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Ecological Divergence and Convergence in Freshwater Fishes

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Ecological Divergence and Convergence in Freshwater Fishes KIRK O. WINEMILLER . IOLOGISTSAND POUCYMAKERSARE RAONG the clock to document EcologicalDivergence and preservebiological diversity in the faceof a growing glob- and Convergencein al human population and alterationsof naturalhabitats. Given FreshwaterFishes the magnitude and rapid pace of changesto natural ecosys- , biologists are challengedwith the taSkof identifying essentialfeatures in the organizationof biological communi- Strong relationships between basic ties. Notwithstanding the inherent uniquenessof different geographical fonn and ecologicalfunction in fISh- regionsand the evolutionaryhistories of their flora and fauna,the recogni- es allow the comparative study of tion of commonfeatures among communities and the speciesthat comprise ecological relationships using mor- them is sometimespossible. This study contributesto our understandingof phological features. Within-fauna biodiversity by further examining the ecologicalcharacteristics of natural variation and between-fauna simi- freshwaterfish communitiesfrom around the world. larities of ecomorphological charac- One of ecology'sgreatest challenges is to explain patternsof regional teristics of freshwater fish assem- speciesdiversity as well as the ecological,morphological, and behavioral blagesfrom five regions around the variation among and within populations. Patternsof ecologicaland mor- world were compared and contrast- phological diversification are of interest to both ecologistsand phyloge- ed. Ecomorphologicalfeatures netic systematists.Systematists use interspecificvariation as the raw mate- allowed identification of a number rial for constructing phylogenies. Ecologists would like to know if of ecologically convergent spedes interspecificvariation in a given trait arisesrandomly during evolutionary from different regions. Based on a divergencefollowing speciation, or whether this variation is correlated relative scale of convergence,tropi- with ecological function. This difficult task is further complicated by cal fish assemblagesexhibited a unique events.Traits adaptivein a particular ecosystemmay be neutral or greater degree of ecological conver- maladaptive in a new environment. If a maladaptivetrait is genetically gence than temperate assemblages. correlatedwith one or more strongly adaptivetraits, then it might persist The ecomorphological comparison in the population for many generations.In one form or another, evolu- supports the view that biotic inter- tionary biologists and ecologistsseek to identify, interpret, and model pat- actions have a greater relative terns of functional (adaptive) variation.21Similarities in patterns of indi- influence on evolution in the tropics vidual, population, and community-levelvariation amongdifferent faunas compared with temperate regions. provide evidence that, in many cases,the same deterministic processes organizebiological systems. Biological convergenceis the independentevolution of the samefea- ture (physiological,morphological, ecological,etc.) within two divergent Figure1. phylogenetic lineages. Divergence is the evolution of dissimilar traits The body shape.posidon of fins. and between closely related lineages.Remarkable examples of morphological nwuth structure of afish influenceits eco- and ecological divergence are well-known in vinually all higher taxa. logicalperformance. Trout possessa gen- Examples from the independent adaptive radiation among marsupial eralited nwrphologythat allows them to mammals of Australia and eutherian mammals of the other continents usethe endre water columnand a wide variety offood items in habitats that con- appearin most introductory biology textbooks.Convergence of ecological tain reladvelyfew compedngspecies. niches is seenin rodents and marsupial mice; shrewsand marsupialinsec- PAlNTING BY WALTER A. WEBER Q NGS tivores; and eutherian canines and marsupial wolves. The bony fishes 308 NATIONAL GEOGRAPHIC RESEARCH &. EXPLORAllON 8(3):308-327; 1992 I am preparing a report on (Osteichthys) are a much older and speciosephylogenetic lineage and as patterns of resource utiliza- might be expected,exhibit numerous examplesof evolutionary conver- gence throughout the world (Figure 1). Within the Characiformes, tion and guild organization Curimata (=Steindachnerina) argentea is an epibenthic mud-feeder; among diversefish assem- Hydrocynusvittatus is a large, roving midwater piscivore;Metynnis argen- blagesin Central and teus is a midwater planktivore/granivore;and Charaddiumfasciatum is a South America, North small, bottom-resting invertebrate-picker.Within the PerCidae,Etheostoma chlorosomumis a small, bottom-resting invertebrate-picker; and Perca America and Africa. In the flavescensis an epibenthic and mid water insectivore/piscivore.A remark- interest of communicating able convergence of body form and ecological function has occurred with a diverse scientific between the phylogeneticallydistant Characidiumand Etheostomaspecies audience,I have attempted (cylindrical body with flattened ventrum, broad pectoral fins, pelvic fins to make the analysis fairly locatedanteriorly; subtenninal mouth, etc.). Despitethe number of instancesof remarkableconvergence that can be straightforward. This is identified within most large taxonomic groups (Figure 3), the conver- sometimesdifficult to gencephenomenon is difficult to study quantitatively;This is especiallyso achievewith numerical for small, closely related groupings. Our ability to recognizeevolutionary analysesaimed at revealing convergenceis entirely dependenton knowledgeof phylogeneticrelation- ships. Becausewe usually have more confidence in our phylogenetic temporal trends within groupings at broader taxonomic scales, ecological convergencecan be speciesassemblages con- identified with greaterease. Within smaller and more recent phylogenetic taining 30 to 70 species. lineages,it is often difficult to distinguish divergenttraits from convergent CORRESPONDENCEtraits. In fact, a great deal of the methodology of modern phylogenetics seeks to identify the suite of traits that is uniquely divergent versus the suite that is convergent.2Phylogenies are basedon the information con- tained in uniquely divergent (derived) characteristics.Convergent traits are seen as potentially confounding characteristicsin phylogentic con- structions. Among closely related taxa, the distinction between derived and convergent traits is sometimesunclear and only becomesapparent after phylogenetichypotheses have been generatedand tested. Here I present results from a comparative study of morphology and ecologyof freshwaterfish assemblagesfrom five widely separatedregions around the world. Study sitesspan a latitudinal gradientfrom Alaska (650 N) to the neotropics (90 N), and a tropical longitudinal gradient from Central America (830W) to Africa (230 E). Like many taxonomic groups, freshwater fIShesshow a strong latitudinal gradient in speciesrichness, with the highest regional diversity appearing in the neotropics. R. H. Lowe-McConneW5counted> 1300 freshwaterfish speciesfor the Amazon River basin comparedwith 192 for all of Europe.Since that time, new dis- coverieshave causedmost estimatesfor South American fIShesto more than double, while the record for European speciesremains essentially unchanged. Whereas climatic and biogeographicalhistory are probably the primary underlying agents that produce global patterns of regional diversity;the basicmechanisms that foster coexistenceof largenumbers of speciesin tropical faunasare open to debate.The extent to which assem- blagesfrom different regions are organizedin the samemanner indicates the degreeto which detenninistic processesregulate the populations that comprisecommunities.33 Moreover, the relativeamount of ecologicalcon- vergenceamong taxa from different regionsshould indicate the strength of KIRK O. WINEM1U.ER, assistant professor. Departmentof Wildlife and Fisheries similar forms of directional selection in relation to random evolutionary Sciences.Texas A&M University. College changeand evolutionary stasisdue to geneticconstramts. Here I contrast Station.TX 77843-2258. patterns of morphological divergencewithin the five local fISh assem- 310 RESEARCH &. EXPLORAnON 8(3); 1992 Curimata Hydrocynus Metynn;s Characidium Etheostoma ~ Figure 2. Phylogenyshowing evolutionary diver- gencr.within chtuadform and perdform fishes, as well as ecologicalconvergence betweenthe lebiasinidgenus of small, bottom-resting.invertebrate-pickers, Cbaracidium (C. fasciatum,South AmtTica),and the percidgenus of small, blages. A substantial literature illustrates the correlations between a num- bottom-resting.invertebrate-pickers, ber of basic morphological features of fishes and ecological performance Etheostoma(E. chlorosomum,North (summarized in P:W Webb21),and morphology has been used successful- America). ly as a surrogate for ecology.6-8.11,20.26.28.32In addition, I compare the rela- tive degree of ecomorphological convergence between fish species from different regional faunas. Methods SAMPLE COLLECTION Each of five study sites was sampledon at least two separatedates, and most locations were visited numerous times as part of long-term field studies.29-31Only numerically dominant specieswere used for the mor- phological analysis. Dominant speciesare defined in this study as the most common speciesat a site that (by number of individuals), when summedby order of rank. formed 99% of all individual fIShescollected over the full samplii1gperiod. Rare specieswere defined as those com- prising the 1% tail of smallest relative
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