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On Ecological Fitting Author(S): D Nordic Society Oikos On Ecological Fitting Author(s): D. H. Janzen Source: Oikos, Vol. 45, Fasc. 3 (Dec., 1985), pp. 308-310 Published by: Wiley on behalf of Nordic Society Oikos Stable URL: http://www.jstor.org/stable/3565565 . Accessed: 13/09/2013 19:33 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Wiley and Nordic Society Oikos are collaborating with JSTOR to digitize, preserve and extend access to Oikos. http://www.jstor.org This content downloaded from 169.237.66.239 on Fri, 13 Sep 2013 19:33:00 PM All use subject to JSTOR Terms and Conditions OIKOS 45: 308-310.Copenhagen 1985 Dan Janzen'sthoughts from the tropics1 On ecologicalfitting I've grownup on a dietof Darwin,Wallace and their occupiedby specieswith the following cyclical pattern culturaloffspring. But I'm facedwith the reality of the to theirhistories. Furthermore, these species are in the biologyof 100 km2of lowlanddry forest (Santa Rosa evolutionarilyquiescent stage in thecycle during their NationalPark) in northwestern Costa Rica. Someof my occupancyof the region.Initially, a speciesis a small backgrounddoes notprepare me for this: almost all the populationoccupying a small area. Sucha restrictedoc- ecologyI see aroundme couldquite easily come to be cupationoccurs through foundling establishment, habi- withvirtually no evolutionhaving occurred at Santa tat fragmentation,habitat shrinkage, etc. This popu- Rosa. lation evolutionarilypasses througha successionof Santa Rosa is a veryordinary place. The fine-scale genotypes.This change occurs because the habitat occu- mosaicof habitatsranges from 30-m-tall nearly ever- piedchanges and becausethe small population is being greenforest to 2-m-talldeciduous thorn scrub, with a evolutionarilyfine-tuned by selectionto theparticular varietyof anthropogenicsuccessional seres. There is a traitsof the habitatit occupies.Then, the population 0-350 m elevationrange, 900-2300 mm annualpre- abruptlyexpands out of its local habitat in theduration cipitation,and twodry seasons, one ofwhich is about6 ofa fewgenerations. The expansionoccurs because the monthslong and usually rain-free. Santa Rosa's habitats evolvingpopulation happens upon some genotype that are quite literallycrawling with complex biotic inter- is serendipitouslyrobust with respect to thechallenges actions.The participantsnumber at least650 species of ofliving throughout some large geographic area. It may plants,3100 species of Lepidoptera, 200-plus species of also occurbecause some geographic barrier has been re- seed-predatorbeetles, 58 speciesof mammals, 250-plus laxed(or bypassed),or becausemajor abiotic changes speciesof birds, and many more. It's decidedly tropical. makea smallhabitat into a widespreadone. The species The problemis thatat least98 percentof thesespe- is now widespread.By virtueof being huge, wide- cieshave geographic ranges covering tens of degrees of spread,on an adaptivepeak, and subject to a multitude latitude.While the majorityare commonand wide- of selectivepressures that are fine-scaleheterogeneous spreadin the dryNeotropics, another majority ranges andcontradictory, the species now becomes evolutiona- widelyover low elevationrainforest habitats as well. rilystatic. It is onlylikely to evolutionarilychange when Over theirwide ranges, these species interact in many a newgenotype appears that raises fitness throughout complicatedways with many other species that do not muchof the species'range. It thencontinues to be a occurat SantaRosa. However,these wide-ranging spe- widespreadspecies until some sortof majorperturba- cies are madeup ofsimilar, if not apparently identical, tionoccurs throughout its range,whereupon it is ex- individualsin quitedifferent parts of theirranges. tinguishedor reducedto somesmall population that is The implicationsof these observations are difficultto againevolutionarily labile. Likewise, during its tenure reconcilewith the commonplaceview of a speciesas as a widespreadspecies, isolates occur from time to evolutionarilylabile. This difficulty leads me to wonder time,and a veryminute fraction of these start the cycle ifa profitableexcursion in evolutionaryecology might again. be to explicitlyintroduce some heterogeneityinto the It appearsto me thatSanta Rosa is almostentirely abilityof natural selection to moldgenomes. As I write occupiedby species with such cyclical histories and that that,I suddenlyrealize that I have blunderedthrough theyare occupyingthe site during the widespread phase thefront door of the turmoil over punctuated equilibria. of theirhistories. Furthermore, it appears to me that We don'thave to dig at the fossilrecord; punctuated mostcontinental habitats are occupied as is SantaRosa. equilibriaare righthere in frontof us, representedby Whensuch a speciesbreaks out of its local status mostof thespecies that you and I haveanything to do somewhereand arrivesat a site such as Santa Rosa, with. whathappens? Think in termsof Africanizedhoney I am hereconcerned with any region that is primarily bees,cattle egrets, feral cattle and guanacaste trees (or 308 OIKOS 45:3 (1985) This content downloaded from 169.237.66.239 on Fri, 13 Sep 2013 19:33:00 PM All use subject to JSTOR Terms and Conditions gypsymoths, ring-neckedpheasants, feral house cats sures are sufficientlysimilar to maintain this single and kudzu). It willcome to occupysome, but not all the phenotype. habitatsat the site. If it is a plant,its flowerswill be pol- 2) The phenotypeis evolutionarilyfrozen and the linated to various degrees and qualities in different mothoccurs wherever its realized fitness is greatenough habitats.Likewise, its seeds will be dispersedto various for persistence. In some habitats, the eye spots are degrees and into various patternsof seed shadows, highlyfunctional in deterringpredators, but the mothis whichwill in turnimpinge upon a wide varietyof habi- absentbecause its larvalhost plants are absent. In some tats. Only some of these habitatswill containmembers habitats,the eye spots and the behaviourof displaying of theplant population. Its foliagewill be fedon to vari- themsimply lead to increasedpredation by (too) smart ous degrees in differenthabitats and its fitnesswill be predators,but the mothremains a commonmember of variouslyreduced in these differenthabitats as a conse- the fauna because there are massive amountsof para- quence. Of the habitatsit occupies, in some it will be sitoid-freehost plants. In some habitats,the eye spots verycommon and breed, in some it willbe less common are highlyfuntional and thatis whythe mothis present. and breed, and in othersit willbe presentonly as strays Finally,in some habitatsthe eye spotsare simplyirrele- (be they adults or juveniles). These various levels of vant because thereare no significantvisually-orienting habitatoccupation will also be generatedby the more predators,but the mothis onlyoccasionally present be- traditionallyconsidered variations in physicalenviron- cause only in some years is there enough rain for the ment and impact of other plants. If an animal, all the host plant's seeds to germinate.So, when a mutantap- above kinds of interactionswill occur, and with the pears thatmodifies the eye spot so as to raise the moth's same consequences. In short, our newcomer will be fitnessin but one of the many habitatsit occupies, it patchilydistributed, variably abundant, and have differ- seems reasonablethat it does not spread throughoutthe ent interactionswith different organisms in the various moth'srange. However, it also seems likelythat such a habitatsof the Park. mutantmight persist in some small isolate of Automeris No evolutionarychange was necessaryor likelyfor io occupyingonly one habitat. our invadingspecies to incorporateSanta Rosa into its If thisapproach is not all nonsense,then in determin- range,and to develop the heterogeneousmicrodistribu- ing the processes that lead to structurein habitatswe tion and microexpressionof its fitnessthat it now has. should concernourselves with the ecological outcomes Furthermore,there is no reason to expect that the es- of successiveintroductions into a habitat,as well as with tablishedresidents did anythingother than ecologically the evolutionaryadjustments among species. There is, readjustto thisact of ecological fitting.All imaginable then,a strongesoteric rationale for the study of habitats levels of readjustmentmay occur, from going extinct to richin introducedspecies. On the one hand, it is tempt- no response.Almost all thecomplex interactions now at ing to criticallynote thateasily 99 percentof the papers Santa Rosa maybe nothingmore than the consequences in esotericecological journals today deal with"native" of a long successionof ecological fittingsof one wide- animalsand plants. On the otherhand, easily the same rangingspecies afteranother. 99 percentapplies to widespreadorganisms in habitats While seeminglyinnocuous, trite and obvious, I have so modifiedby recenthumans that the organismsmight constructedan argumentthat roughlysuggests that a as well have been introduced.The rationalethat I pre- major part of the earth's surfacemay be occupied lar-
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