biotas have evoked many hypotheses, some of which have been extrapolatedto con-
tinental regions. The comprehensive theory needed to explain the welter of geographical
patternsformed by the hundredsof thousandsof plant and animalspeciesis still incomplete.
Islandscontinue to be drawn on as heavily as ever for general models, which have come to
include mathematicalformulations.The boldest recent attempt in this line is a formidable
little book by Robert H. MacArthurandEdward 0. Wilson,' which is having a great impact
in some quartersand which requirescareful consideration.To put it in perspective, some
previous theory will be sketched as background.
EARLY BIOGEOGRAPHICALTHEORY

Islandshave been regardedboth as isolated microcosms and as samplesdrawn from the

world outside; conflicts between these opposing views, and attempts to reconcile them,
have been responsiblefor most of the theoretical interest in island biotas. When seafaring
naturalistsbegan to explore remote islands,they were preparedto find them populated by
products of special creation or at least by antediluvianrelics. In the wild state specieswere
supposed to be essentially static, both geographically and genetically, perhaps spreading
gradually and coherently around centers of origin or of refuge during ancient catastrophes.
The extent of natural long-range dispersalswas unappreciated,and land organisms were
assumed to be incapable of crossing oceans unless carriedby man. The explorers did find
some raraeaves among island creatures,of course, but most speciesturned out to be similar
to, and many were identical with, those on other islands and mainland coasts of the same
ocean basin. This was bound to stimulatespeculationabout common origins and thus about
processesof evolution and migration.
Three biogeographicalcomparisonsmade by CharlesDarwin in the Galapagosin 1835
led directly to what Sir Julian Huxley has called "the greatest of all revolutions in human

Robert H. MacArthur and Edward 0. Wilson: The Theory of Island Biogeography (Princeton,
1967).

)DR. SAUERis professorof biogeographyat the University of California,Los Angeles.

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 OCEANIC ISLANDS 583

thought, greaterthanEinstein'sor Freud'sor even Newton's, by simultaneouslyestablishing

the fact and discovering the mechanism of organic evolution."2 At first Darwin saw the
archipelago as a little world apart,most of its speciesbeing endemic, but when he compared
them with othershe had met earlieron the voyage, he was struckby the evident relationship
to South American species of quite differentenvironmentsand by their lack of resemblance
to those of the Cape Verde Islands,which in climate and geology are very like the Galapagos.
The last, crucial geographical clue was given to Darwin just before he sailed from the
Galapagos,when a local residenttold him that tortoisesfrom each islandwere recognizably
different.Darwin wrote that he had never dreamedthat islandsso similarand mostly in sight
of one another might be endowed with distinct creatures,and that he had been indiscrimi-
nately mixing specimensfrom differentislands. But fortunately some of his bird and plant
collections happened to remain unmixed, and he was able to look into the unlikely notion
and to discoverlocal modificationsof basicstocksthat are still classiccasesofspeciation. Many
additionalobservationsand much thought went into the maturationof his theory of evolu-
tion, but the idea that island endemics were derived from immigrant progenitorswas at its
core.
For Darwin, the primarycause of divergence of islandpopulationswas not isolation but
selection in unlike physical and biotic environments-in the Galapagos strong differences
between the islands and the mainland, subtle differencesamong the islands. He was well
aware that geographical separationcould promote divergence, but he rejected the facile
model promulgated by Moritz Wagner and his followers, which equatesa gap on the map
with absoluteisolation as a prerequisiteto speciation.There was uncanny agreementon this
score,just as on naturalselection,between the conclusionsreachedindependentlyby Darwin
and by the other great pioneer of island zoogeography, Alfred RusselWallace. Darwin had
a persistentcuriosity about long-range dispersal,including transportof seeds by birds and
ocean currents,and began observationsand experimentson this cryptic process,which others
have continued to the present. The question is no longer whether overseasdispersaloccurs,
but ratherits probability in differenttaxa over various distancesand routes.
Darwin, strongly seconded by Wallace, drew a clear distinction between continental
shelf islands,where drowning of former land connectionshas to be considered,and oceanic
islands,such as the Galapagos,built up as volcanic peaksfrom the sea floor. Another pioneer
biogeographer, Sir Joseph Dalton Hooker, who classifiedDarwin's Galapagosplant speci-
mens, agreed that the archipelago'sflora was derived from overseasmigrants, but he toyed
with the idea that other oceanic island floraswere relics isolated by sinking of land masses.
Other writers have since spun virtual webs of land bridges acrossthe world's oceans, usually
as ad hoc explanationsfor disjunctrangesin particulartaxa. The variousconstructshave not
generallyconverged, either in time or in space,and many are geologically fantastic.3Clusters
of drowned atolls and guyots indicate possible stepping stones for Cretaceousand Tertiary
migrations in certain oceanic regions, but continental drift or other crustalmovement suf-

2Julian Huxley: Charles Darwin: Galapagos and After, in The Galapagos: Proceedings of the
Symposia of the Galapagos International Scientific Project (edited by Robert I. Bowman; Berkeley and Los
Angeles, 1966), pp. 3-9; reference on p. 3.
3 For a
confrontation between some taxonomists and geologists on this issue, see J. Linsley Gressitt,
edit.: Pacific Basin Biogeography: A Symposium (Tenth Pacific Science Congress; Honolulu, 1963), pp.
193-253.

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584 THE GEOGRAPHICAL REVIEW

ficientto bridgeoceanbasinsdoesnot seemto fit withintheprobabletimespanof thetaxain

question.
OF ISLANDSAS UNITS
BIOTICCOMPARISONS
In spiteof somedurablecontroversies, moderntheoryof islandbiogeographytendsto
follow Darwinin lookingto ordinaryprocessesof migrationandevolutionratherthanto
catastrophic can, of course,be
eventsin a lost world. The doctrineof uniformitarianism
areignored,but it hasbeen
pushedtoo farif possibleresidualeffectsof pastenvironments
healthy for biogeographersto renounce them as explanations of first choice. Filtering of
immigrantsby differentialdispersalcapacity seems to explain adequatelythe famous biotic
impoverishmentand disharmonyof oceanic islands-that is, the limited number of species
and the absence of major taxa, such as mammals, amphibia, conifers, or dipterocarps,that
are presentin comparablecontinentalareas.The equally famous endemism of large oceanic
islands may be a consequenceof this filtering: from the relatively few initial stocks, suites
of speciescapableof exploiting the unoccupiedniches have evolved by adaptiveradiation.
Such generalizationsabout impoverishment,disharmony,and endemism of islandbiotas
emerged early from fragmentaryinformationon distributions.It became possibleto sharpen
and quantifythem for certaintaxaand regions asadditionalrecordsaccumulated.The number
of speciesper islandhas commonly been graphedas a function of some unitary characteristic
of the islands,such as area.If such regressionsare plotted for physicallysimilarislandswithin
a given region, they show that speciesnumberstend to increasewith islandsize at a regularly
diminishing rate, so that if logarithms are used the average curve approaches a straight
line. This resemblesthe familiarspecies-areacurve obtained by sampling nonisolated areas
of differentsize, but the islandcurve is steeper.In a mathematicallybrilliantand biologically
cloudy treatment of such curves, F. W. Preston suggested that any isolated region should
have an equilibriumspeciesnumber strictly relatedto the total number of individualsof all
species that it can support.4He pointed out that at equilibrium small isolates should have
fewer species than nonisolated sample areasof the same size, which are constantly enriched
by immigrantsfrom adjacentareas.Thus an island with all the speciesit could carry might
still appear impoverished by comparison with a continental area flooded with transients.
Whether a given islandis in fact saturatedwith speciesis a questionwe are in no position to
answer by theoretical calculations. Quantification of the species-area relationship is still
empiricaland will always be impreciseinsofaras other variablesaffectingspeciesnumber are
uncontrolled.Pure areaeffects are hard to sort out from effects of increasingenvironmental
diversity as the sampledareais expanded. When whole islandsare taken as units, only crude
statisticalcontrol of environmental diversity can be achieved, for instance by classifying
islands as high or low.

ECOGEOGRAPHICALANALYSIS OF ISLAND BIOTAS

Meanwhile, a different approach-namely, attention to local species distributions in

diverse island habitats-has revealed increasingly definite coalitions. For example, such

4 F. W. Preston: The CanonicalDistribution of Commonness and Rarity, Ecology, Vol. 43, 1962,
pp. 185-215 and 410-432.

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 OCEANIC ISLANDS

coalitionswere clearly recognized by A. F. W. Schimperin a monumental work on pioneer

plants of tropical seashores.5The capacity of these species for dispersalby ocean currentsis
roughly relatedto how frequentlytheir characteristichabitatsare disturbedby the sea. Some
outpost species of storm beacheshave seeds capable of floating acrossthousandsof miles of
ocean, and their rangesarenaturallypantropical.Lessextreme pioneersof mangrove swamps
and beach-ridge forests are generally adaptedfor drift dispersalwithin a single ocean basin
or a part of it. Like the wide-ranging sea birds, the pioneer coastal flora contributesa cos-
mopolitan element to island biotas, subject to weak filtering by distance and to little local
evolutionary divergence. Some coastal pioneers have spread inland or have been taken to
distantregions by man as weeds or crops (notably the coconut and the New World cottons),
but within their natural habitats the effects of human intervention have been minimal.
Some inland pioneers, particularlyfreshwateraquaticsand marsh plants,also have wide
and disjunct ranges on islandsjust as on continents; but they depend mainly on migratory
birds as dispersalagents and do not have the same geographicalpatterningas the coastal
pioneers. In contrast,speciesbelonging to climax rain forestsor to other stablecommunities
characteristicallyhave coherent, circumscribeddistributionsand are not adaptedfor regular
transoceanicdispersal.The colonization of the new islands of Krakatoasupplies a nice case
history of the rapidarrivalof coastaland inlandpioneersas comparedwith the poor showing
of rain-forestspecies,though it is not entirely clear how much filtering was effectedby dis-
persaland how much by differentialsurvivalin the raw habitats.6
A few of the endemics that dominate native forests and other stable communities on
ancient oceanic islandsmay have evolved from wide-ranging pioneers. However, most en-
demics are derived from less likely immigrantsbrought by freakishand irregulardispersals,
for example by raftingor straybirds. Some areconceivably relicsof speciesthat have become
extinct in the outsideworld, but their peculiaritiesaremainly explainableby divergentevolu-
tion in situ. Local adaptationto stable habitatsshould theoreticallyinvolve loss of dispersi-
bility, so that isolation and divergence would reinforce each other in a feedback process.
As would be expected of makeshift communities patched together from limited raw ma-
terial, they have been extremely vulnerable to invasion by exotics introduced by man.
Island endemics, particularlyland birds and forest plants, are strongly representedon lists
of recently extinct or endangeredspecies.
The subdivision of island biotas on the basis of habitat and dispersalhas helped explain
broad similaritiesand differencesbetween whole islandstaken as units. Low atolls and sand
cays, which offer almost nothing but coastal pioneer habitats,tend to have simple, cosmo-
politan biotas, largely irrespectiveof remoteness, size, age, or human intervention. High,
rugged islandshave, in addition to this common pool, individualisticcomplementsof inland
species, sensitive both in kind and in number to each island'slocation, physical character,
geological antiquity, and human history. Patternsof this sort have been brought out clearly
by Sherwin Carlquistin a seriesof comparativestudies of island biotas. In one of these, he
tabulated the probable modes of dispersalof immigrants ancestralto the native floras of

5 A. F. W.
Schimper: Die indo-malayische Strandflora, Botanische Mitt. aus den Tropen, No. 3, 1891.
6 W. M. Docters van Leeuwen:
Krakatau, 1883 to 1933, Annales duJardin Botanique de Buitenzorg,
Vols. 46-47, 1936.

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586 THE GEOGRAPHICALREVIEW

variousPacific islandsand computed percentagesof differentmodes for each island or archi-

pelago.7Quantitativedifferencesbetween dispersalmodes were shown to be relatedprimarily
to the physicalcharacteristicsof the islandsand secondarilyto their location.
Innumerableworkers have contributed other general insights of this sort. Concern
with dispersaland survival processesshaping species distributionson islandshas provided a
meeting ground for museum, field, and experimentalstudiesby diversetechniques.The dis-
tribution of intensivework, however, has been scattered,taxonomically and geographically,
and some crucial problems, such as plant-animal interactions,are almost untouched. The
mood of exploration of the unknown can still be strong even in such a mecca for scientific
expeditions as the Galapagos,where evidence of unsuspectedsymbiotic relationshipswas
found recently between the most conspicuousplants and animals on the islands, the giant
cacti and the giant tortoises.8Aldabrais also famous for its giant tortoises,which have been
looked at by numerous scientific expeditions, but a careful survey of availabledata on the
atoll, preparedas backgroundfor the Royal Society Expedition of 1967-1968, shows how
little is actuallyknown of tortoiseecology andhow much lessis known about other creatures.9
Some of the most dynamic and consequentialinterspeciesrelationshipson islands,those
between man and other organisms,have been leastadequatelyinvestigated.Expeditionshave
gravitatedtoward islandsleast affectedby man, such as the Galapagosand Aldabra.On in-
habited islands, ecologists and other biologists have traditionallyimmersed themselves in
the closest approximationsto naturalcommunities they could find. Thus the grand task of
describing and interpretingnaturaland artificialpatternsin island biogeography is still in
full swing and is far from completion.

MODELSFORBIOGEOGRAPHY

MacArthur and Wilson's book"?is not intended as a summary or outgrowth of this

gradualaccretionof knowledge, though they have previously contributedto it by work on
West Indian birds and Pacific ants, respectively.Their avowed purpose is, by using islands
as an opening wedge, to lead biogeographyout of the natural-historystageand to reformulate
it in terms of first principlesof population ecology and genetics. Their method is primarily
logical deduction from mathematicaland graphicalmodels, dipping into empirical data for
occasionalexamples.
Their startingpoint is Preston'streatmentof the relationshipbetween areaandthe equilib-
rium speciesnumber, at which the additionof new species by immigration or evolution is
balancedby extinctions." Postponing evolutionary processesto the end of the book, Mac-
Arthur andWilson present a graphicalmodel that relatesimmigration and extinction rates
on a hypothetical island to the number of species present: the more species, the lower the
rate of immigration of new speciesand the higher the rate of extinctions.The immigration

7 SherwinCarlquist:The Biota of Long-DistanceDispersal:Part 5, PlantDispersalto PacificIslands,

Bull. TorreyBotan.Club, Vol. 94, 1967, pp. 129-162.
8E. Yale Dawson: Cacti in the GalapagosIslands,with SpecialReferenceto Their Relationswith
Tortoises, in The Galapagos[see footnote 2 above], pp. 209-214.
9 David R. Stoddart,edit.: Ecology of AldabraAtoll, Indian Ocean, Atoll ResearchBull. No. 118,
1967.
I0 MacArthurand Wilson, op. cit. [see footnote 1 above].
" Preston, op. cit. [see footnote 4 above].

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 OCEANIC ISLANDS 587
andextinctioncurvesintersectat the equilibriumor saturation number.Immigration is de-
fined as the arrivalof a propagule,perhapsa matedfemalebird or a singleseedof a self-
pollinatingplant,whichbelongsto a speciesnot presenton theisland;successful
establishment
or reproduction is not required.Speciesmustbe countedasimmigrantsif absentat the be-
ginningand presentat the end of whateverintervalis used, and must be countedas ex-
tinguishedwhen the reverseis true. Immigrations cancelledby extinctions(or vice versa)
withinan intervalaredisregarded, but a speciesmay repeatedlyimmigrateand go extinct
in successiveintervals.
A similarmodelis then presentedto show differentimmigrationcurvesfor nearand
farislandsanddifferentextinctionratesforlargeandsmallislands;everythingelseis assumed
to be constant,includingthe pool of speciescapableof reachingthe islands.Sincelow im-
migrationratesare shown for far islandsand high extinctionratesfor smallislands,the
equilibriumnumbersarelowestfor small,remoteislands.Also,sincethe equilibrium num-
bers are low on far islands,their logarithms decreaserelatively rapidly with decreasingarea,
which agrees with the theoretical effect of isolation on the slope of the species-areacurve
noted by Preston.i2Similarmodels are used to show how the size and spacingin hypothetical
islandclustersmight affectrelativeequilibriumnumbers.The authorsnext demonstratehow
the colonization curve for a new island-that is, the buildup of speciesnumber through time
-could be drawn by integratingthe differencethrough time of immigration and extinction
rates. The hypothetical example shows a rapid initial colonization rate, with the curve flat-
tening graduallyas equilibriumis approachedand thereafterlevelling off as a constantturn-
over rate is reached, with extinction of some speciesbalancedby replacementby others.
Up to this point in the book, species are treated as interchangeableand island biotas as
differing only in species number. In succeeding chapters, attention is shifted to qualitative
differencesbetween speciesin dispersal,in demography,and in ecological niches. The authors
speculatethat taxa may differnot only in mean dispersaldistance,but also in the form of the
frequency distributions of the propagules that reach various distances. These probability
distributionscan be uniform, exponential, or normal, depending on randomnessof pathway
and survival time of propagulesin transit.Mathematicaland graphicalmodels are used to
show how such differencesmight produce island biotas with radicallymixed geographical
origins (for example, New Guinea,which has primarilyAustralianvertebratesand primarily
Asian insects and plants). Other models show the relative importance of islandsof various
sizes and spacings that act as stepping-stones for propagules. These models suggest that
stepping-stonesare far more important for speciesin which dispersalof propagulesfollows a
normal frequency distribution,as would be expected with lost birds or drift seeds or rafts
liable to eventual sinking, than for species with exponential dispersal,such as long-lived
spores borne by a steady wind. This comparison assumesthat mean dispersaldistancesare
equal; the role of stepping-stonesdecreasesrapidly as dispersalpower increases.
A chapter on the strategy of colonization considersthe demographic characteristicsof
populationsin evaluatingprobabilitiesof successfulcolonization. It is assumedthatany colony
is subject to random fluctuationin numbers, and that extinction is therefore ultimately in-
evitable. By use of per capitabirth rate, death rate, and carrying capacity of an island for a
given species, models are formulatedto predict time to extinction. These models show that
I2
Ibid., pp. 197-201.

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588 THE GEOGRAPHICALREVIEW

the chances are excellent that descendantsof a single propagule will reach the maximum
population size that the island can carry, if the specieshas at least a moderate capacity for
reproductiveincrease.They also show that once a population reachesthis size, it has an ex-
cellent chance of surviving through geologic time, provided the island can carry more than
a few individualsof the species.Of course, the carryingcapacityfor a given speciesdepends
partly on the rest of the biota and changes as colonization proceeds.The authorsnote that
a speciesthat arrivedbefore the islandwas saturatedmay expandecologicallyinto temporarily
uncontestedniches. Even after the biota equilibrates,speciescan be expected to have larger
populationsand to be less compressedecologically thanin comparablemainlandareas,where
predators,competitors, and parasitesare more numerous. This effect should be stronger if
colonists are filtered through stepping-stoneislands,in which competitive selection among
ecologically similar speciesreduces the number and increasesthe relative diversity of those
passingthrough.
Finally,evolutionarychangesfollowing islandcolonizationare divided into three phases.
First come founder effects due to limited genetic diversity of the propagule and to shifts in
selective pressureduring the initial buildup of the population; these are assigneda relatively
minor and fleeting role. The much longer and more consequentialsecond phase concerns
adaptationof species to peculiaritiesof the island, a process that is exceedingly difficult to
generalize.The authorsdo not attempt to constructcomprehensivemodels but ratherspec-
ulate informallyabout the effect of islandlife on certaincommon evolutionarytrends,partic-
ularlycharacterreleasewhen speciesinvade areaswith few competitorsand characterdisplace-
ment when they meet new competitors. MacArthur and Wilson conclude that as species
evolve adaptationsto their new environmentsand associatesmore immigrantscan be packed
into an island ecosystem. Thus the equilibriumnumber may creep upward indefinitely, and
the initial models, in which evolution was excluded, representonly a quasi equilibrium.
Speciation is assignedto the third phase. The authorsadopt the standardmodel of al-
lopatric speciation: spatialseparationis the necessaryand sufficientcause of origin of two
speciesfrom one. Occurrenceof closely relatedspeciesin separatehabitatson the sameislandis
attributedto accumulationof immigrantsfrom neighboring islandsor other discreteoutside
sources. The authorsstate that, given enough time, all island populationswill evolve away
from the mother populationand from each other and will become distinctspecies.As a logical
corollary, they suggest that any nonendemic speciesare recent immigrantsand that the per-
centage of nonendemic speciescan thus be used to measureratesof extinction and replace-
ment in equilibrialbiotas.
The book closes with a prospect of biogeography being in a position to enter a new
experimentaland theoreticalphase.Experimentalintroduction of exotic speciesto islandsis
suggested,and also poisoning or removal of existing biotas on some small islandsto generate
artificialKrakatoas.Immigrationand other ratescould then be measuredin sufficientreplica-
tion to yield statisticallysound results.The authorsvisualize a meshing of new experimental
data with quantitativetheory that will galvanize biogeography and, expanding to include
continentsas well as islands,lead to a grandsynthesiswith ecological and evolutionarytheory.

EVALUATION OF THE MODELS

Before we analyzethis prospect,we should considerhow well MacArthurand Wilson's

approachservestheir basic purposesof launchingbiogeography into quantitativetheory and
of identifying the kinds of dataneeded for its furtherdevelopment. The book is most admir-

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 OCEANIC ISLANDS 589

able on the level of formal mathematicalargument, where any assumptionsthat are not
contradictory can be made, and any conclusions that are reached without internal error in
logic are valid. However, their constructscannot survive transferfrom this fairylandinto a
world where assumptionshave to be evaluated and conclusions have to be verified. This
would be irrelevantif some predeterminedset of relationshipswere the object of study, but
the stated goal is recognition of significantfactors in real species distributions.The authors
admit to certainweaknessesand cruditiesin their models, and the implicationis that these can
be remediedby tinkeringand polishing. The defectsseem to me to be more fundamentalthan
that.
The basic equilibriummodel assumesthat for a given islandimmigration and extinction
rates vary as functions of the number of species already present. The number at the start
of an interval does, of course, set a strict limit on the number of potential new immi-
grants that remain in a fixed pool and also on the number that could become extinct dur-
ing the interval, but these are merely semantic ceilings. Actual rates may fall anywhere
between these ceilings and zero. For predictingrateswithin these ranges,the number of other
speciesis totally irrelevantfor speciesunconditionallybarredfrom the islandby the absence
of suitablehabitatsand also for those uniquely adaptedto availableniches. The number of
other speciespresentis vaguely relevantfor specieswhose successis contingenton competitive,
symbiotic, or prey-predator relationships,but no sums of species numbers can properly
measurethese diverse and highly specific interactions,especiallywhen a speciesrepresented
by a dormant seed is counted equally with an ecological dominant. The immigration and
extinction curves shown in the basic model are not derived from mathematicalformulasbut
are artisticcompromisesbetween extremely wide possibilities.
MacArthurand Wilson proceed to comparativeequilibriummodels tor large and small,
near and far islands, in which they manipulateindividual parametersand keep everything
else constant.They assume,for instance,that distanceaffectsthe rate of immigration, but not
the pool of speciesthat can reach an island. In a concrete case, the artificialintroductionand
naturalizationof speciesin Hawaii are explained solely on the basisof an increasedimmigra-
tion rate, as if the pool of immigrantsand the extinction rateshad been constant.Islandsize is
supposedto affectextinction ratesbut not immigrationrates,and distanceis supposedto have
the converse effect. Actually, a tricky feedback that interferes with this neat distinction
between immigration and extinction is built into the models. A large target is likely to be
hit sooner and more often by propagules;if unsuccessful,thesewould be countedin the model
as repeatedimmigrationsand extinctionsup to the point where the islandwas so large that it
was never free of them. Similarly, extinction ratesas defined in the models are not actually
independentof distance;an islandclose to the sourceof the propagulesmight never be free of
them, and extinction would be zero. In any case, overall immigration and extinction rates
can be related only loosely to island remotenessand size, since remotenessis relative to the
source regions of each species,and size is an impure measureof habitatdiversity mixed with
direct areaeffects. It is no wonder that the regressionsof speciesnumber on island areasthat
the authors occasionally offer as empirical examples show distressingscatter; islands with
similarspeciesnumbersdiffer in size by severalordersof magnitude, and islandsof equal size
differwidely in speciesnumber, even within assortmentsof islandsselectedfor comparability.
The authorsare driven to invoke controls not included in the models to explain some of this
scatter.
The theoretical colonization curve, as an integral of immigration and extinction rates

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590 THE GEOGRAPIICAL REVIEW

in successiveintervals,sharesall these defectscompoundedthrough time. The curve presented

as an example assumesrates that are plausible for a pool of species whose propagules are
chronically unsuccessfulon the island. Other species would tend to produce qualitatively
differentcurves, depending on whether they were quickly and continuingly successful,as in
the case of coastal pioneers not subject to competitive displacement;or were temporarily
successful,as in the case of inland pioneers eliminated by succession;or were increasingly
successful,as in the case of speciesbelonging to communitiesthat take time to develop. Since
the proportionalrepresentationof each of these groups dependson the location andcharacter
of the island, the shape of the theoreticalcolonization curve can vary enormously.
The only actualcase history of island colonization cited, the familiarstory of Krakatoa,
does not follow the authors'predicted convex curve. From reports of various expeditions
between the eruption of 1883 and 1934, it appearsthat the buildup of bird species began
slowly, acceleratedduring the middle of the period, and then slowed again, while plant
colonization continued at a more or less steady rate.13The authorsreachthe plausibleempir-
ical conclusion, not anticipatedby their theory, that extinction rates actually declined as
vegetational successionproceeded.
In short, the equilibriummodel and its derivativessufferfrom extreme oversimplification
by treating islandsas functionalunits with no attention to internalhabitat diversity and by
treating species as interchangeablewith no allowance for genetic or geographicaldiversity.
This is not even good as a first approximation,becauseit filters out the interpretablesignal
instead of the random noise. The authorsare in such a hurry to abandonthe particularsof
naturalhistory for universalgeneralizationthat they lose the grand theme of naturalhistory,
the shaping of organic diversityby environmentalselection. A model that visualizesvarious
sizes of assemblagesof characterlessspecies on various sizes of featurelessplains is essentially
absurd, since it excludes the very basis of genesis and continued coexistence of multiple
species.
As noted above, the authorsdo later considersome aspectsof speciesdiversity and in so
doing reachwhat is certainlythe high point of the book, with some stimulatingand satisfying
formulations, such as those exploring effects of different dispersalpatterns.However, it is
impossible to insert these into the monolithic structureof the equilibriummodel. Neither is
it possible to integratethe evolutionary concepts with the rest of the theory. This is partly
becausethe authorsregardevolution and migrationas operatingon quite differenttime scales,
and partlybecausethey believe speciationrequiresessentiallycomplete geographicalisolation.
In most of the book, propagulesfrom a constant species pool are supposedto arrive on an
islandwith regularfrequencies.In the model of speciationby geographicalisolation, dispersal
and extinction are shut off for agesuntil speciationis completed.Then the deus ex machinaof
absolute isolation is conveniently banished so that dispersaland extinction can resume to
concentraterelatedendemicson single islands.In this awkwardscheme, the lesssaidthe better
about loss of dispersibilityor local adaptationwhile the endemics were evolving in their
native homes.
Beyond all the weaknessesin conceptual structurelie troubles with calibrationof the

'3 Later observations, not cited by MacArthur and Wilson, show no sign of deceleration in plant
colonization. See J. van Borssum Waalkes: Botanical Observations on the Krakatau Islands in 1951 and
1952, Annales Bogoriensis, Vol. 4, 1960, pp. 5-64.

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 OCEANIC ISLANDS 591

models so that they can be used for quantitativepredictions.The authorsnote that appropriate
observationaldata are presentlyscarcebut suggest that the book may help identify the kinds
needed in the future. Actually, measurementsof most of the terms used in their formulations
arenot scarcebut nonexistentand areprobablyunobtainablefor anythinglargerthan an islet.
The book offersone example of empiricallydeterminedextinctionrates,basedon comparison
of plant speciestabulatedin 1904 and 1916 on certainkeys in the Dry Tortugas.The percent-
ages of speciesseen in 1904 that were not seen in 1916 are plotted as functions of island area
and initial species number. The resultsare taken to representnaturalextinction rates in an
equilibrialflora, but the case leaves something to be desired.As the authorsnote, the surveys
were not carried out by comparable methods, and there is also a problem of sorting out
naturalextinctionsfrom those attributableto constructionactivities;they do not mention the
great hurricaneof 191o that alteredthe keys and even the surroundingseafloor, nor do they
take into account the ordinaryseasonalcomings and goings of some of the ephemeralbeach-
pioneer species.
MacArthur and Wilson recognize the impracticality of measuring immigration and
extinction ratesin their models but suggest that these ratesmight be deduced from variation
among islandsin equilibrialspeciesnumber and speciescomposition. They are also hopeful
about direct measurement of colonization rates. However, all these approachesrequire
synoptic tabulationsof entirebiotas.Availabletabulations,includingmost of those the authors
cite, are usually accumulationsof observationsover extended periods of time and, even so,
omit parts of the biotas. For an island of any size and complexity, a complete census-type
enumerationwould be a fantasticundertaking,particularlyif the criterionof speciespresence
were basedon propagulesratherthan on establishedpopulations.The whole approachsmacks
of ornithology and is reasonableenough for creaturesthat fly about advertisingtheirpresence,
especiallyif they areidentifiableby a body of volunteer watchers.However, enumeratingthe
entire biota would requirea massivecollectingjob to be undertakenby specialists.The entire
biomasswould have to be screened,the forestsfelled, and the soil sifted for seeds,spores,and
whatnot, with each tested for viability and cultured until mature enough to be identified.
Each census would produce the suggested artificialKrakatoaand whatever was left of the
biota could hardly be pursuedto the next census as an equilibrialsystem.
Measurementof other constantsand variablesin the theoreticalformulationsis even more
hopeless.To evaluatethe size of the speciespool capableof reachingan islandand the distance
propaguleshave to travel supposesomniscienceabout speciesrangesand dispersalbeyond the
island. Calibrationof the demographic models demands measurementsthat have not been
made on any naturalpopulationsand, if attempted,would so disturbthe populationsthat the
resultswould be artifactsof the investigation.

PROSPECT
With models incapableof organizing real data, it seems prematureto talk about galvan-
izing biogeography by meshing new experimentaldatawith quantitativetheory. MacArthur
and Wilson's book may contributeto theoreticaldevelopment simply becauseit is provoca-
tive and will stir up interest, but I believe that the approachthey have chosen leads into a
morass ratherthan into the heraldedbreakthrough.In a field as complex as biogeography,
pure deduction that starts with gross assumptionsand a few casually selected variables is
unpromising, no matter how skillfully done. Paradoxically, it resembles pure induction,

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592 THE GEOGRAPHICALREVIEW

startingwith thoughtlesslydesignedobservationsand experiments,in that discoveryof

significantrelationships is mainlya matterof luck.This is not trueof the middleground
whereempiricalobservations andworkinghypothesesconstantlyinteractthroughfrequent
takingof bearingsandcoursecorrections. Suchinteraction cutsoffthecollectionof uselessor
redundantdata,a vital economyin a field that lackssecondhanddatafrom government
bureausandcannotgeneratedatacheaplyfromquestionnaires or hiredhelp.Thebasicbitsof
informationon occurrence of speciesat pointsin timeandspaceoftendependon taxonomic
competencethatis a scholarlyaccomplishment of highorder.Itspossessors wouldbe miscast
as regimentedobservers,contributing rawdatato testor confirma prioriconstructs, rather
thanas participants in synthesis.Clearly,widerexpertparticipation in developingbiogeog-
raphywill be invitedif the focusis directlyon speciesdistributions and not on synthetic
regional indices.
We alreadyhave a familiarconceptualmodel of the generalmechanismthat shapes
speciesdistributions: thecentrifugaldispersal of propagules thattendsto expandthe rangeis
opposedby theforce of elimination, which is both random and contingenton environmental
variables.When dispersaland eliminationare in equilibrium,speciesbordersare static;
migrations areinitiatedby changesin dispersal or elimination, owingeitherto environmental
change or to evolution of the species itself.Insights into these processes do not requirecom-
plete, coordinated synopticsurveys but can be gainedgraduallyby independently conducted
and
sampling experiments, and can be progressively sharpened to whatever degreeseems
worthwhile.The modelcan be fully integratedwith evolutionarytheory,sincesuccessor
failureof propagulesdependson both theirlocationandtheirheredityandsimultaneously
affectsboththegeographical andgeneticpatternsof theirspecies.It canalsobe readilyrelated
to ecologicaltheory,sincecommunitystructure anddynamicsdependon, andacton, species
distributions.
In anintuitiveandinformalway, thisconceptof biogeographical patterning hasalready
been integratedwith evolutionaryand ecologicalmodelsandwas, in fact,crucialin their
formulation. Thetheoryof naturalselectionis stillbasedmoreon spatialcorrelations between
environmentalfactorsand biologicalvariationpatternsthan on genetic case histories.
Similarly,bioticcommunitiesandecosystemshaveusuallybeenrecognizedon the basisof
spatialassociations of speciesratherthan by detectinginteractionsbetweenthem. H. C.
Cowles'sclassical modelof primarysuccession leadingto climaxvegetationwaslinkedto the
Davisianerosioncycle;it wasinferredfromspatialcorrelations betweenspeciesandedaphic
factorsandwas not derivedfromhistoricalevidence.
Morerecentlyecologyhasdevelopedan independentempiricalbaseby statisticalsam-
plingmethodsthatdivergefromthegeographical traditionof dealingdirectlywithpatterns of
gradientsandborders.Ecologistscommonlyrecognizeabstract typesof associations,
ecosys-
tems,or habitatsandpool samplesfrom scatteredstudyareasthatthey regardas acceptably
homogeneousandrepresentative of thetype.MacArthur andWilsonhave,in effect,proposed
the substitution of a grossandsimplifiedversionof thisapproachfor the geographical treat-
ment of distributionpatterns.In theirequilibriummodels,whole islandsbecomesample
quadratsin which the only datarecordedarecountsof speciespresent.In termsof a data
matrixin whichthecolumnsrepresent differentsitesandtherowsrepresent differentspecies,
theylumpallsiteson anisland into a singlecolumn and sum the speciespresentin thecolumn.
All informationon patternswithinrowsis thrownout with the arithmetical bath.

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 OCEANIC ISLANDS 593

It is true that, compared to ecology, biogeography is in a primitive, nonquantitative

state, but the reason may be found simply in geography's persistentpreoccupationwith the
superficialphysiognomy of vegetational formations. In view of the few personscommitted
to studying distributionsof speciesother than man and his domesticates,the resultshave been
excellent, and we would be foolish to neglect our effective approachto ape the ecologists in
techniquesdevised for other ends. We might better copy their willingness to make firsthand
acquaintancewith interestingspeciesand to work up a body of quantitativedatafrom original
field observations.
At present, I believe, biogeography would accomplish more by using its concepts and
tools thanby redesigningthem. No matterhow cleverly derived, our models will remainsoft
and amorphousuntil calibratedwith real values.We need to work out enough solid casesof
species patternsand the processesshaping them to get beyond vague banalitiesand isolated
details.This is a grand enterprisein which any number canjoin. Eventually, work will have
to be pushed with some taxonomically difficult organisms in some unpleasanthabitats.At
the moment there are still plenty of easily recognized speciesand many beautifulislandsthat
no biogeographerhas claimed. There are good practicalreasonsfor continuing to draw on
islands for case studies, though there is no longer any excuse for segregating them from
continents in biogeographicaltheory.

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