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frontiersofbiogeography
vol.3,n3november2011
thescientificmagazineofthe InternationalBiogeographySociety
ISSN19486596freelyavailableathttp://www.biogeography.org/
thescientificmagazineoftheInternationalBiogeographySociety
volume3,issue3November2011 ISSN19486596
frontiers of biogeography is published by the International Biogeography Society (IBS), an international and interdisciplinary society contributingtotheadvancementofallstudiesofthegeographyofnature frontiersofbiogeographyisavailableonlineattheIBSwebsite:http://www.biogeography.org/html/fb.html frontiers of biogeography aims to be a forum for biogeographers and a way to disseminate research in biogeography to the general public;ourscopeincludesopinions,perspectives,andreviews,symposiaproceedings,letterstotheeditor,bookreviews,researchupda tes,interviews,andarticlesonhowtoteach,disseminateand/orapplybiogeographicalknowledge.Letterstotheeditorandsymposium proceedingsmayincludenovelanalysesoforiginaldatasets(seeeditorialinstructions).Manuscriptsshouldbesubmittedtofrontiersof biogeography@gmail.com.EditorialenquiriesshouldbemadetotheEditorinChiefatibs@mncn.csic.es. frontiers of biogeography usesapublicationagreementbasedontheCreativeCommonsschemetoensurethattheauthorsretainfull intellectualproperty(IP)rightsontheirwork,andthatthisisfreelyavailableforanynoncommercialuse.Underthisagreement,theIBS retainsonlythecopyrightofthejournalcompilationunderaCreativeCommonsAttributionNonCommercialNoDerivatives(CCANCND) license.TheauthorshavefullIPovertheirtextsunderanuniversalCreativeCommonsAttributeLicense(CCAL),beingabletodistribute theirwork(includingtheoriginalPDFs)activelyfromthedayofpublication,andpassivelyfromoneyearafter(seethefulllicenseinfor mationattheendoftheissue). youcanfindinformationabouttheInternational Biogeography Societyathttp://www.biogeography.org/;forthelatestjobannounce ments and other news please visit also the IBS blog (http://biogeography.blogspot.com/), and the IBS facebook group (http:// www.facebook.com/group.php?gid=6908354463).
editorialboard
deputyeditorsinchief:
editorinchief:
Joaqun Hortal Museo Nacional de Ciencias Naturales (CSIC), MichaelNDawsonUniversityofCalifornia,Merced,USA RichardFieldUniversityofNottingham,UK SpainandUniversidadeFederaldeGois,Brazil
associateeditors:
AntjeAhrendsRoyalBotanicGardenEdinburgh,UK JanBeckUniversityofBasel,Switzerland JessicaBloisUniversityofWisconsin,Madison,USA ChrisBurridgeUniversityofTasmania,Australia MarcusV.CianciarusoUniversidadeFederaldeGois,Brazil MarkusEichhornUniversityofNottingham,UK RoyErkensUniversiteitUtrecht,TheNetherlands CamillaFljgaardAarhusUniversity,Denmark DanGavinUniversityofOregon,USA MatthewJ.HeardBrownUniversity,USA DavidG.JenkinsUniversityofCentralFlorida,Orlando,USA FrankA.LaSorteCornelllabofOrnithology,USA RichardLadleUniversidadeFederaldeAlagoas,BrazilandOxford University,UK RichardPearsonAmericanMuseumofNaturalHistory,USA ThiagoF.RangelUniversidadeFederaldeGois,Brazil WillemRenemaNCBNaturalis,TheNetherlands NriaRouraPascualUniversitatdeGironaandCentreTecnolgic ForestaldeCatalunya,Spain SpyrosSfenthourakisUniversityofPatras,Greece
editorialassistant:
LaurenSchiebelhutUniversityofCalifornia,Merced,USA
advisoryboard:
Miguel B. Arajo Museo Nacional de Ciencias Naturales (CSIC), SpainandUniversidadedevora,Portugal Lawrence R. Heaney Field Museum of Natural History, Chicago, USA DavidG.JenkinsUniversityofCentralFlorida,Orlando,USA RichardLadleUniversidadeFederaldeAlagoas,BrazilandOxford University,UK MarkV.LomolinoStateUniversityofNewYork,USA IBSV.P.forPublicAffairs&Communications
InternationalBiogeographySocietyofficers20112012
President:LawrenceR.Heaney PresidentElect:RosemaryGillespie VPforConferences:DanielGavin VPforPublicAffairs&Communications:MichaelNDawson VPforDevelopment&Awards:GeorgeStevens Secretary:RichardField Treasurer:LoisF.Alexander Directoratlarge:CatherineGraham Directoratlarge:KathyWillis Studentatlarge:AnaM.C.Santos FirstPastPresident:JamesH.Brown SecondPastPresident:MarkV.Lomolino ThirdPastPresident:BrettR.Riddle FourthPastPresident:VickiFunk FifthPastPresident:RobertJ.Whittaker Upcomingmeetinghost(exofficio):KennethFeeley PastGraduatestudentrepresentative(exofficio):MatthewHeard
cover:Floweringredbuglosses(Echiumwildpretii,alsonamedtajinastesrojosinSpanish)infrontofMount Teide(Tenerife,CanaryIslands).PhotographbyAnaM.C.Santos.
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Speciesareacurvesandtheestimationofextinctionrates
The speciesarea relationship (SAR) is one of the longestknown,mostintuitiveandempiricallybest provenpatternsofbiodiversity(Arrhenius1921). Variousauthorsdeterminedtheoreticallythatthe SARcanbeapproximatedasapowerlawfunction (i.e.,S=cAzwhereSisspeciesrichness,Aisarea and c and z are constants; Preston 1962, May 1975,Harteetal.1999),withz0.25incontinen tal areas but higher when dispersal barriers are involved (e.g., island speciesarea relationship). Empirical data suggested lower z in continental areas(0.130.18)andvaluesupto0.35forisland systems (Rosenzweig 1995). Dengler (2009) re centlycametotheconclusionthatthepowerlaw fits empirical data best in most cases (see also Dengler & Odeland 2010). Various authors ob served further systematic variations of z, such as whenconsideringspatialscaleorsamplingdesign (Plotkin et al. 2001, Scheiner 2006, Tjrve 2006, Dengler 2009). Kinzig & Harte (2000) pointed out the difference between SAR and the endemics area curve (EAR), which considers only species endemictoapartoftheregionunderanalysis.So what could He & Hubbell (2011) report that was so novel and generally relevant about SARs to meritrecentpublicationinNature? Sinceareaseemsalwaystoaffectbiodiver sity,nomatterwhattaxon,systemorscale,SARs have frequently been used to estimate species richnesslossresultingfromanthropogenichabitat destruction,i.e.extinctionratesinaconservation context.Thelossofacertainamountofarealeads to fewer species existing in a region at least some regional extinctions occur and the shape of the SAR has typically been used to retrieve quantitative estimates of how many species will go(regionally)extinct. Providingempiricalevidencefortheextinc tionofaspeciesischallengingandestimatingex tinction rates across a community even more so (Ladleetal.2011,thisissue).Yetthisisneededfor many conservation applications, such as schemes foroffsettingbiodiversityloss(Curranetal.2011) or,notleast,forpoliticalargument.Itistherefore notsurprisingthatSARbasedestimatesofextinc tion have been welcome despite critical studies that often found lower extinction rates than pre dicted (e.g., Kinzig & Harte 2000). It was argued, reasonably,thatontopofimminentextinctionin some species, others will be doomed to future extinction because of reductions in their popula tion size, and that this extinction debt explains apparent misfits. Other sources of uncertainty of the SARbased estimates are the (often false) as sumptionof acompletelyinhospitablematrixbe tween remaining habitat patches (Koh & Ghazoul 2010)ortheuseofdefaultslopevalues(z)inthe absenceofsystemspecificfitteddata. He & Hubbell (2011) pointed out that a backward interpolation of SARs is a flawed con ceptofmeasuringextinctionrates(seealsoKinzig & Harte 2000). This is because the area gain neededtoencounterthefirstindividualofanew species (which shapes the SAR) is always smaller thanthearealossneededtoremovethelastindi vidual.Toshowthis,theyformulatedbothasspa tiallyexplicitsamplingprocesses(SARforfirsten counters, EAR for last encounters). They con cluded that SARderived estimates of imminent extinctionwillalwaysbetoohigh,unlessindividu als are randomly distributed (i.e., no aggregated occurrenceofindividualswithinaspecies),which isanunrealisticassumption.He&Hubbell(2011) also showed that the EAR is a good predictor of empiricalextinctionratesevenifnospatialaggre gation is modelled, which offers an alternative (butamorechallengingone)forestimatingimme diateextinctionofendemicsfromarealoss. He & Hubbell (2011) clearly acknowledged that there is an anthropogenic extinction crisis and that habitat loss causes extinction. Further more, they did not claim that small population sizes of remaining species could not lead to fur ther, lagged extinction (in He & Hubbells view, EARsmodelonlyimminentextinctionandsodo SARs, but wrongly). Despite this, He & Hubbell (2011) already anticipated that pointing out this error in estimating extinctions would not be 81
frontiersofbiogeography3.3,20112011theauthors;journalcompilation2011TheInternationalBiogeographySociety
newsandupdate greetedwithenthusiasmamongconservationists, andthecorrespondenceonthepaper(Evansetal. 2011, Brooks 2011; see also online comments at http://www.nature.com/nature/journal/v474/ n7351/full/474284b.html) seems to confirm that. Thepaperisviewedasirresponsiblyundermining conservationeffortsbyallowinganticonservation groupstoclaimthatthingsarenotasbadasprevi ously asserted (fossil fuel lobbying in the climate change discussion is cited as example of this tac tic). Conserving nature is not only about science, butitistoalargedegreepoliticsandcorrecting anerrorleadstobettersciencebutmightweaken political success. I think scientists must correct themselves and not hold on to preconceived ideas,evenifitcreatessuchdilemmas. However,He&Hubbell(2011)studiedarea effects as a sampling problem in continental re gions,whichisprobablyappropriateforcapturing immediate extinction in many conservation set tings which occur at the regional or landscape scale. It remains to be understood and tested whethertheirconclusionsthat(a)EARestimates extinction better than SAR (cf. Kinzig & Harte 2000,Pereiraetal.2012)and(b)zdifferssystem aticallybetweenSARandEAR(whichispresented confusingly)aregeneralities.Thusitremainsto be seen whether SARs always overestimate ex tinction,asHeandHubbell(2011)claimed.Afur ther task will be to quantitatively estimate how many more species may go extinct after a time lag: how large the extinction debt really is (see alsoPereiraetal.,inpress).Inthiscontext,itmay be worthwhile to thoroughly investigate under whichcircumstances,ifany,theconsequencesof area lost to habitat destruction could be under stood solely on the basis of island biogeographic mechanisms (Rosenzweig 2001) that is, species richness as equilibrium between immigration + speciationandextinction.Thespatialandtempo ralscalesofanalysis,amongotherfactors,maybe relevantforthis.Undersuchcircumstances,SARs mayestimatethenewequilibriumstate,account ingforimminentandtimelaggedextinctions.
JanBeck
UniversityofBasel,Dept.EnvironmentalScience (Biogeographysection),Basel,Switzerland. email:jan.beck@unibas.ch; http://www.biogeography.unibas.ch/beck
References
Arrhenius,O.(1921)Speciesandarea.JournalofEcol ogy,9,9599. Brooks, T.M. (2011) Extinctions: consider all species. Nature,474,284. Curran, M., De Baan, L., de Schryver, A.M., van Zelm, R., Hellweg, S., Koellner, T., Sonnemann, G. & Huijbregts, M.A.J. (2011) Toward meaningful end points of biodiversity in life cycle assess ment. Environmental Science and Technology, 45,7079. Dengler,J.(2009)Whichfunctiondescribesthespecies arearelationshipbest?Areviewandempirical evaluation. Journal of Biogeography, 36, 728 744. Dengler, J. & Oldeland, J. (2010) Effects of sampling protocol on the shapes of species richness curves.JournalofBiogeography,37,16981705. Evans, M., Possingham, H. & Wilson, K. (2011) Extinc tions:conservenotcollate.Nature,474,284. Harte, J., Kinzig, A. & Green, J.(1999)Selfsimilarity in the distribution and abundance of species. Sci ence,284,334336. He,F.&Hubbell,S.P.(2011)Speciesarearelationships alwaysoverestimateextinctionratesfromhabi tatloss.Nature,473,368371. Kinzig,A.& Harte, J. (2000)Implicationsof endemics area relationships for estimates of species ex tinctions.Ecology,81,33053311. Koh, L.P. & Ghazoul, J.(2010) A matrixcalibrated spe ciesarea model for predicting biodiversity losses due to landuse change. Conservation Biology,24,9941001. Ladle, T.J., Jepson, P., Malhado, A.C.M., Jennings, S. & Barua, M. (2011) The causes and biogeographi calsignificanceofspeciesrediscovery.Frontiers ofBiogeography,3,111118. May, R.M. (1975) Patterns of species abundance and distribution. In Cody M.C. & Diamond J.M. (eds.), Ecology and evolution of communities, pp.81120;BelknapPress,Cambridge(Mass.). Pereira,H.M.,BordadeAgua,L.&Martins,I.S.(2012) Geometry and scale in speciesarea relation ships.Nature,inpress. Plotkin,J.B.,Potts,M.D.,Yu,D.W.,etal.(2000)Predict ingspeciesdiversityintropicalforests.Proceed ings of the National Academy of Sciences USA, 97,1085010854.

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Preston,F.W.(1962)Thecanonicaldistributionofcom monness and rarity: Part I. Ecology, 43, 185 215. Rosenzweig,M.L.(1995)Speciesdiversityinspaceand time.CambridgeUniversityPress,Cambridge. Rosenzweig, M.L. (2001) Loss of speciation rate will impoverish future diversity. Proceedings of the National Academy of Sciences USA, 89, 5404 5410.
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Scheiner,S.M.(2003)Sixtypesofspeciesareacurves. GlobalEcologyandBiogeography,12,441447. Tjrve,E.(2006)Shapesandfunctionsofspeciesarea curves: a review of possible models. Journal of Biogeography,30,827835.
EditedbyJoaqunHortal
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Extinctorextant?Woodpeckersandrhinoceros
Biogeographical research needs accurate data on thedistributionofspecies.Formanyspeciesthisis exceedinglydifficulttoobtain,leadingtoalackof globalinformationcollectivelyknownastheWal lacean shortfall. Fortunately, new tools are being developed that allow conservationists and bio geographerstodeterminetheexistenceofextant populationswithmuchgreateraccuracy. Foremost among these new tools is the in creasinguseofgeneticanalysis.Thiswasrecently used to great effect to confirm the extinction of theJavanrhinoceros(Rhinocerossondaicusanna miticus) in Cat Tien National Park in Vietnam (Brook et al. 2011). Despite their enormous size, Javan rhinoceros are remarkably shy forest dwelling animals that are difficult to see under natural conditions and were only rediscovered in mainlandAsiain1988.Giventhedifficultyoftra ditional surveying techniques, scientists from WWF and the Cat Tien National park had been monitoring the population by conducting genetic analysisofdungsamplescollectedintheparkbe tween2009and2010.Theanalysisindicatedthat all the dung belonged to a single individual, the bodyofwhichwasfoundApril2010,therebycon firmingtheextinctionofthepopulation. Of course, genetic analysis is costly, time consuming and requires some form of biological tissue(hair,dung,etc.).Formanyrareanimalsthe onlyinformationthatexistsistheoccasionalsight ing, the reliability of which is often highly ques tionable. Andrew Solow and his colleagues have recently come up with an ingenious method to account for this inevitable uncertainty (Solow et al. 2011). They use Bayesian (probabilitybased) statistics to model changes in the rate of valid sightings and to assess the quality of uncertain sightings for the ivorybilled woodpecker (Campephilus principalis) in North America. The woodpecker was controversially rediscovered in 2005, but a lack of clear documentary evidence and the failure of subsequent intensive surveys have led many scientists to doubt the veracity of this claim. The Bayesian model applied by Solow to68historicalsightings(29ofwhichwereclassi fiedasuncertain)stronglysuggeststhatthebirdis indeedextinct,andthe2005sightingwassadlya caseofmistakenidentity.
RichardLadle
FederalUniversityofAlagoas,InstituteofBiological SciencesandHealth,BrazilandOxfordUniversity, SchoolofGeographyandtheEnvironment,UK. email:richard.ladle@ouce.ox.ac.uk; http://www.geog.ox.ac.uk/staff/rladle.html
References
Brook, S., de Groot, P.V.C., Mahood, S. & Long, B. (2011) Extinction of the Javan Rhinoceros (Rhinoceros sondaicus) from Vietnam. WWF Report. Available at: http:// www.worldwildlife.org/who/media/press/2011/ WWFBinaryitem24584.pdf Solow,A.,Smith,W.,Burgman,M.,Rout,T.,Wintle,B. and Roberts, D. (2011), Uncertain sightings and the extinction of the ivorybilled woodpecker. Conservation Biology. doi: 10.1111/j.1523 1739.2011.01743.x
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Climatewars
Links between climate and societal instability, conflictandwarhaveincreasinglybeensuggested andanalyzed(Diamond2005),therebyfusingtra ditionally distinct academic disciplines such as (bio)geography, (agro)ecology and economics, history and peace research. Studies exploring these relationships are particularly pertinent in timesofanthropogenicclimatechange. Recent research has provided quantitative support for such climateculture linkages, but most of these studies have either been based on correlativeevidence(e.g.,Zhangetal.2007),ana lyzed shortterm climate fluctuations (e.g., Burke et al. 2009) or addressed specific hypotheses on the causes of human conflict (Beck and Sieber 2010).However,inordertomakeconflictpredic tionsunderclimatechangescenariosreliableand toengageinconflictpreventionormitigation,itis importanttobecertainaboutcausalrelationships and to fully understand the mechanistic links be tween past climatic changes and historical con flicts.Twonewstudieshaveattemptedthis. Hsiangetal.(2011)madeuseoftherecur ring yet irregular El Nio Southern Oscillation (ENSO) climatic changes as a natural experiment. Thisallowedthemtoshow,onaglobalscaleand foratimeperiodofmorethanhalfacentury,that (withinthesamelocalitiesandsocieties)civilcon flicts were more likely to arise during El Nio events as compared to La Nia periods. Further more, no such effect was observed for countries outsidetheENSOaffectedzoneoftheworld.This provides strong evidence that climate is indeed causaltotheseevents.However,theauthorscan only speculate on a variety of mechanisms for how(warmeranddrier)ElNioperiodscouldlead toconflict.Effectsmediatedbydecreasedagricul tural productivity and/or economic disturbance (e.g., resulting from increases in natural disasters and diseases) seem plausible, but psychological effects of unusual weather conditions on a large number of individuals may also increase a soci etysconflictpotential. Zhang et al. (2011) presented a detailed causality analysis based on a time series of cli matic fluctuations over a 300 year period in pre industrialEurope.Theyprovidestrongsupportfor theideathatclimaticvariationcausedfluctuations in agricultural productivity, and hence food avail abilityandprices.Thelatterwasidentifiedasthe root cause for a number of societal phenomena suchasmigrations,epidemics,populationgrowth and war. A temperaturebased model based on thesemechanismscouldsuccessfullypredictperi odsofcrisisandharmonyforpasteraswithless detailedhistoricalrecords. Animportantfuturedirectionofresearchin this field will certainly be the identification of natural factors and societal traits that explain variation around such climatedetermined pat terns. Demography and economic performance have sometimes been analyzed in this context (Samsonetal.2011,Hsiangetal.2011).However, itwillrequirethefurtherintegrationoftheabove mentioned disciplines to sort out the ultimate causes of why certain regions and/or societies navigated smoother and less violent routes through times of crisis than others (my current location, Switzerland, is a prime example within thelastfewcenturies).
JanBeck
UniversityofBasel,Dept.EnvironmentalScience (Biogeographysection),Basel,Switzerland. email:jan.beck@unibas.ch; http://www.biogeography.unibas.ch/beck
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Zhang,D.D.,Lee,H.F.,Wang,C.,Lie,B.,Pei,Q.,Zhang, J. & An, Y. (2011) The causality analysis of cli mate change and largescale human crisis. Pro ceedings of the National Academy of Sciences USA,108,1729617301. Zhang,D.D.,Brecke,P.,Lee,H.F.,He,Y.Q.&Zhang,J. (2007) Global climate change, war and popula tion decline in recent human history. Proceed ings of the National Academy of Sciences USA, 104,1921419219.
References
BeckJ.,&Sieber,A.(2010)Isthespatialdistributionof mankinds most basic economic traits deter minedbyclimateandsoilalone?PLoSONE5(5): e10416. Burke, M., Miguel, E., Satyanath, S., Dykema, J. & Lo bell, D. (2009) Warming increases risk of civil warinAfrica.ProceedingsoftheNationalAcad emyofSciencesUSA,106,2067020674. Diamond, J. (2005) Collapse: how societies choose to failorsucceed.Viking. Hsiang,S.M.,Meng,K.C.&Cane,M.A.(2011)Civilcon flictsareassociatedwiththeglobalclimate.Na ture,476,438411. Samson,J.,Berteaux,D.,McGill,B.J.,Humphries,M.M. (2011)Geographicdisparitiesandmoralhazards in the predicted impacts of climate change on human populations. Global Ecology and Bio geography,20,532544.
EditedbyRichardLadle
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Emergingresearchopportunitiesinglobalurbanecology
Biogeographershaveexaminedhowhumanactivi ties have affected patterns of biological diversity fromavarietyofperspectives,withspecialatten tion often given to oceanic islands. With the cur rent accelerating pace of environmental change, these effects are increasingly evident at global scales. Human industry, commerce, agriculture and transportation all have the potential now to affectnaturalsystemsgloballythroughanassort ment of drivers; primary among these are land use change, species introductions and climate change. Human activities and their consequences cometoauniquefocusinurbanareas,anexpand ing form of land use that is attracting increasing research attention from ecologists (Grimm et al. 2008).Urbanareascontainsimilarenvironmental conditions worldwide and act as a focal point for species introductions and extinctions. These hu mandominated environments offer unique op portunities to investigate the broadscale dynam ics of humanmediated biotic interchange (La Sorteetal.2007),itsconsequencesfordiversity (LaSorteetal.2008)andtheregionalfactorsand biologicaltraitsassociatedwithnativespeciesex tinctions (Hahs et al. 2009, Duncan et al. 2011). Urban areas typically contain spatially heteroge neouscollectionsofnativeandnonnativespecies (McKinney 2008); these unique assemblages can be examined based on their compositional (Niemeletal.2002)andphylogeneticstructures (Ricotta et al. 2009). Three nested sampling ap proaches are currently used to investigate urban systems at broad spatial scales: urban plots or transects, the entire urban matrix and the urban matrix embedded within a regional context (Werner 2011). Each sampling approach provides a unique inferential basis, although the third al lows for more refined interpretation, controlling forregionaldifferences. A recent study in Global Ecology and Bio geography adopts a novel perspective and exam ineshowavianassemblagessampledwithinplots ofintactvegetationinurbanandseminaturalar eas differ based on several common mac roecological relationships. Pautasso et al. (2011) compiled data on species composition and abun dance from all around the globe, although the majority of the samples are from Europe and NorthAmerica.Aprimaryfindingofthestudywas a lack of evidence for differences in the species area,speciesabundanceorspeciesbiomassrela 85
newsandupdate tionships between urban and seminatural locali ties. The number of exotic bird species in urban areas is low, suggesting that these relationships are defined primarily by native species in both environments.Thesefindingshighlighttheimpor tance of maintaining intact vegetation within ur banlandscapesandtheroleofurbandiversityasa tool for promoting conservation initiatives and biological awareness, as emphasized in many ur banecology studies. Nevertheless, the findings from Pautasso et al. (2011) contrast with current expectationsonhowurbanizationaffectspatterns of diversity, and should be a motivating factor in promotingfurtherresearch.Theincreasingpreva lence and quality of global data sources provides anexcitingbasistoexaminethestructureandde terminantsofthesemacroecologicalrelationships acrossmorerefinedtemporal,spatialandanthro pogenicgradients. By taking a global perspective, novel in sightscanbegainedontheuniquepositionurban areashave,bothasasourceforglobalchangeand as regions capable of maintaining important as pects of biological diversity. Global comparative studies also have the potential to bolster and re fine current recommendations about how to maintain biological diversity within human dominated landscapes. Specifically, the preserva tionorrestorationofpatchesofintactvegetation within urban areas is as valuable in maintaining basic macroecological patterns of avian diversity asconductingtheseactivitiesoutsideurbanareas. Importantly,thisworktakesthefocusawayfrom EuropeandNorthAmerica,wherethevastmajor ity of the research has been conducted, allowing foramoreinclusivesetofinferencesandrecom mendations. Urban data are becoming increas ingly available through remote sensing activities, citizen science initiatives and broader collabora tive efforts. Exploring how anthropogenic activi tiesareimpactingnaturalsystemsgloballyiscriti cal in supporting a truly comprehensive under standing of the current dynamics and longterm consequencesofglobalenvironmentalchange.
FrankA.LaSorte
CornellLabofOrnithology,Ithaca,NY,USA. email:fal42@cornell.edu; http://www.birds.cornell.edu/
References
Duncan, R.P., Clemants, S.E., Corlett, R.T., Hahs, A.K., McCarthy, M.A., McDonnell, M.J., Schwartz, M.W., Thompson, K., Vesk, P.A. & Williams, N.S.G.(2011)Planttraitsandextinctioninurban areas: a metaanalysis of 11 cities. Global Ecol ogyandBiogeography,20,509519. Grimm, N.B., Faeth, S.H., Golubiewski, N.E., Redman, C.L., Wu, J., Bai, X. & Briggs, J.M. (2008) Global change and the ecology of cities. Science, 319, 756760. Hahs, A.K., McDonnell, M.J., McCarthy, M.A.,et al. (2009) A global synthesis of plant extinction ratesinurbanareas.EcologyLetters,12,1165 1173. LaSorte,F.A.,McKinney,M.L.&Pyek,P.(2007)Com positional similarity among urban floras within and across continents: biogeographical conse quencesofhumanmediatedbioticinterchange. GlobalChangeBiology,13,913921. LaSorte,F.A.,McKinney,M.L.,Pyek,P.,Klotz,S.,Rap son, G.L., CelestiGrapow, L. & Thompson, K. (2008)DistancedecayinsimilarityamongEuro peanurbanfloras:theimpactsofanthropogenic activitiesondiversity.GlobalEcologyandBio geography,17,363371. McKinney, M.L. (2008) Effects of urbanization on spe cies richness: a review of plants and animals. UrbanEcosystems,11,161176. Niemel,J.,Kotze,D.J.,Venn,S.,Penev,L.,Stoyanov,I., Spence, J., Hartley, D. & Montes de Oca, E. (2002)Carabidbeetleassemblages(Coleoptera, Carabidae) across urbanrural gradients: an in ternational comparison. Landscape Ecology, 17, 387401. Pautasso, M., BhningGaese, K., Clergeau, P., et al. (2011)Globalmacroecologyofbirdassemblages in urbanized and seminatural ecosystems. GlobalEcologyandBiogeography,20,426436. Ricotta,C.,LaSorte,F.A.,Pyek,P.,Rapson,G.L.,Celesti Grapow, L. & Thompson, K. (2009) Phyloecol ogyofurbanalienfloras.JournalofEcology,97, 12431251. Werner,P.(2011)Theecologyofurbanareasandtheir functions for species diversity. Landscape and EcologicalEngineering,7,231240.

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Beyondtaxonomicalspace:largescaleecologymeetsfunc tionalandphylogeneticdiversity
Communityecologytraditionallyfocusesonhypo theticaldeductive and experimental approaches and often is criticized for narrowing our under standingofnaturetolocalidiosyncrasies,ignoring the importance of historical explanations. On the otherhand,approachestakenbymacroecologists and biogeographers have been excessively ex ploratory and correlative, with limited success in elucidatingthemechanismsresponsibleformany of the largescale patterns we observe in nature (see Gaston & Blackburn 1999, Ricklefs 2008 and references therein). Recognizing that both ap proaches can learn from each other is pivotal in the challenge of integrating data from different scalesinordertounraveltheecologicalandevo lutionary mechanisms that influence current pat ternsinbiodiversityandecosystemfunctioning. Species richness has been the most com mon metric used to represent all aspects of bio logical diversity (from genetic and taxonomic to phenetic diversity). However, species richness alone cannot describe the processes involved in species coexistence and ecosystem functioning and also does not describe properly the differ ences in community structure. In contrast, phy logenetic and functional diversities allow us to understand the relative importance of species composition in terms of evolutionary history and ecological similarities. Phylogenetic diversity (PD) is a biodiversity measure that accounts for the phylogeneticrelationship(henceevolutionaryhis tory)amongspecies,whereasfunctionaldiversity (FD) represents how species are distributed in a multidimensional niche space defined by ecologi caltraits. Phylogenetic and functional approaches to community ecology emerged as prominent fields of research in the last decade (Fig. 1), but some how independently and without much crossover in the first years. Early PD measures were pro posed as a tool to select conservation areas, but later the idea was extended to understand how communitiesareassembledfromaregionalpool. FD,whichinitiallywasconsideredtheholygrailof the biodiversityecosystem functioning agenda, alsowasrapidlyappliedasametricforinvestigat ing assembly rules (see Pavoine & Bonsall 2011). Howcouldmacroecologyandbiogeographybene fitfromthesetwoapproaches?Theanswerliesin understanding what FD and PD should represent andhowtheyrelatetoeachother:whilephyloge netic community ecology links evolutionary and biogeographic history to presentday ecology, functional diversity (as any traitbased approach) linksnichetheorytolargescaleapproaches,such as macroecology, biogeography or phylogeogra phy.Therefore,combiningecologicalandphyloge neticframeworkstoexplainlargescalepatternsof biodiversityisanimportantstep,takenrecently. Largescale studies involving PD and FD seems to be increasing at similar rates (Fig.1). Recently, it was shown that both measures can be decom posed into gamma (regional), alpha (local) and beta(turnover)components.Whereaslargescale studies and anyscale studies follows a similar trend for betaPD, there were few studies with betaFD (none at largescale). This is perhaps be cause biogeographers and macroecologists were more aware of evolutionary and historical hy potheses, so the conceptual framework of beta PDwaslikelytobeabsorbedfirst.Also,thiscould reflecttheassumptionthatcloselyrelatedspecies should be ecologically more similar than distant related species and, thus, PD should be a good surrogate for FD (in fact this is what most large and localscale PD studies used to assume). This traditionalassumptionisnowdebated(e.g.Losos 2008),andthesetwomeasuresmaybeviewedas complementary, rather than competing, ap proaches (Gmez et al. 2010, DinizFilho et al. 2011, Meynard et al. 2011, Pavoine & Bonsall 2011,Safietal.2011). WhilesomelargescalestudiesinvolvingPD andFDareexploratory(e.g.Meynardetal.2011) others have presented hypotheses and predic tions. Safi et al. (2011) investigated global pat 87
newsandupdate ternsofmammalPDandFDandfoundthatwhen controlling mammal assemblages for their evolu tionaryhistorythetropicswerecharacterizedbya FDdeficit.Thissuggeststhatmorespeciescanbe closelypackedintotheecologicalspaceintropical than in temperate regions (see figure 3 in their paper), a paradoxical situation in which competi tionseemstolimittraitevolutioninagroup,but does not decrease the cooccurrence of species with similar trait values (Wiens 2011). There are several nonmutually exclusive mechanisms that couldberesponsibleforthispattern(seeFigure1 in Safi et al. 2011). In temperate regions, for ex ample, if resources are limited, species need to occupywiderecologicalnichesinordertosecure theirenergydemandsandthereforecommunities would show signs of overdispersion in functional traits. In addition, high environmental heteroge
200
6
neity could also result in an overdispersion in FD because coexisting species could adapt and spe cializetothedifferentenvironmentalconditions. Some light has been shed on betaPD pat ternsbyGmezetal.(2010),studyingNeotropical Forestantbirdsatdifferentspatialscales.Ifspeci ationoccurredmainlyamongecoregions,thereis a lower probability of sister species cooccurring in the same ecoregion, resulting in phylogenetic evennessatthissmallerscale.Ifso,wewouldex pecthighspeciesturnover(taxonomicbetadiver sity) and low phylogenetic turnover (betaPD) amongecoregions,becausespecieswouldtendto becloserelatives.Analternativescenarioiswhen phylogenetic structure at the regional scale is a product of limited dispersal of lineages. In this casewewouldexpectbothhighspeciesturnover and high betaPD among regions, because each
180 160 140

4
Any spatial scale

FD 2 PD beta-FD 0 2007 2008 2009 2010 2011 beta-PD
published studies
120 100 80 60 40 20 0 1975
Large spatial scale

FD PD beta-PD
1980
1985
1990
1995
2000
2005
2010
year
Figure1.ThenumberofarticlespublishedinpeerreviewedjournalsindexedbyISIwithfunctionalandphylogenetic diversityinthetitle,abstractorkeywordsfrom1976to2010.Anyspatialscalemeansallstudiespublishedinallsub disciplines of ecology and evolutionary biology, irrespectively of scale. Large spatial scale are those studies con strainedbythesearchexpressionTopic=(geograph*ORmacroecol*ORbiogeogr*),thatis,thosestudiesmostlikely toberelatedtomacroecologyandbiogeography.FD=anystudywithtopicfunctionaldiversity;PD=anystudy with topic phylogenetic diversity; betaFD = any study with topic functional beta diversity or functional turn over;betaPD=anystudywithtopicphylogeneticdiversityorphylogeneticturnover.Theinsetisprovidedto showcurrentlystartingpublicationtrendsconcerningbetaPDandbetaFD.Therewasnolargescalestudyinvolving betaFDupto2010;butafewwerepublishedin2011orareinpress.
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newsandupdate region would contain distinct clades, with inde pendent diversifications. Finally, if observed val uesofspeciesturnoverandbetaPDdonotdiffer from what would be expected by chance (using nullmodels where random assemblages are built from the species pool), phylogenetic structure at the regional scale is unlikely to be the result of historical processes. In that case using FD should be better because nichebased processes are more likely to explain the pattern. For example, alongastrongenvironmentalgradientwherespe ciesaresortedfromtheregionalpoolaccordingto theirtraits,weexpectbothspeciesandfunctional turnover. However, if the species pool is com posed of ecologically similar species an indica tion that species were sorted according to their traitsatahigherspatialscale(forexample,dueto a climatic filter or historical processes) we should expect low functional turnover because the pool already contains very similar species. Also,intheabsenceofenvironmentalfilters,spe ciesturnovershouldoccurindependentlyoffunc tional turnover (Mouchet et al. 2010). Neverthe less,speciestraitsshouldhaveatleasttosome extentsomephylogeneticsignaland,therefore, partitioningtherelativecontributionofevolution ary history to trait dissimilarities among species may be important. A potential, and unexplored, solution is to decouple functional diversity into phylogenetic structured and specific (ecological) components. This would help us to betterunderstandhistoricalandrecentprocesses onbiodiversitypatternsandassemblyrules(Diniz Filhoetal.2011). The ground is reasonably well settled to start rebuilding community ecology from func tional traits (McGill et al. 2006) and merging community ecology with evolutionary biol ogy (CavenderBares et al. 2009). Yes, there are some methodological challenges how to prop erly define the species pool and null models, whichtraitsshouldbeused,whatisthemostsuit able measure of PD and FD, and so on (see Pavoine&Bonsall2011),butweshouldavoidbe coming locked into a blinkered debate about methodological issues. For example, in the last decademorethantwomeasuresofPDorFDwere proposed, each year! This may come at the ex pensesofthemoreimportant(andexciting)steps of doing science: how can we move forward the theorybyusingnovelapproaches? All existing hypotheses that have been ap plied to taxonomic diversity can be extended to phylogeneticandfunctionaldiversity(Meynardet al.2011).However,PDandFDcanbeusedtocre atemorerigorousanddirectpredictionsformost of the hypotheses in macroecology and biogeog raphy,suchasattemptstoexplainlatitudinalpat terns of biodiversity (Willig et al. 2003). These metrics also present an opportunity to formulate new hypotheses about how species evolutionary history and trait diversity are distributed across communities at different scales. For example, Wiensetal.(2011)showedsituationswhereafter amajorevolutionaryradiationwithinaregion,the region can still be invaded by ecologically similar speciesfromanotherclade,challengingthepara digm that communities are saturated. Large scale phylogenies and trait databases are cur rently becoming available for a wide range of taxonomicgroups,facilitatingestimatesofFDand PD. Including these two aspects of biological di versitywillbecrucialifwewanttoadvancefrom exploratory studies which report interesting rela tionships between biodiversity and environment toalsoidentifyingtheircausalmechanisms.
Acknowledgements
IthankJoaqunHortal,ThiagoRangel,andMichael Dawson for valuable comments on the manu script.ThisworkwassupportedbyCAPES(project #012/09).
MarcusV.Cianciaruso
DepartamentodeEcologia,InstitutodeCinciasBiol gicas,UniversidadeFederaldeGois,Goinia,GO, Brazil.email:cianciaruso@gmail.com; http://www.wix.com/cianciaruso/home
References
CavenderBares, J., Kozak, K., Fine, P. & Kembel, S. (2009) The merging of community ecology and phylogenetic biology. Ecology Letters, 12, 693 715.
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DinizFilho,J.A.F,Cianciaruso,M.V.,Rangel,T.&Bini,L. (2011) Eigenvector estimation of phylogenetic andfunctionaldiversity.FunctionalEcology,25, 735744. Gaston, K.J. & Blackburn, T.M. (1999) A critique for macroecology.Oikos,84,353368. Gmez, J.P., Bravo, G.A., Brumfield, R.T., Tello, J.G. & Cadena,C.D.(2010)Aphylogeneticapproachto disentangling the role of competition and habi tatfilteringincommunityassemblyofNeotropi cal forest birds. Journal of Animal Ecology, 79, 11811192. Jenkins, D.G. & Ricklefs, R.E. (2011) Biogeography and ecology: two views of one world. Philosophical Transactions of the Royal Society of London B, 366,23312335. Losos, J.B. (2008) Phylogenetic niche conservatism, phylogenetic signal and the relationship be tween phylogenetic relatedness and ecological similarity among species. Ecology Letters, 11, 9951003. McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. (2006) Rebuilding community ecology from functional traits. Trends in Ecology and Evolu tion,21,178185. Meynard, C.N., Devictor, V., Mouillot, D., Thuiller, W., Jiguet, F. & Mouquet, N. (2011) Beyond taxo nomic diversity patterns: how do , and componentsofbirdfunctionalandphylogenetic diversity respond to environmental gradients across France? Global Ecology and Biogeogra phy,20,893903. Mouchet,M.A.,Villger,S.,Mason,N.W.H.&Mouillot, D. (2010) Functional diversity measures: an overviewoftheirredundancyandtheirabilityto discriminate community assembly rules. Func tionalEcology,24,867876. Pavoine,S.&Bonsall,M.(2011)Measuringbiodiversity to explain community assembly: a unified ap proach.BiologicalReviews,86,792812. Ricklefs, R.E. (2008) Disintegration of the ecological community.AmericanNaturalist,172,741750. Safi, K., Cianciaruso, M.V., Loyola, R.D., Brito, D., Ar mourMarshall, K. & DinizFilho, J.A.F. (2011) Understanding global patterns of mammalian functional and phylogenetic diversity. Philoso phical Transactions of theRoyal Society of Lon donB,366,25362544. Wiens,J.J.(2011)Theniche,biogeographyandspecies interactions. Philosophical Transactions of the RoyalSocietyofLondonB,366,23362350. Wiens, J.J., Pyron, R.A. & Moen, D.S. (2011) Phyloge neticoriginsoflocalscalediversitypatternsand thecausesofAmazonianmegadiversity.Ecology Letters,14,643652. Willig, M.R., Kaufmann, D.M. & Stevens, R.D. (2003) Latitudinal gradients of biodiversity: pattern, process, scale and synthesis. Annual Review of Ecology, Evolution, and Systematics, 34, 273 309.
EditedbyThiagoF.Rangel
Remember that being a member of IBS means you can get free online access to four biogeo graphyjournals:Journal of Biogeography,Ecography,Global Ecology and Biogeographyand DiversityandDistributions.Youcanalsoobtaina20%discountonthejournalsOikosandJour nalofAvianBiology. Additionalinformationisavailableathttp://www.biogeography.org/.
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bookreview
Amangrovecompendium
Worldatlasofmangroves,byMarkSpalding,MamiKainumaandLornaCollins(editors) 2010,Earthscan,336pp.ISBN:9781844076574 Price:65(Hardback);http://www.earthscan.co.uk/
TheWorldatlasofmangroves,anupdatetoSpal ding et al. (1997), is a musthave publication for everyone loving and working with, in, or near to mangroves. It celebrates the wonderful world of these beautiful forests with astonishing figures and photographs. The informative maps and ta blesprovidecaptivatingfactsabouttheecological andeconomicvaluesofmangrovesandtheconse quencesoftheirloss. The atlas scores with the presentation of recent findings on carbon sequestration, showing that mangroves store more carbon than tropical forests(Donatoetal.2011);andwiththesuitabil ity of intact mangroves for protecting coastal re gions against tsunamis (Wibisono and Suryadipu tra2006).Thiswillarm(withpowerfularguments) ecologists, conservation biologists and policy makers, who urgently need to communicate this knowledge in order to increase public awareness and political willingness to protect and rehabili tate one of the most vulnerable ecological sys temsonearth. As indicated by its title, the World atlas of mangrovesgivesacomprehensiveoverviewofthe globaldistributionofmangrovespeciesatcountry level. A detailed description of the particular status of mangrove systems in each country, ac companied by information about their specific threats, level of degradation and extent of reha bilitation programs guides the reader through a multitudeofdistinctfeatures,whilekeepingsimi laritiesandgeneralprinciplesinmind. Mangrove experts of international repute contribute boxes on particular topics of interest, such as mangroves responses to climate change (Gilman,Dukeetal.)ortheirfunctioninginhighly dynamic coastal regions (Fromard and Proisy). They summarise uptodate research as well as the hot topics that will be developed in the near future. In addition, the annexes containing tree species descriptions, national species lists and countryfactsheetsserveasanexcellentcompen dium and make this atlas perfect as a quickstart guide for students as well as experienced re searchersapproachinganewregion. Considering the presentation of global trendsasthemainpurposeoftheWorldAtlasOf Mangroves, this book fulfils expectations. Unnec essaryuncertaintiesanderrorsintheintroduction to the ecology of mangroves leave, however, a drop of bitterness. The first chapters (Mangrove ecosystems and Mangroves and people) notably omit explicit references to any publications. The authors state that these chapters and the boxes therein draw heavily on the relevant literature, but information presented is confusing or even erroneous, and does not always reflect the con tentofthepublicationslooselymentionedatthe end of each subchapter, nor established knowl edgeavailableintextbooks(e.g.Tomlinson1986) or extended reviews (e.g. Feller et al. 2010). For example, the classification of mangroves into fringing mangroves, basin mangroves, and over washmangrovesisneedlesslyincomplete;itcould be easily improved by following standard man grove literature (e.g. Lugo & Snedaker 1974, Woodroffe1992).Theheterogeneoushandlingof outdatedtheoriesanddebatedhypothesesabout the functioning of mangroves is also surprising. For instance, the editors correctly do away with theperspectivethatthelandcreatesthecapabil ityformangroveformation,butthenpresentele vationandthesubsequentgradientofinundation as the only factors driving patterns of species zonation. There are, however, four other major hypotheses to explain this striking feature: geo morphological influences, propagule dispersal, predationandspeciescompetition(seee.g.Smith III 1992 for detailed discussion). Further errors in theclassificationofaeratingrootsandalsointhe systematicsandgeographicaldistributionofsome mangrove species have been already listed and 91
newsandupdate discussed in detail by DahdouhGuebas (2010). It remains a mystery why these chapters have not been written or carefully revised by the leading mangrove experts mentioned above, or the nu merousotherswhocontributedtothisbookwith specificboxes. This volume appears 14 years after Man groves The forgotten forest between land and sea(Mastaller1997).Itseemsthattheworldhas changedandtheforgottenforesthasbeenredis covered.Obviouslyneitherthesimpleexistenceof this remarkable ecosystem, nor its fascinating functioningbasedonadaptationtotheharshcon ditions of tidal zones, were sufficient to convince people that it is worth protecting mangroves againstaquaculture,agriculture,landuseandthe many types of waste water we produce. The monetary expression of the value of mangroves (US$ 20009000 ha1 yr1 according to the statis tics in this book), and the change from the eco logical perspective to the human perspective in termsofcoastalprotectionagainsthurricanesand tsunamis and in carbon sequestration, is neces sary to improve public awareness about the im portance of mangroves for our present life and a critical part of our response to the challenges of environmental changes, including sea level rise andclimatechange.TheWorldatlasofmangroves is a strong contribution towards this goal and, I hope,anothersteptowardsusheringinanewera where mangroves are valued for their beauty in thesamewayasmanyrainforestsorcoralreefs. In summary, if you are working in the field ofmangroveconservationorrelatedissuesinthe contextoftropicalcoastalzones,orifyourworkis targeted towards practitioners, stakeholders or usersofatriskmangroveecosystemservices,the Worldatlasofmangrovesisyourbook;itwillsup port your daily work with easytounderstand in formation and strong facts about the ecological and economic values of this forest. If you are a mangrove ecologist, this book should also be on your shelf because it provides you with a quick overview of mangrove distribution and current statusonEarth.Italsoactsasanenormoussource of suitable maps and material to round off your lectures. This should convince your students that 92 mangrove research is a challenge, an urgent de mand for mankind and that being involved is an accolade.Ontheotherhand,ifyouarelookingfor a general text spanning the interdisciplinary as pectsofmangroveecology,thisisnotthebookfor you.Therootsofthisbooklargelycomefromge ographyandremotesensing.Ifyouaresearching foranuptodatetextaboutthepresentscientific understanding and recent findings in mangrove research, I recommend supplementing the atlas with textbooks, recent reviews or more detailed publications on mangrove ecosystems and peo plesdepencyontheirhealthandfunctioning.
UtaBerger
InstitutfrWaldwachstumundForstlicheInformatik, TechnischeUniversittDresden email:uta.berger@forst.tudresden.de; http://www.forst.tudresden.de/SystemsAnalysis/utaberger
References
DahdouhGuebas,F.(2011)WorldAtlasofMangroves: MarkSpalding,MamiKainumaandLornaCollins (eds).HumanEcology,39,107109. Donato,D.C.,Kauffman,J.B.,Murdiyarso,D.,Kurnianto, S., Stidham, M. & Kanninen, M. (2011) Man groves among the most carbonrich forests in thetropics.NatureGeoscience,4,293297. Feller,I.C.,Lovelock,C.E.,Berger,U.,McKee,K.L.,Joye, S.B. & Ball, M.C. (2010). Biocomplexity in Man grove Ecosystems. Annual Review of Marine Science,2,395417. Lugo,A.E.&Snedaker,S.C.(1974).Theecologyofman groves.AnnualReviewofEcologyandSystemat ics,5,3964. Mastaller,M.(1997)Mangrovestheforgottenforest betweenlandandsea.TropicalPressSdn.BhD. KualaLumpur,Malaysia.189pp. Smith III, Th.J. (1992). Forest Structure. In: Tropical mangroveecosystems(ed.byA.I.Robertsonand D.M. Alongi), pp.101136. American Geophysi calUnion,Washington. Spalding,M.,Blasco,F.&Field,C.(1997).Worldman grove atlas. The International Society for Man groveEcosystems,Okinawa,Japan.178pp. Tomlinson,P.B.(1986).Thebotanyofmangroves.Cam bridgeUniversityPress,Cambridge,UK.419pp. Wibisono,I.T.C. & Suryadiputra, N.N. (2006). Study of lessons learned from mangrove/coastal ecosys tem restoration efforts in Aceh since the tsu nami. Wetlands International Indonesia Pro gramme,Bogor.86pp.
ISSN19486596
Woodroffe,C.D.(1992).Mangrovesedimentsandgeo morphology. In: Tropical mangrove ecosystems (ed. by A.I. Robertson and D.M. Alongi), pp.7 41.AmericanGeophysicalUnion,Washington.
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EditedbyMarkusEichhorn
bookreview
A comprehensive foundation for the application of biogeogra phytoconservation

Conservationbiogeography,byRichardJ.LadleandRobertJ.Whittaker(editors) 2011,BlackwellPublishing,301pp.ISBN:9781444335033 Price:95(Hardback)/34.95(Paperback);http://eu.wiley.com/
Itisbecomingincreasinglyclearthatthediversity of plant and animal species in the world is con tinuingtodeclineinspiteofambitioustargetsset by governments to prevent this (Butchart et al. 2010).Itisalsobecomingevidentthatthecontin ued functioning of ecosystems depends on this diversity (Isbell et al. 2011). In order to conserve whatisleftofbiodiversity,itiscrucialthatweun derstandthediversityoflifeandhowitisdistrib uted across the biomes and ecosystems of the world.Sinceunderstandingthedistributionofbio diversity is a central tenet of biogeography, it seems obvious that the field of biogeography shouldbeofcentralimportanceinconservation. In this volume, Richard Ladle and Robert Whittakerbringtogetherchaptersbyanumberof biogeographerstosummariseprogresstodatein applyingtheprinciplesofbiogeographytoconser vation and to identify areas where there is still worktobedone.Thebookisacomprehensivebut digestible summary of the field of conservation biogeographyandshould makeessentialreading, not only for the students at whom it is primarily aimed, but also for more experienced scientists. Theeditorsprofessattheoutsetthattheaimwas to achieve a degree of coherence among the chapters,anaimthatisachievedremarkablywell togiveaverycoherenttext. Thefirstsectionofthebookprovidesabrief but interesting history of the conservation move mentandthecontrastingvaluesheldbydifferent sectors of this movement (Chapters 2 and 3), as wellassomebackgroundtothefieldofconserva tion biogeography (Chapter 1). A distinction is madebetweenapproachesthatfocusonthecom positionofbiologicalcommunitiesandthosethat focus on ecosystem function through an under standingofecosystemprocessessuchasnutrient cycling(p.31).Aninterestingandgrowingfieldin ecology, which receives little attention in the book, uses the functional traits of species to ex plainthelinkbetweenthecompositionofbiologi cal communities and the function of the ecosys temsthatcontainthem.Functionaltraitssuchas bodymass,diet,habitataffinityanddevelopment mode of animals, and height and photosynthetic pathwayofplantscanhelpexplainhowspecies contribute to the processes underlying the func tioningofecosystemsandcanalsohelpinpredict ing how ecosystems will respond to environ mentalchange(McGilletal.2006). Thesecondsectionreviewsourcurrentun derstanding of the distribution of biodiversity, summarises the history of the global protected areas network and describes the methods avail able for more systematically representing biodi versityinfutureextensionstothisnetwork.There isastrongterrestrialfocushere,indeedthrough out the entirety of the book, which the authors acknowledge and which is owing to a less com pleteunderstandingofthedistributionofdiversity in the oceans and in freshwater habitats. It is worth noting, though, that the Census of Marine Life, an ambitious $650 million project that fin ished recently, has made huge progress towards understanding the biogeography of the oceans 93
newsandupdate (e.g.seeTittensoretal.2010).Evenintheterres trial realm, knowledge about the number and identity of the worlds species and how they are distributed remains very far from complete: the Linnaean and Wallacean shortfalls respectively (Chapter 4). A recent paper (Joppa et al. 2011) addressedbothoftheseknowledgegapssimulta neously by predicting the spatial distribution of undiscovered plant species, predicting that most new plant species will be discovered in areas al readyidentifiedashotspotsofplantdiversity,em phasising the importance of these areas for con servation. Chapter 5 provides an excellent sum mary of the many different types of protected areasintheglobalnetworkandthedifferentval uesthatunderpinthese,whileChapter6provides ausefulandsuccinctreviewoftheenormousand evergrowing literature on systematic conserva tionplanning. Thethirdsectionofthebookdescribeshow thetoolsofbiogeographycanbeusedtoplanfor environmentalchangeinconservation.Thisisthe only part of the book where the chapters appear somewhatdisjointed,butthisisprobablyowingto the attempt to summarise a vast literature in a very small number of chapters. Nevertheless, the chaptersinthissectionprovideexcellentdescrip tionsofsomeoftheavailablemethods,fromphe nomenological models that infer future changes fromcurrentpatterns(Chapter7)tomoreprocess based models that use the theory of island bio geographytopredicttheconsequencesforbiodi versityofshrinkingandincreasinglyisolatednatu ral habitat patches (Chapter 8). Chapter 9 deals with invasive species, which are an important driver of environmental change, and the homog enisation of biological communities, i.e. the ero sionofbetadiversity.Mostofthestudiesinvesti gating broadscale patterns of diversity have fo cusedoninventorydiversity,commonlymeasured as species richness, and it is only recently that studieshaveattemptedtomapbetadiversity(e.g. McKnight et al. 2007) and to relate it to spatial and environmental factors (e.g. Ferrier et al. 2007). Withagrowingneedtounderstandchanges in the natural environment and the impact of these changes on human society, the emerging fieldofconservationbiogeographyislikelytobe comeincreasinglyimportantinprovidingthenec essary theoretical basis and tools for doing so. This book provides an excellent foundation for that field and is highly recommended reading for students,scientistsandpractitionersofconserva tion.
TimNewbold
UnitedNationsEnvironmentProgrammeWorldCon servationMonitoringCentre,Cambridge,UK email:Tim.Newbold@unepwcmc.org; http://www.unepwcmc.org/timnewbold_368.html
References
Butchart, S.H.M., Walpole, M., Collen, B. et al. (2010). Globalbiodiversity:indicatorsofrecentdeclines. Science,328,11641168. Isbell, F., Calcagno, V., Hector, A. et al. (2011). High diversity is needed to maintain ecosystem ser vices.Nature,477,199202. Joppa,L.N.,Roberts,D.L.,Myers,N.etal.(2011).Biodi versityhotspotshousemostundiscoveredplant species.ProceedingsoftheNationalAcademyof Sciences of the United States of America 108, 1317113176. McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. (2006). Rebuilding community ecology from functionaltraits.TrendsinEcology&Evolution, 21,178185. McKnight, M.W., White, P.S., McDonald, R.I., Lam oreux, J.F., Sechrest, W., Ridgely, R.S. & Stuart, S.N. (2007). Putting betadiversity on the map: broadscale congruence and coincidence in the extremes.PLoSBiology,5,e272. Tittensor, D.P., Mora, C., Jetz, W., Lotze, H.K., Ricard, D.,VandenBerghe,E.&Worm,B.(2010).Global patterns and predictors of marine biodiversity acrosstaxa.Nature,466,10981101.
OneofthebenefitsopentoIBSmembersistheopportunitytohavejobopeningspostedonthe IBSblog(http://biogeography.blogspot.com/).Ifyouhaveapositionyouwouldliketohavead vertised, please contact Karen Faller (faller@wisc.edu) or Michael Dawson (mdawson@ucmerced.edu)withdetails.
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Anewencyclopediaforbiologicalinvasions
Encyclopediaofbiologicalinvasions,byDanielSimberloffandMarcelRejmnek(editors) 2011,UniversityofCaliforniaPress,792pp.ISBN:9780520264212 PriceUS$95(Hardbackorebook);http://www.ucpress.edu/
Despite existing in some form for many decades (Davis 2005), invasion ecology/biology is in many waysanascentandemergingfield,andisstillen gendering discussion regarding whether it indeed trulyexistsasafieldordisciplineinitsownright, or is rather a particularly focused aspect of com munityecologyorbiogeography(e.g.Marris2009, Pyek and Hulme 2009). As with many ecological disciplines, invasion ecology has seen fundamen taldisagreementsoveraspectsrangingfromcore definitions (including invasion itself; Falk Petersenetal.2006,RicciardiandCohen2007)to level of scientific objectivity (e.g. Larson 2007). Thefieldisatastageinitsdevelopmentwhere(1) dedicated journals exist (e.g. Biological Invasions) and there is a substantial number of academic articles published every year (for example a search of invasive species in Web of Knowledge returns 1181 articles published in 2010 alone), 2) thereisclearandsignificantinternationalinterest and action in relation to invasions and (3) an ex tendedpeercommunityisinvolvedinresearching andmanagingthethreatofinvasivespecies,from worldleading academics at researchintensive universitiestolocalgovernmentandconservation volunteers.Theresultoftheburgeoninginforma tionandunevenlevelsofunderstandingandfocus across the peer community is confusion and un certainty,rightfromthefundamentals(whatisan invasivespeciesexactly,andwhyisitinvasive?)to thespecifics(whatisthebesttechniqueforreduc ing populations of Crassula helmsii in my pond, andhowdoesthatdifferfrommanagingspreadin the local lake?). The time is ripe therefore for an encyclopaedia such as this one by Daniel Simber loff and Marcel Rejmnek to form a baseline for futuredefinitionsanddiscussions. The book is one of University of California Press Encyclopedias of the Natural World series, andaswiththeothervolumeshasawiderangeof entriesthatareeffectivelyshortessaysorsumma riesofkeytopicsrelating(inthiscase)tobiologi cal invasions, without citations but with relevant further reading at the end. The entries vary in length from 1 to 8 pages, and often incorporate usefulfiguresandoccasionallytables.Thebookis impressively glossy (all figures are in full colour) andwellpresented,whichisallthemoreremark able considering the relatively modest price. The editors, Daniel Simberloff and Marcel Rejmnek, areleadinginvasionecologistsandarewellquali fied to compile such a text; this is reflected not justinthebroadrangeofwellselectedtopicsthat thevolumeincludes(ofwhichthere are153)but also the rollcall of esteemed contributors that havesuppliedtheentries(ofwhichthereare197, manyofthemhighprofileinternationalresearch ers). The book is aimed not just at an academic audience, however, and the articles are written withtheinterestedandeducatedgeneralpublicin mind. Theindividualarticlescovervariousaspects of invasions,ranging from particular attributes of invasive species and invaded ecosystems to im pacts and management, interesting case studies andhistoricalperspectives.Clearlyitis notpossi bleto cover alloftheentriesinareviewsuchas this,butIdidfindseveralarticlesespeciallyinter esting, particularly because they highlight the many socioecological factors that complicate our relationshipswithpotentiallyproblematicspecies. TheentryonXenophobiaforexampledoesanex cellentjobofsummarisinghowsocietysrelation shipwithnonnativespeciesisconstructedincer tain ways by the use of loaded terms or cultural metaphors, for example the negative personifica tion of zebra mussels as outlaws on the west coast of the US, or the badging of harmful or distasteful species with appellations that note their foreign status (Japanese knotweed, Chinese mitten crab, English sparrow and so on). As a starting point for a discussion of scientific objec 95
newsandupdate tivity related to invasion biology it works excep tionallywell,andisexactlytherightsizefordiges tionbystudentsorinterestedamateurs. Indeed,oneofthebestusesIfindforrefer ence works such as these are as opening forays intotopicsforclassdiscussions,whetheratgradu ate or undergraduate level. Good examples in clude the entry on Succession, which very effec tivelyandconciselysummariseskeyconceptsthat take up whole chapters in many textbooks, and although invasion biology is only addressed to wards the end, it is clear how the two link to gether. Likewise, the discussion on Native invad ers, in which issues of invasive terminologies (and when they are appropriate) are covered, is excellently written and illuminating at a range of levels, particularly in relation to the many exam ples of invasion given. Certainly students and researchersnewtothesubjectwillhaveanyinitial confusion over what is meant by invasions dis pelledbythearticle,anditwillalsohelpthemto think objectively about whether a species really maybeconsideredinvasiveornot.Allofthearti clesIreadthroughwereofahighqualityandwell written/edited, with very little wasted space for suchalargevolume(althoughonoccasionfigures arenotalwaysrelevantImnotsurewhyanim age of Frank Buckland physicking a por poise(page2)isworthyofinclusionforexample, despitehisroleinfoundingthemainUKacclimati sationsociety). Ofcourse,itisalwayshardtogettheright balance between conciseness and detail in such entries, and to retain the relevant focus. The opening entry, Acclimatisation societies is a case inpoint:thearticledoesanexcellentjobofsum marisingthedevelopmentandimpactofsuchso cietiesindifferentcountries,manyofwhichwere responsible for the introduction of significant numbers of nonnative species around the globe before dying out in the face of increasing legisla tion, awareness of ecological risk from introduc tionsandlackofinterestfromthegeneralpublic. The article elegantly conveys how originally be nevolent intentions, such as the introduction of nonnatives to improve food resources, control pests and to soothe homesick colonists (among 96 otherreasons),inmostcasesfailedtoberealised and also (with some notable exceptions) that manysocietieswereunsuccessfulinactuallynatu ralising many species at all. But much is left un said: in some cases one is left wanting to know more about whether species referred to as released became naturalised, whether regions suchasSouthAmericamaintainedanysuchsocie ties (these countries are ignored, while others such as Germany and Italy receive only one sen tence)andultimatelywhethersuchsocietiesindi rectly provided evidence to force their own dis continuation.Asatastertowhettheappetite,the articlesucceedsverywell(andrelevantbookson the subject are provided in the Further Reading section),butitisnotanauthoritative,encyclopae dicsummaryinitself. As with any vast topic, covering all aspects inasinglevolumeisdifficultinthiscasethereis differential coverage of ecosystems (e.g. entries forcanals,lakes,riversandwetlands,butnocov erage of urban ecosystems, despite these being importantpointsofintroductionforsomeinvasive taxa);hypotheses(e.g.EnemyReleaseHypothesis, Novel Weapons Hypothesis, but no Tens Rule); geographical areas (Australia, the Great Lakes, Hawaiian islands, the Mediterranean, the Ponto Caspian, New Zealand and South Africa receive a particular focus) and species (good examples of somekeyspeciesorgroupssuchaszebramussel, earthworms and fishes, but understandably not comprehensive coverage). This is entirely reason able, and is not a criticism of the volume it is impossibletocoverthevastrangeoftopicsasso ciatedwithbiologicalinvasionsinsufficientdepth in a single volume, and the material that is in cludedisimpressive.Thedivisionofthebookbe tweeninvaderattributes,processes,taxa,ecosys tems,pathwaystoinvasionandsoonisverywell doneandrepresentsahugeeffortonthepartof theeditors,forwhichtheyshouldberoundlycon gratulated. I would encourage consideration of a second volume, however, at least with regard to key concepts and hypotheses. The opening guide to the Encyclopedia notes that there is a website with a list of articles, sample entries and so, and notesthatthesitewillevolvewiththeadditionof
ISSN19486596 new information, p. xxii). The web address has sincechangedandIwasunabletolocatethenew one.ThoughIhappilyagreethatthiscouldpoten tially be a very useful resource, given the rapidly changing environment of the internet, the publi cation of a second volume would perhaps be the mostreliableoption. In summary, this is an excellent reference work that combines readability with academic rigourthroughout.Itsbroadcoverageofthefield, high quality of production and reasonable price makes it an essential purchase for any university with departments teaching or researching within thebroadspectrumofecology,aswellasforindi vidualresearchersofspeciesinvasions.
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References
Davis, M.A. (2005) Invasion biology 19582004: the pursuitofscienceandconservation.In:Concep tual ecology and invasions biology: reciprocal approaches to nature (ed. by Cadotte, W.M, McMahon, S.M. and Fukami, T.) , pp. 3564. KluwerPublishers,London. FalkPetersen, J., Bhn, T. & Sandlund, O.T. (2006) On thenumerousconceptsininvasionbiology.Bio logicalInvasions,8,14091424. Larson, B.M.H. (2007) An alien approach to invasive species: objectivity and society in invasion biol ogy.BiologicalInvasions,9,947956. Marris,E.(2009)Theendoftheinvasion?Nature,459, 327328. Pysek,P.&Hulme,P.E.(2009)Invasionbiologyisadis ciplinethatstooyoungtodie.Nature,460,324 324. Ricciardi, A. & Cohen, J. (2007) The invasiveness of an introduced species does not predict its impact. BiologicalInvasions,9,309315.
RobertA.Francis
DepartmentofGeography,KingsCollegeLondon email:robert.francis@kcl.ac.uk;http://rg.kcl.ac.uk/ staffprofiles/staffprofile.php?pid=1961
bookreview
ApiscinehistoryoftheNeotropics
HistoricalbiogeographyofNeotropicalfreshwaterfishes,byJ.S.AlbertandR.R.Reis(editors) 2011,UniversityofCaliforniaPress,408pp.ISBN:9780520268685 Price59(Hardback);http://www.ucpress.edu/
The Neotropics leave an indelible impression on everyone who visits them. The seeds of some of themostimportantconceptsinecologyandevo lutionweresownduringtheSouthAmericantrav els of influential 19th century thinkers. For exam ple, the latitudinal gradient of diversity, now rec ognized as ecologys oldest pattern (Hawkins, 2001),wasfirstidentifiedbyvonHumboldt,while Batesdocumentedthevarietyandadaptationsof species in Amazonian forests, and Wallace and Darwinponderedthemechanismsresponsiblefor the myriad forms of life they encountered. Al though the Neotropics have played a crucial role in our understanding of the diversity of life on earth,inmanywaystheycontinuetorepresentan unexploredfrontier.Thisisparticularlyclearinthe case of Neotropical freshwater fish, a group esti mated to consist of more than 7000 species, and thataccountsforoverhalfthefreshwaterfishon the planet and around 10% of all vertebrate spe cies. JamesAlbertandRobertoReisgoalasedi torsoftheHistoricalBiogeographyofNeotropical Freshwater Fishes is to examine the evolutionary forces responsible for this diversity. In doing so theymakethecasethatmultipleprocessesofdi versification were involved and that these oper atedoverlongperiodsoftimeaswellasonacon tinental scale. The book itself is divided into two parts, the first of which examines current knowl edgeonthebiogeographyoftheregion,whilethe secondisaregionalanalysisthatlinkscontempo rarygeographicalpatternswithgeologicalhistory. The book is ambitious in scope and brings to getherpreviouslyfragmentedmaterialtoprovide anauthoritativeoverviewofthisimpressivegroup offish.AndwhileafisheyeviewoftheNeotropi cal ichthyofauna is inevitably drawn to the Ama 97
newsandupdate zon,thebookhasbroadcoverage,embracingthe AndesandextendingthroughCentralAmericaand intosouthernMexico.Asitmakesclear,itisnec essarytohaveacontinentalperspectivetounder standthediversityanddistributionofthisimpres sivegroup. I particularly liked the care and thought in volvedinputtingthebooktogether.Itisabeauti fully presented volume with informative tables and figures, many of them in colour. However, moreimportantthanthisisthattheeditorshave a strong sense of what the important issues are and how these should be best dealt with. Indeed the book is an essential reference for anyone wanting to learn more about the diversity or his toryofSouthAmericanfishes. One of the most challenging questions in ecology is explaining why different habitats sup portdifferentnumbersofspecies.Theextentofa habitat accounts for much of the variation but SouthAmericahasanexcessofspeciesrelativeto itsarea.Thecoreofthecontinent,particularlythe Amazon, is responsible for a disproportionate amountofthisdiversity.Itistemptingtoattribute thisexceptionalrichnesstotheuniquegeological andenvironmentalfeaturesoftheAmazon.How evermanyofthefishesthatinhabitthisriversys temareolderthantheAmazonBasinitself.More over,theAmazonianichthyofaunahasbeenaccu mulated gradually through tens of millions of years. The explanation, Albert, Petry and Reis ar gue, is rooted in the repeated subdivision and mergingofadjacentriverbasinsandtheirfaunas, withdispersallimitationandenvironmentalfilter ingplayingimportantroles.Theexceptionallyhigh diversity seems to be less to do with exceptional speciationratesthanwithlowratesofextinction. However, diversity is not just a measure of the numbers of species that cooccur but also of the typesofspeciesthatarefoundtogether.Auniver sal feature of natural assemblages is that some familiescontributeamuchhigherfractionofspe ciesthanothers.TheNeotropicsarenoexception. Ten families of fish account for 75% of the Neotropical icthyofauna. Characidae (including piranhasandtetras)andCichlidae(suchasdiscus) are particularly big hitters. One possibility is that this unevenness is simply the result of chance. Alternatively, historical and biological factors, ei therseparatelyortogether,couldcontribute.E.O. Wilson (2003) has argued that an ancient origin, combined with small body size, widespread geo graphic distribution and key innovations contrib utetothesuccessofsomegroupsrelativetooth ers. On the basis of the evidence presented by Neotropical fish, Albert, Bart and Reis conclude that these features are necessary but not suffi cient.Indeedtheynotethatcladescanbeancient (e.g. Arapaima, which is of Cretaceous origin), widespread (Arapaima again) or with small body size(e.g.Amazonsprattus)yetberepresentedbya handfulofspeciesatmost.Ontheotherhandsex ual and trophic innovation may play a role. Eco logical specialisation is also important. For exam ple,Cramptonnotesthatgroupsofcloselyrelated Gymnotiform electric fish species tend to be foundinanarrowrangeofhabitattypesbutmay bespreadacrosslargegeographicareas.Thefac torsthatunderpindiversificationarethesameas those that come into play in the explosive speci ationthatcharacterizestheAfricanriftlakes.The differencehereisthatthegameisplayedoutona continentalscaleasopposedtoalocalarena. Ofcourse,muchremainstobelearntabout the phylogenetic histories of Neotropical fishes andofthegeologicalcontextinwhichthesespe cies evolved. Nonetheless, as this book makes clear, the nature and timing of key events is be coming much better understood. The contribu tions to the book demonstrate how the growing body of molecular data, and its integration with ecological theory and earth sciences, has under pinned the recent and rapid progress in under standingthissystem.
Yourparticipationinfrontiersofbiogeographyisencouraged.Pleasesendusyourarticles,com mentsand/orreviews,aswellaspictures,drawingsand/orcartoons.Wearealsoopentosug gestionsoncontentand/orstructure. Pleasecheckhttp://www.biogeography.org/html/fb.htmlformoreinformation,orcontactusat ibs@mncn.csic.esandfrontiersofbiogeography@gmail.com.

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ISSN19486596 There have been many studies of tropical diversitybutuntilnowNeotropicalfishesfishhave received relatively little attention. This contrasts withSouthAmericanbirds,agroupthathasbeen prominentintestsofmacroecologicalhypotheses (e.g.Rahbeketal.,2007).Fishareresponsiblefor morediversityanddeservetobemorefullystud ied. This book provides the knowledge that will informtheseexcitingresearchopportunities.
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References
Hawkins,B.A.(2001).Ecologysoldestpattern.Trends inEcologyandEvolution16,470. Rahbek,C.,Gotelli,N.J.,Colwell,R.K.,Entsminger,G. L.,Rangel,T.F.L.V.B.andGraves,G.R.(2007). Predicting continentalscale patterns of bird species richness with spatially explicit models. Proceedings of the Royal Society B: Biological Sciences274,165174. Wilson, E.O. (2003). The origins of hyperdiversity. pp. 1318inPheidoleintheNewWorld:ADominant HyperdiverseAntGenus,Wilson,E.O.(ed).Har vardUniversityPress.
AnneE.Magurran
UniversityofStAndrews email:aem1@standrews.ac.uk; http://biology.standrews.ac.uk/magurran/
booksnotedwithinterest
GuidetostandardflorasoftheWorld: An annotated, geographically ar F.StuartChapinIII,PamelaA.Matson&Peter ranged systematic bibliography of the M.Vitousek principal floras, enumerations, check 2011,2ndedition,Springer,529pp. listsandchorologicalatlasesofdiffer 135(Hardback),44.99(Paperback) entareas ISBN:9781441995032/9781441995025 Principlesofterrestrialecosystemecol ogy
http://www.springer.com/
An outstanding textbook which, after definitions, setsthestagewithprimersonEarthsclimatesys tem and geological processes. What follows is a magisterial and comprehensive account of the movements of water, energy, carbon and nutri entsthoughnaturalsystems.Alongwithstandard generalisations, the authors delve into the finer detail and explain how biological processes can haveimportantmodulatingeffectsthroughspace andtime.Afinalreflectivepairofchaptersconsid ersglobalchangesandtheimplicationsforecosys tem management. The book is well written throughout and punctuated with excellent colour illustrations; noone from undergraduates to es tablished researchers can fail to learn something fromit.
DavidF.Frodin 2001,2ndedition,CambridgeUniversityPress, 1100pp. 198(Hardback),90(Paperback),US$120(e book) ISBN:9780521790772/9780521189774 http://www.cambridge.org/

Whilenotgenerallyourpolicytofeaturereprints, this standard text has newly appeared in paper back, bringing it within affordable reach of a greater number of researchers. It does exactly whatitsaysonthecover,makingitthedefinitive reference for anyone commencing work on the floraofanewregion.Despiteitsnotreceivingany further updates and its coverage ending in 1999, there remain no resources to rival it, either in print or online. It also contains insightful reviews onthehistoryoffloristicdescription.Anessential book which belongs in the library of every plant biogeographer. 99
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Field guide Afghanistan: Flora and Communityecology PeterJ.Morin vegetation

SiegmarW.Breckle&M.DaudRafiqpoor 2011,ScientiaBonnensis,Bonn,864pp. Price:Contactpublishers ISBN:9783940766304 http://www.scientiabonnensis.com/
Thefloraofthisvast,environmentallydiverseand biogeographically central country has yet to be fully catalogued, but this field guide represents a landmark accomplishment on the path to doing so,fillingananomalousgapatthejunctionofsev eralfloristicrealms.Itcontainsapictorialguideto over1200species(>25%oftheflora)plusgeneral chapters onvegetative formations and should fa cilitate both local and international study. Copies have been freely distributed to universities and institutes throughout Afghanistan as well as her baria and museums worldwide. A feature on this projectisplannedforafutureeditionofFrontiers ofBiogeography.
2011,2ndedition,WileyBlackwell,407pp. 90(Hardback),34.99(Paperback) ISBN9781444338218/9781405124119 http://www.wiley.com/

Community ecology straddles conventional inter actionbased ecology and biogeography; recent heateddebateinthepagesofAmericanNaturalist hasevendisputedwhethercommunitiestrulyex ist as natural entities. Unsurprisingly the author makes a strong case for communities, stressing patterns and processes that can only be under stood at this level, and pleasingly devotes equal attention to both models and experimental data. The textbook is intended for a graduate course and represents a major update on the previous edition.Onemightquerythebalanceofcoverage ofvarioustopicsbutneverthelessthisremainsthe only textbook exclusively devoted to this scale of study.

Editorialpolicyforbookreviews
MarkusEichhorn
BookReviewEditor. email:Markus.Eichhorn@nottingham.ac.uk
FrontiersofBiogeographywillpublishindepthreviewsofrecentlypublishedbooks(typicallylessthanone yearold)onbiogeographyorofinteresttobiogeographers,alongsideaNotedwithInterestsectionprovi dingbriefdetailsofnewpublications.Authors,editorsorthirdpartiesareinvitedtosuggestbooksforre viewtotheBookReviewEditor,DrMarkusEichhorn,SchoolofBiology,UniversityPark,NottinghamNG7 2RD, United Kingdom; telephone ++44 (0)115 951 3214; email markus.eichhorn@nottingham.ac.uk. We welcomeofferstoreviewbooksforFrontiersofBiogeography,butwillnotacceptanoffertoreviewaspeci ficbook.Anyonewishingtoreviewbooksshouldsendabriefcurriculumvitae,descriptionofcompetencies, andastatementofreviewingintereststotheBookReviewEditor.Reviewsshouldbeinanessaystyle,ex pressinganopinionaboutthevalueofthebook,itsfocusandbreadth,settingitinthecontextofrecent developmentswithinthefieldofstudy.Textbookreviewsshouldconsidertheirutilityasresourcesfortea chingandlearning.Avoiddescribingthebookchapterbychapterorlistingtypographicalerrors.Thelength shouldnormallybe1000words(1500wordsforjointreviewsofrelatedtexts)includingamaximum10refe rences.Authorsmaysuggestashortheadingforthereview,followedbythetitleofthebook(s),theaut hors/editors,publisher,publicationdate,price,hbk/pbk,pages,ISBNandwebsite(whereavailable).Figures ortableswillnotordinarilybeincluded.Authorsofreviewsmustverifythattheyhavenotoffered(andwill notoffer)areviewofthesamebooktoanotherjournal,andmustdeclareanypotentialconflictofinterest thatmightinterferewiththeirobjectivity.Thismayformabasisforeditorialdecisionsandsuchdisclosures maybepublished.Bookreviewswillusuallygothroughalighteditorialreview,thoughinsomecircumstan cesalsowillbeconsideredbyoneormorereferees.
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thesisabstract
Applyingspeciesdistributionmodelingfortheconservationof Iberianprotectedinvertebrates
RosaMaraChefaoui
PhDThesis,DepartamentodeBiodiversidadyBiologaEvolutiva,MuseoNacionaldeCienciasNaturales, c/JosGutirrezAbascal2,28006Madrid,Spain. email:rosa.chef@gmail.com;http://www.biogeografia.org/ Abstract. This article outlines the approaches to modeling the distribution of threatened invertebrates usingdatafromatlases,museumsanddatabases.SpeciesDistributionModels(SDMs)areusefulforesti matingspeciesranges,identifyingsuitablehabitats,andidentifyingtheprimaryfactorsaffectingspecies distributions. The study tackles the strategies used to obtain SDMs without reliable absence data while exploring their applications for conservation. I examine the conservation status of Copris species and Graellsia isabelae by delimiting their populations and exploring the effectiveness of protected areas. I showthatthemethodofpseudoabsenceselectionstronglydeterminesthemodelobtained,generating differentmodelpredictionsalongthegradientbetweenpotentialandrealizeddistributions.Afterassess ing the effects of species traits and data characteristics on accuracy, I found that species are modeled moreaccuratelywhensamplesizesarelarger,nomatterthetechniqueused. Keywords: Environmental niche modeling, Iberian Peninsula, invertebrates, predictive accuracy, species distributionmodels The rapid disappearance of habitats and species starklycontraststheneedtoconservebiodiversity against our inability to inventory and protect all speciesindividually.Knowledgeaboutbiodiversity remainsinsufficientbecausemanyspeciesarestill not described (the "LinneanShortfall";Brown andLomolino 1998) and the distributions of de scribed species often are inadequately defined (the "Wallacean Shortfall"; Lomolino 2004). It is thereforeessential to identify threatened species and describe their distributions using approaches thatovercomethetimeandbudgetconstraintsof systematicconservationplanning. Arajoetal.(2007)demonstratedtheneed for additional protected areas for the effective conservation of the diversity of plants and verte brates in the Iberian Peninsula. Preliminary data suggestthattheexistingnetworkofreservesalso would be ineffective in representing invertebrate species (Verd and Galante 2009).Unfortunately, the conservation of invertebrates faces serious challengesduetotheirhighdiversity,complexlife cyclesanddifficulttaxonomy,amongotherfactors (seeNew1998). Geographic Information Systems (GIS) sig nificantly advanced the conservation of endan gered species because they allow us to delimit species potential distributions (e.g.Hortaletal. 2005), to control their populations (e.g.Daviesetal. 2005), to analyze their niche (Peterson et al. 2002), design networks of pro tectedareas (e.g.PearceandBoyce2006),and to forecastthefuture(e.g.Hilletal.2002).Together, the databases taken from atlases, museums and herbaria have emerged as a valuable source of species occurrence records (e.g. Elith and Leath wick 2007). Unfortunately, these data from het erogeneous sources may containerrors or havebeen obtained using a biasedsampling pro cedure (Hortal et al. 2007, 2008, Newbold 2010).Besides, they do not usually provide reli able absences needed to perform consistent pre dictive models (Anderson et al. 2003, Lobo et al. 2007), so alternatives have been sought generat ing models based only on presences (Hirzel et al. 2002, Pearce and Boyce 2006), sometimes em ploying pseudoabsences obtained in different ways (Zaniewski et al. 2002, Engler et al. 2004, 101
SDMappliedtoinvertebrateconservation Loboetal.2006,2010). Formydoctoralthesis,Ievaluatedtheutil ity of SDMs for the conservation of threatened invertebrates in the Iberian Peninsula (Chefaoui 2010).The majority of thespecies studied herehave been designated by the European Un ion as species of community interest requiring protectionandconservation(HabitatsDirective).I used presenceonly dataon Iberian threatened invertebrates obtained frommuseums, atlases and databases. I applied presenceonly methods such as ENFA (Ecological Niche Factor Analysis) and MDE (MultiDimensional Niche Envelope), in additiontoothermethodsthatrequirepresences and absences (here, pseudoabsences): GAM (Generalized Additive Models), GLM (Generalized LinearModels)andNNET(NeuralNetworksMod els).Iapproachedmethodologicalissuesconcern ing the difficulties associated with predicting the distributionofspecieswhenreliableabsencedata arenotavailable,andexploredthepossibilitiesof SDMs as a tool for conservation of endangered and threatened Iberian invertebrates. In this res pect, I explored the applications of SDM to esti matespeciesranges,identifysuitablehabitatsand the primary factors affecting species distribution in order to assess the conservation status of threatenedinvertebrates. Dung beetle populations, which are in de cline in the Iberian Peninsula, play a critical eco logical role in extensive pasture ecosystems by recyclingorganicmatter.Wedelimitedthepoten tial distribution of the two species of Copris (Coleoptera, Scarabaeidae) that inhabit the Ibe rian Peninsula using ENFA (Chefaoui et al. 2005). ENFA is a presenceonly method that compares the environmental values of the localities where thespecieshasbeenobservedwithrespecttothe environmental values of the territory studied (Hirzel et al. 2002). We explored the environ mental niche occupied by each species in a small region,theCommunityofMadrid(CM),torestrict the role of dispersal constraints discriminating possible areas of cooccurrence and identifying thespecificenvironmentalcharacteristicsofeach species.Weidentifiedthatsolarradiationandthe presence of calcareous soils are critical to the 102 presence of Copris hispanus, while Copris lunaris requires siliceous soils and high rainfall.Both Co prisspeciesaredistributedalongageographicand environmental gradient from the Tajo basin (warmer,dryer,withstrongannualweathervaria tions) where only C. hispanus is found, towards the mountain slopes of the Sistema Central (colder,higherrainfall)whereC.lunarispredomi nates. The environmental niches of both species aredistributedalongaDryMediterraneantoWet Alpine axis, and overlap in areas of moderate temperatures and precipitations in the north of CM. Wealsostudiedthedegreeofprotectionof keypopulationsofC.hispanusandC.lunaris,mak ing a proposal to improve their conservation. To evaluatetheconservationstatusofCoprisspecies, wetookintoaccountthesizeofprotectedsitesas well as the values of habitat suitability in each protected natural site and Natura 2000 network. We found that Copris species were poorly con served in the previous protected sites network: forC.hispanusonlytwoprotectedsitesmeasured around 30 km2, and for C. lunaris a single area measured183km2.However,protectionprovided bySitesofCommunityImportance(SCIs)seemsto improve the general conservation status of these species in CM because the area and connectivity of protected sites have been increased substan tially. Chefaoui and Lobo (2008) assessed the ef fects of pseudoabsences on model performance whenreliableabsencedataarenotavailable.We compared seven procedures to generate pseudo absencedatatobeusedinGLMlogisticregressed models. These pseudoabsences were selected randomly or by means of presenceonly methods (ENFA and MDE) to model the distribution of a threatened endemic Iberian moth species (Graellsiaisabelae).Ourpurposewastoshowthe possibilityofachievingdifferentforecasteddistri butionsdependingonthemethodandthethresh oldusedtoselectthesepseudoabsences. The results showed that the pseudo absence selection method greatly influenced the percentage of explained variability, the scores of theaccuracymeasuresand,mostimportantly,the
RosaM.Chefaoui predicted range size. As we extracted pseudo absences from environmental regions further from the optimum established by presence data, the models obtained better accuracy scores, and overprediction increased. Conversely, the profile techniquesthatgeneratedwiderunsuitableareas, produced functions with lower percentages of explained deviance and poorer accuracy scores, but more restricted predictive distribution maps, similar to the observed distribution. The random selectionofpseudoabsencesgeneratedthemost constrainedpredictivedistributionmap. Based on results of the aforementioned work, we identified the environmental variables most relevant for explaining the distribution of Graellsia isabelae and assessed this species con servation status (Chefaoui and Lobo 2007). We modeledthepotentialdistributionoftheinsectby performing GLM with pseudoabsence data se lected from an ENFA model. We found that the best predictor variables were summer precipita tion(rangingfrom1250mmto3250mm),aridity, andmeanelevation.This speciesprefershabitats withmidrangemountainconditions.Withrespect to host plants, the presence of G. isabelae was associated mainly with Pinus sylvestris and P. ni gra. Moreover, we found 8 areas exclusively in theeasternIberianterritory,andalargerunoccu piedhabitatinthewesternIberianPeninsula,indi cating that this species is probably not in equili brium with its environment because of historical factors (Chefaoui and Lobo 2007). We sug gestedthatthecurrentdistributionofthespecies was associatedwith the dynamismof itshost plantsduring glacial periodsof the Holocene, whenthe forests of Pinussylvestrisdecreased strongly inthe northwestern part ofthe penin sula.Afteranalyzingthepossibilityofconnectivity andfragmentationoftheeightpopulationsdelim itedaswellasthedegreeofprotectionofG.isa belae on the SCIs, we found that the SCIs under protectiondidnotseemsufficienttomaintaincu rrent populations. Moreover, our study rejected theideathatthespecieswasexpandingitsrange duetoreforestation.Becausetheconservationof G.isabelaedependsontheforestsofPinussylves tris and P. nigra located both inside and near to SCIs, we suggestedthat the reintroduction of the speciesinthesehabitatscouldimproveitsconser vation. Tounderstandthelimitationsandpossibili ties of SDM techniques, we evaluated the effects of species traits and data characteristics on the accuracy of SDMs for redlisted invertebrates (Chefaoui et al. 2011). We applied three SDM techniques (GAM, GLM and NNET) using pseudo absences to model the distribution of 20 threa tened Iberian invertebrates. We correlated the accuracyoftheobtainedmodelswithseveraldata characteristics and species ecological traits. We examinedtwodatacharacteristics,theamountof data (N) and the relative occurrence area (ROA), andbothsignificantlyaffectedtheaccuracyofthe models.GreaterAUCvaluesandhighersensitivity scores were obtained from samples for which there were more than 200 records. In general, specieswhosedistributionsweremostaccurately modelledwerethosewithagreatersamplesizeor smaller ROA. In addition, species related to habi tatsthatareproblematictodetectusingGISdata, such as riparian or humid areas, seemed to be moredifficulttopredict.
Summary
TheperformanceofSDMsdependsonthetypeof data and the characteristicsof the species. Pres enceonly methods (ENFA and MDE) achieved worse validation results and overpredicted more than techniques using pseudoabsences. Never theless,presenceonlymethodscanbeveryuseful for obtaining pseudoabsences and discovering the environmental response of species. The method of pseudoabsence selection strongly de terminedthepredictedrangesize,generatingdif ferent model predictions along the gradient be tween potential and realized distributions. There isanaddeddifficultyinobtainingpredictionsthat closely approximate the realized distribution of species under nonequilibrium conditions, be cause both presence and absence data may be possible under similar environmental conditions. Irrespective of the approach used, species distri butionsaremodelledmoreaccuratelywhensam 103
SDMappliedtoinvertebrateconservation ple sizes are larger. Species in habitats that are difficulttodetectusingGISdata,suchasriparian species, thus may tend to be more difficult than mosttopredict.
status of the silverspotted skipper butterfly (Hesperia comma) in Britain: a metapopulation successstory.BiologicalConservation,124,189 198. Elith,J.&Leathwick,J.R.(2007)Predictingspeciesdis tributionsfrommuseumandherbariumrecords using multiresponse models fitted with multi variate adaptive regression splines. Diversity andDistributions,13,165175. Engler, R., Guisan, A. & Rechsteiner, L. (2004) An im proved approach for predicting the distribution ofrareandendangeredspeciesfromoccurrence and pseudoabsence data. Journal of Applied Ecology,41,263274. Hill,J.K.,Thomas,C.D.,Fox,R.,Telfer,M.G.,Willis,S.G., Asher,J.&Huntley,B.(2002)Responsesofbut terflies to twentieth century climate warming: implications for future ranges. Proceedings of theRoyalSociety,269,21632171. Hirzel, A., Hausser, J., Chessel, D. & Perrin, N. (2002) EcologicalNiche Factor Analysis: How to com pute habitatsuitability maps without absence data?Ecology,83,20272036. Hortal, J., Borges, P.A.V., Dinis, F., et al. (2005) Using ATLANTIS Tierra 2.0 and GIS environmental information to predict the spatial distribution andhabitatsuitabilityofendemicspecies.Direc oRegionaldeAmbienteandUniversidadedos Aores, Horta, Angra do Herosmo and Ponta Delgada,Horta,Faial. Hortal, J., Lobo, J.M., & JimnezValverde, A. (2007) Limitationsofbiodiversitydatabases:casestudy on seedplant diversity in Tenerife (Canary Is lands).ConservationBiology,21,853863. Hortal,J.,JimnezValverde,A.,Gmez,J.F.,Lobo,J.M., & Baselga, A. (2008) Historical bias in biodiver sity inventories affects the observed realized nicheofthespecies.Oikos,117,847858. Lobo, J.M., Verd, J.R. & Numa, C. (2006) Environ mental and geographical factors affecting the Iberian distribution of flightless Jekelius species (Coleoptera: Geotrupidae). Diversity and Distri butions,12,179188. Lobo,J.M.,Baselga,A.,Hortal,J.,JimnezValverde,A., & Gmez, J.F. (2007) How does the knowledge about the spatial distribution of Iberian dung beetle species accumulate over time? Diversity andDistributions,13,772780. Lobo, J.M., JimnezValverde, A., & Hortal, J. (2010) The uncertain nature of absences and their im portance in species distribution modelling. Eco graphy,33,103114. Lomolino, M.V. (2004) Conservation biogeography. FrontiersofBiogeography:newdirectionsinthe geographyofnature(ed.byM.V.Lomolinoand L. R. Heaney), pp. 293296. Sinauer Associates, Sunderland,Massachusetts.
Availabilityofthesis
Printed and PDF copies are available in the Sci ence Faculty Library, Universidad Autnoma de Madrid (http://biblioteca.uam.es/ciencias/). A PDFcopyisalsoavailableatrequestfromtheau thor.
Acknowledgements
Iwouldliketothankmytwosupervisors,JorgeM. LoboandJoaqunHortalfortheirsupportanden couragement.
References
Anderson,R.P.,Lew,D.&Peterson,A.T.(2003)Evaluat ing predictive models of species distributions: criteria for selecting optimal models. Ecological Modelling,162,211232. Arajo,M.B.,Lobo,J.M.&Moreno,J.C.(2007)Theef fectiveness of Iberian protected areas in con serving terrestrial biodiversity. Conservation Biology,21,14231432. Brown,J.H.&Lomolino,M.V.(1998)Biogeography,2nd edn.SinauerPress,Sunderland,Massachusetts. Chefaoui,R.M.,Hortal,J.,&Lobo,J.M.(2005)Potential distribution modelling, niche characterization and conservation status assessment using GIS tools:acasestudyofIberianCoprisspecies.Bio logicalConservation,122,327338. Chefaoui,R.M. & Lobo, J.M.(2007) Assessingthe con servation status of an Iberian moth using pseudoabsences. The Journal of Wildlife Man agement,8,25072516. Chefaoui, R.M. & Lobo, J.M. (2008) Assessing the ef fectsofpseudoabsencesonpredictivedistribu tion model performance. Ecological Modelling, 210,478486. Chefaoui,R.M.(2010)Modelospredictivosaplicadosa la conservacin de invertebrados protegidos iberobaleares. Ph.D. Thesis. Universidad Aut noma de Madrid, Departamento de Biologa, FacultaddeCiencias,196pp. Chefaoui,R.M.,Lobo,J.M.&HortalJ.(2011)Effectsof speciestraitsanddatacharacteristicsondistri butionmodelsofthreatenedinvertebrates.Ani malBiodiversityandConservation,34,(inpress). Davies, Z.G., Wilson, R.J., Brereton, T.M. & Thomas, C.D. (2005) The reexpansion and improving
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New,T.R.(1998)Invertebratesurveysforconservation. OxfordUniversityPress,NewYork. Newbold,T.(2010)Applicationsandlimitationsofmu seum data for conservation and ecology, with particularattentiontospeciesdistributionmod els.ProgressinPhysicalGeography,34,322. Pearce, J. & Boyce, M.S. (2006) Modelling distribution andabundancewithpresenceonlydata.Journal ofAppliedEcology,43,405412. Peterson,A.T.,Ball,L.G.&Cohoon,K.P.(2002)Predict ingdistributionsofMexicanbirdsusingecologi calnichemodellingmethods.Ibis,144,E27E32. Verd,J.R.&Galante,E.,eds.(2009)AtlasdelosInver tebrados Amenazados de Espaa (Especies en peligro crtico y en peligro). Direccin General para la Biodiversidad, Ministerio de Medio Am biente,Madrid,340pp. Zaniewski, A.E., Lehmann, A. & Overton, J.M. (2002) Predicting species spatial distributions using presenceonlydata:acasestudyofnativeNew Zealand ferns. Ecological Modelling, 157, 261 280.
EditedbyRichardPearson
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opinion and perspectives

opinion
ISSN19486596
Politicalerosiondismantlestheconservationnetworkexisting intheCanaryIslands
JosMaraFernndezPalaciosandLeadeNascimento
Island Ecology and Biogeography Group, Instituto Universitario de Enfermedades Tropicales y Salud PblicadeCanarias(IUETSPC),UniversidaddeLaLaguna(ULL),Avda.AstrofsicoFranciscoSnchezs/n, 38206,LaLaguna,(Tenerife),Spain email:jmferpal@ull.es;http://webpages.ull.es/users/jmferpal Abstract. The outstanding nature of the Canary Islands has been recognized by European, national and regionaladministrationssincethearrivalofdemocracyinSpain.Fortyfivepercentofitsemergedterri toryhasbeendeclaredasNaturalProtectedAreas,fourCanarianNationalParkswereincludedwithinthe Spanish network, more than 200 endemics were listed in the Spanish catalogue of endangered species, and450specieswerelistedintheCanariancatalogueofprotectedspecies.However,inrecentyears,po liticaldecisionshavestarteddismantlingthissplendidconservationnetwork,whichimpedesconstruction oflargeinfrastructure,golfcoursesandresorts,despitetheadviceofthescientificcommunity.Canarian natureisnowfacingtwothreats:delistinganddowngradingofnumerousendangeredspecies,andtrans ferofthemanagementofCanarianNationalParkstotheregionaladministration. Keywords:Biodiversityloss,endangeredspecies,NationalParks,naturalprotectedareas,politicalcorrup tion,scientificcommunity,speciesdelisting Recently the CanarianParliament hasapproveda new version of the Canarian catalogue of pro tected species (see Box 1) that reduces substan tially both the number of species included (from 466 species in the 2001 list to 361 species in the 2010 list) and the protection afforded (from 381 threatenedspeciesto160,andfrom85protected speciesto18).Thesereductionshavebeenwidely criticized by environmental NGOs and the local scientific community1, mainly due to the absence ofarigorousscientificprocessinitsdevelopment. Although certainly the first version of the cata logue could be improved, the main reasons be hind the new revisions were not conservation is suesbutratherstrictly political. Thereasonsmay include, for instance, the development of large infrastructures, such as industrial harbours and golfcourses,whichuntiltherevisionswereforbid denduetotheirimpactsonprotectedspeciesin cludedintheoriginalversionoftheCanariancata logue. Changesintheenvironmentallegislationof the Canary Islands entail a serious threat to the nature of this region of biogeographical interest (FranciscoOrtega et al., 2000; Juan et al., 2000; FernndezPalacios & Whittaker, 2008). Thus, we believe it is important to share our appraisal of thecurrentsituationwiththeinternationalscien tificcommunity. Within the new revised catalogue a com pletely new criterion for protection has emerged especies de inters para los ecosistemas canar ios(literally:speciesofinterestforCanarianeco systems),comprising152species(seeBox1).The phrase is poorly chosen. It is supposed to apply onlytoendangeredspecies,consequentlythefre quent and abundant species which usually struc ture and dominate the ecosystems are explicitly not listed, leading to a curious paradox: the Ca narian pine (Pinus canariensis) is not a species of interest for the Canarian pine forest, the Macaronesian Laurel (Laurus novocanariensis) is
1. Seedifferentreactionsathttp://www.nodescatalogacion.com,http://www.wwf.es,http://www.greenpeace.org, http://www.atan.org, http://www.ecologistasenaccion.org, http://especiesamenazadascanarias.blogspot.com, http://ecooceanos.blogspot.com,http://www.seo.org,.
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JosMaraFernndezPalaciosandLeadeNascimento
Box1
Law 4/2010, June 4, of the Canarian Catalogue of Protected Species (see the original Spanish text at http://www.gobiernodecanarias.org/boc/2010/112/) Article3.Canarianprotectedspecies 2)SpeciesofinterestforCanarianecosystems The Canarian Catalogue of Protected Species will also include species of interest for Canarian ecosys tems"whicharethosethat,withoutbeinglistedinthethreateningsituationsabove(endangeredorvul nerable),areworthyofparticularattentionforitsecologicalsignificanceinareasoftheCanarianNetwork ofNaturalProtectedAreasorNatura2000network. 2.EffectsofinclusionintheCatalogue b)ThelegalregimeforprotectionofspeciesofinterestforCanarianecosystems"willbeapplicableonly intheterritoryoftheCanarianNetworkofNaturalProtectedAreasorNatura2000Network.Tothisend, applicablemeasuresshallbeprovidedbythemanagementplansofNaturalProtectedAreasandHabitats oftheNatura2000Networkinwhichtheyarelocated.Suchplansshallincludethedeterminations,con trolandmonitoringtoensureeffectivenessofprotection,orwhereapplicable,thejustificationthatthere isnoneedforplans.(...)Inthecaseofactionspromotedbyreasonsofpublicinterestandpriorityaffect ingthespeciesofinterestforCanarianecosystems"theseactionscouldbepossibleaslongastheydo notaffecttheecosystemsubstantially,underthetermsinparagraphs4to7oftheArticle45oftheLaw 42/2007,December13,ofNaturalHeritageandBiodiversity. notaspeciesofconcernfortheLaurelforest,and so on. This is not to say that the most common structuring species of the Canarian ecosystems have to be included in the catalogue, but we wouldliketodrawattentiontotheinadequacyof theconcept. But this conceptual shortcoming pales in comparisonwiththerealrepercussionofthenew criterion, which is that those species listed here are only protected if present in an already desig nated Natural Protected Area (NPA). (In the Ca naries,thatmeansineithertheCanarianNetwork ofNPAsortheEuropeanUnionNatura2000Net work, which overlap extensively). If a listed spe cies, for instance the woodcock (Scolopax rusti cola) or the coot (Fulica atra) which are both in cludedunderthenewcriterion,dwellswithinthe limits of the protected area they are safe; but if any birds cross those limits (which are not that obvioustobirds,unfamiliarastheyarewithGIS), theycanbeshotlegallybyhunters.Thesamein consistency affects, for instance, ca. 10 endemic species of sea lavenders (Limonium spp.) pro tectedincertainravines,butnotinothers. The new law could have negative implica tionsforconservationbiogeography,andthiscan beillustratedwithsomeexamplesoftheCanarian flora and fauna. The endemic legume Cicer ca nariensis, previously considered as vulnerable in the2001Canariancatalogue,isnowincludedun der the criterion species of interest. From its 12 locations (ten in La Palma and two in Tenerife), the six populations in the North of La Palma2 are outside NPAs and therefore unprotected accord ing to the new law. Metapopulation dynamics in thisspeciescouldbeaffectedbythisnewcriterion if source populations within these northern loca tions are threatened, endangering sink popula tions included in NPAs. The same could apply to theAbaloneorCanarianclam(Haliotistuberculata ssp.coccinea)ortheSeaHorse(Hippocampuship pocampus). Both are marine species with sparse populations in the meso and infralittoral, which donotalwayscoincidewiththegeographicalloca tionofthemarineSpecialAreasforConservation, whichoccupymainlyleewardfringesontheArchi pelagos coasts. Collection and capture of both speciesisprohibitedbytheRegulationoftheFish
2. AccordingtotheevaluationofthisspeciesbytheCanarianGovernment(ServiciodeBiodiversidad2009),there aresixpopulationnucleiintheNorthofLaPalma,distributedinthreelocationsmorethan10kmdistantonefrom eachother. frontiersofbiogeography3.3,20112011theauthors;journalcompilation2011TheInternationalBiogeographySociety
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Canarianconservationnetworkdismantled eriesLawoftheCanaryIslands,buttheirinclusion inthenewcriterionmayleadtoconfusiononthe fishing ban in populations outside of the reserve networks. The case of the sea grass Cymodocea nodosa is of particular interest for two reasons; thisspeciesstructuresacommunity(sebadales), considered as Natural Habitat of Community In terest by the Habitats Directive, and its presence inthelittoralzoneisoneofthemainobstaclesto theconstructionorenlargementofharbours.The most recent is the Puerto de Granadilla, where conservation of a European priority ecosystem comes into conflict with European funding of a largeinfrastructure.Thesebadalesareakeycom munity from an ecological point of view as they play an important role in the carbon cycle, stabi lize sandy soils, export biomass and act as a fish nurseryarea(Barberetal.2005).Thelatterchar acteristicisalsoveryimportantforthesustainabil ityoflocalfisheries.Also,themarinemeadowsof C. nodosa in the Canary Islands and Mauritania are the most extensive examples at the species southernlimitandcompromisingthemmaythere foreleadtorangecontraction.Theconstructionof Puertode Granadillawill severelydamageoneof themostgeneticallydiversepatchesofsebadales in the Archipelago (Alberto et al. 2008). In 2009, asaprecautionarymeasure,theSuperiorCourtof Justice of the Canary Islands suspended the pro posalsubmittedbytheCanarianGovernment,the Port Authority and the Canarian Company of Gas Transportation,todelistC.nodosa3.Currently,the European Courts have declared admissible the complaintfiledby the NGO EcologistasenAccin asking for the public release of documents that included alternatives to the construction of the harbour (including a renewal of the infrastruc turesofalreadyexistingharbours),thatwerehid den from the European Commission by Spains NationalGovernment. This controversial criterion especies de inters para los ecosistemas canaries is an ad aptation of the criterion species susceptible to habitatdisturbance,fromthepreviouscatalogue. Infact,manyofthespeciesofinterestcomefrom theformerlistofsusceptiblespeciesoraredown gradedthreatenedspecies.Howeverintheformer criteriontherewerenorestrictionsintheprotec tion,suchasthelocationornotinaNPA,andthe main consideration to include a species was that its habitat was threatened, in regression, frag mentedorlimited.Thepreviouscriterionforpro tection was much more appropriate if we think about the design of the Canarian Network of NPAs. Unfortunately the Canarian Network was notbasedonathoroughanalysisofmetapopula tion dynamics, genetic diversity or viability of populations, but simply in protecting less de graded remnants of communities that were still available.Asinmanypartsoftheworld,reserves were not designed to meet the principles of sys tematic conservation planning needed to achieve representativenessandpersistenceofbiodiversity (Margules and Pressey 2000). The situation fur ther worsens in the Canaries when data, trends andviabilityofpopulationsarealmostunknown. The Canarian Network is largely protecting speciesfrommarginalpopulations.Moreover,the protection of species present only in the current Reserve Network inhibits reestablishment of originaldistributions.Agoodexampleisthelaurel forestinAnagaRuralPark,whichnowadaysisthe bestrepresentationofthisforesttypeinTenerife yetstillanimpoverishedfractionofitspastdistri bution throughout the windward slope of the is land.Fromthepointofviewofmitigatingtheef fectsofglobalchange,vulnerabilityofcertainspe ciesoutsidetheNetworkwouldhinderaltitudinal migration,especiallywhenecologicalcorridorsare notincludedinthedesignofNPAs. ThepracticeofprotectingtaxaonlyinNPAs is already working in Catalonia (the only prece dent in Spain). The Catalonian Plan of Areas of Natural Interest includes species of flora and fauna strictly protected in designated areas. To our knowledge no cases of the failure of these practices or public disapproval have been re portedthere,butwesuspectthatthespecieswith restricted protection in the Catalonian NPA Net
3.Seenewsinhttp://www.laprovincia.es. 4.Seehttp://www.laopinion.es,http://www.ecologistasenaccin.org.
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JosMaraFernndezPalaciosandLeadeNascimento work were not demoted from higher protection. In theory, the main aim of the existence of re gional catalogues is ensuring the protection of particular species that are not considered by the National Catalogue. On the other hand several authors have questioned and analysed the effec tivenessofNPAsNetworksinbiodiversityconser vation (Jaffre et al. 1998, Rodrigues et al. 2004) and concluded that reserve networks are geo graphically and taxonomically unbalanced leaving a big proportion of endemic and threatened spe ciesunprotected. This way of thinking may function well when protecting a resource, for instance marine sanctuaries are intended to increase catch in neighbouring areas outside, and this works com petently in the Canaries Marine Reserves with Fishery Interest, but is nonsensical when the aim ofthedeclarationistoprotectathreatenedspe cies.IfaspeciesisprotectedwhenwithinaNPA, but unprotected when beyond the area, what is reallyachievedintermsofprotection?Mightitbe too cynical to suggest the greatest achievement wouldbethepoliticalgoalofinflatingthenumber ofspeciesincludedinthecataloguethusreducing thenumberofcriticsofdelisting?Despitenumer ouspublicprotestsandtheclearoppositionofthe majorityoftheCanarianscientificcommunity,the new catalogue was presented by the leading po litical force in the Regional Parliament. These kinds of conflicts are not exclusive to the Canary Islandsandarenowadaystakingplaceindifferent regions of the world (Possingham et al. 2010, Metzgeretal.2011). Ifthedelistingitselfisnotofsufficientcon cern,othernewsmakestheoutlookevenbleaker. The Canaries harbour four of the 13 National Parks (NPs) in Spain Caadas del Teide (Tenerife), Caldera de Taburiente (La Palma), Ti manfaya (Lanzarote) and Garajonay (La Gomera) despiterepresentingonly1.5%ofthecountrys geographical area. After decentralization of the Spanish State with the arrival of the democracy, the NPs were simultaneously comanaged by the Central Government (Madrid) and the Regional Governments. However, the Spanish Constitu tional Court now has determined that NPs man agement is exclusively a matter for the Regional Governments. Consequently the Central Govern ment has transferred all management to the re gions.InthecaseoftheCanarianarchipelago,this management was intended to be subsequently delegated to the respective island Councils (Cabildos) in 2012, although recently the new deputy of Environment of the Canarian Govern mentexpressedherintentiontodiscussagainthis transfer and to limit the management of the is landCouncilsintheNPs. The transfer to regions is not inherently bad, and for instance would work exceptionally wellinNorthernEuropeancountries.Theproblem is not the law but how it is developed when the main political parties that govern in the Canary Islandsshownointerestsinconservation,andan alarmingnumberofitspoliticians,includingsome who have significant responsibilities in conserva tion, have been charged with environmental crimes5.Althoughsomeimplicationsofdecentrali zationshouldbepositive,forinstancethecreation ofregionallistsandplansconsideringtheparticu larsofeachNPortheproximitytolocalspecialists and technicians with a wider knowledge of the region, the result is exactly opposite. With the proximityofthemanagementcentrestotheNPs, thelikelihoodofpatronageandcorruptionseems likelytoincreasewhileunificationofconservation criteria across the archipelagos four NPs seems destined to decrease, especially if the different islandCouncilsaregovernedbydifferentpolitical parties, which is currently the case. In addition, jointmanagementoftheNPsandtheotherNPAs ineachislandwoulddilutetherigorandresources
5. See press references in http://www.abc.es/20100322/canariascanarias/tresimputadoscoronannueva 20100322.html (last accessed August/2011); http://www.canariassemanal.com/elhierro.html (last accessed Au gust/2011); http://www.eldia.es/20110413/CANARIAS/5Esfrecuentealcaldesestenimputadosdelitos urbanisticos.html (last accessed August/2011); http://www.elpais.com/articulo/espana/corrupcion/presenta/ elecciones/elpepiesp/20110410elpepinac_1/Tes (last accessed August/2011); http://www.europapress.es/islas canarias/noticiaimputadoscanariaslogranmantenerseinstituciones20110524094822.html (last accessed Au gust/2011). frontiersofbiogeography3.3,20112011theauthors;journalcompilation2011TheInternationalBiogeographySociety
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Canarianconservationnetworkdismantled dedicated to NPs. Considering that budgets are notfixedthiswouldimplythatfundingtomanage the NPs could eventually be used in other tasks, moreconsistentwiththe"needsofthemoment". A recently created Commission of Canarian NPs, constitutedmainlyofpoliticiansandwithonlytwo advocatesforenvironmentalissues,leftasidethe present directors and conservators of the NPs. It could also happen that once transferred to the Councils, the election of new directors will not consider the balance between conservation and managementskillsthatsuchpositionrequires. TheislandCouncilsarealreadyinchargeof the management of the Canarian Network of NPAs. While some of these areas have been ac tively managed others lack any type of control. ThesituationofsimilarNPAsvariesamongislands andformosttheactionplanshavebeenpartially or barely fulfilled, so that nowadays (more than tenyearsafteritsdeclaration)itisstilleasytofind dumps, illegal constructions, invasive species, to getherwithotherpotentialemergingthreats.De spite the capacity and good work of environ mentaltechnicians,whostrugglewithbudgetcuts everyyear,theCouncilshavedemonstratedatra jectoryofinefficiencyandlackofcommitmentto the management of NPAs. Within the new Ca narianNPsframework,therabbitswillreceivethe responsibilityoftakingcareofthelettuces.
Acknowledgements
We would like to thank Rafael Loyola and three anonymous reviewers for their comments on the manuscript. We are also grateful to the editorial board of Frontiers in Biogeography for their help improvingthispaper.
References
Alberto, F., Massa, S., Manent, P., DiazAlmela, E., Ar naudHaond, S., Duarte, C.M. & Serro, E.A. (2008) Genetic differentiation and secondary contactzoneintheseagrassCymodoceanodosa across the MediterraneanAtlantic transition region.JournalofBiogeography,35,12791294.
Barber,C.,TuyaF.,BoyraC.,SanchezJerezP.,Blanch I. & Haroun R.J. (2005) Spatial variation in the structural parameters of Cymodocea nodosa seagrassmeadowsintheCanaryIslands:amul tiscaled approach. Botanica Marina, 48, 122 126. FernndezPalacios,J.M.&Whittaker,R.(2008)Canar ies. An important biogeographical meeting place.JournalofBiogeography,35,379387. FranciscoOrtega, J., SantosGuerra, A., Kim, S.C. & Crawford,D.(2000)Plantgeneticdiversityinthe Canary Islands: A conservation perspective. AmericanJournalofBotany,87,909919. Jaffre, T., Bouchet, P., Veillon, J.M. (1998) Threatened plants of New Caledonia: Is the system of pro tectedareasadequate?BiodiversityandConser vation,7,109135. Juan, C., Emerson, B.C., Orom, P. & Hewitt, G.M. (2000)Colonizationanddiversification:towards aphylogeneticsynthesisfortheCanaryIslands. TrendsinEcologyandEvolution,15,104109. Margules, C.R. & Pressey, R.L. (2000) Systematic con servationplanning.Nature,405,243253. Metzger, J.P., Lewinsohn, T.M., Joly, C.A., Verdade, L.M.,Martinelli,L.A.,Rodrigues,R.R.(2011)Bra zilian Law: Full Speed in Reverse? Science, 329, 276277. Possingham,H.P.etal.(2010)OpenlettertothePrime Minister and Leader of the Opposition, Science supporting marine protected areas, signed by 152 Australian scientists. Available from http:// www.ecology.uq.edu.au/docs/Marine% 20Reserve%20Scientist%20Ltr% 2018Aug2010.pdf(lastaccessedOctober/2011) Rodrigues, A.S.L., Andelman, S.J., Bakarr, M.I., et al. (2004)Effectivenessoftheglobalprotectedarea network in representing species diversity. Na ture,428,640643. Servicio de Biodiversidad (2009). Evaluacin de espe cies catalogadas de Canarias: Cicer canariensis [Ciccan06/2009].ConsejeradeMedioAmbien te y Ordenacin Territorial, Gobierno de Cana rias, Las Palmas de Gran Canaria. Available at http://www.gobcan.es/cmayot/ medioambiente/medionatural/biodiversidad/ especies/especies_protegidas_amenazadas/
EditedbyJoaqunHortal&MichaelNDawson
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ISSN19486596
opinionandperspectives
perspective
Thecausesandbiogeographicalsignificance ofspeciesrediscovery
Richard J. Ladle1,2,*, Paul Jepson2, Ana C. M. Malhado1, SteveJennings3andMaanBarua2
1.InstituteofBiologicalandHealthSciences,FederalUniversityofAlagoas,Macei,AL,Brazil.2.School ofGeographyandthe Environment,Universityof Oxford,SouthParks Road,Oxford, OX13QY,United Kingdom.3.OxfamGB,OxfamHouse,JohnSmithDrive,Oxford,UnitedKingdom. *Authorforcorrespondence:DrRichardJ.Ladle,InstituteofBiologicalandHealthSciences,FederalUni versityofAlagoas,PraaAfrnioJorge,s/n,Prado,Macei,AL,Brazil,57010020. email:richard.ladle@ouce.ox.ac.uk;http://www.geog.ox.ac.uk/staff/rladle.html Abstract.Therediscoveryofaspeciesthatwasputativelyconsideredtobeextinctcanprovidevaluable datatotestbiogeographicalhypothesesaboutpopulationdeclineandrangecollapse.Moreover,such rediscoveries often generate muchneeded publicity and additional funds for the conservation of rare species and habitats. However, like extinction, rediscovery is challenging to define. In this perspective wearguethatthelossofaspeciesanditssubsequentrediscoverycanbeunderstoodintermsofthe interplayamongfoursocioecologicalfactors:(1)thestateofknowledgeofspecieslossandrediscovery; (2)thepresenceofpeopleand/ororganizationswiththeinterest,motivation,resources,skillsandtech nologytofindtargetspecies;(3)theaccessibilityoftheareas,habitatsorsiteswherethespeciesare thoughttosurvive;and(4)theeasewithwhichaspeciescanbelocatedwhenitispresentwithinahabi tat. Thus, species are lost from scientific knowledge for different reasons and, consequently, not all rediscoveriesareequallysignificantforbiogeographicalresearchorconservation.Indeed,rediscoveries ofspeciesthatunderwentawelldocumenteddeclineanddisappearanceandarethereforeofgreatest potentialimportanceforbothconservationandbiogeographicalresearchappeartobepoorlyrepre sentedintheliteraturecomparedtorediscoveredspeciesthatwereonlyknownfromahandfulofmu seum specimens. Thus, carefully distinguishing between the causes of temporal gaps in zoological re cordsisessentialforimprovingtheutilityofrediscoverydataforbiogeographicalresearchandconser vationpractice. Keywords:extinction,rangecollapse,rarity,criticallyendangered,monitoring
Introduction
Rediscoveries of putatively extinct species are of great potential interest to both conservationists and biogeographers (Crowley 2011). For the for mer,rediscoverycanbeaconsiderableconserva tion policy and publicity asset (Ladle and Jepson 2008, Ladle et al. 2009) as testified by recent global initiatives: in 2009 BirdLife International launched a global bid to try to confirm the con tinued existence of 47 species of bird that have notbeenseenforupto184years(BirdLifeInter national 2009). The following year Conservation International launched its Search for lost Frogs whichinvolvesadedicatedcampaignandexpedi
tionsto18countriesseekingtolocate40species not seen for a decade or more (Conservation In ternational2010)atthetimeofwriting12spe cies have been rediscovered. Moreover, since re discovered species are typically exceedingly rare and geographically localized, new knowledge on populationstatusanddistributionsupportseffec tive conservation interventions. Finally, rediscov eries remove uncertainty from extinction risk as sessments; a confirmed new record moves the species from extinct or probably extinct and intoanIUCNthreat(ordatadeficient)category. For biogeographers, species rediscovery has both a practical and conceptual significance. From the 111
rediscoveriesinbiogeography practicalperspective,therediscoveryofaspecies that has gone unrecorded for a long period of time improves geographical knowledge about someoftheworldsrarestspecies,helpingtoad dresstheWallaceanshortfalltheinadequacyof ourknowledgeofthegeographicaldistributionsof species(Lomolinoetal.2006,Riddleetal.2011). Theshortfallcanoftenbeextreme,withaspecies knownfromjustoneorafewmuseumspecimens collecteddecadesorevencenturiesearlier.These speciesaresometimesincorrectlyassumedorde clared extinct, a phenomenon which Ladle and Jepson (2008) refer to as a Wallacean extinction. As we discuss later, these extreme examples of theWallaceanshortfallareamongstthemostfre quentlyrediscoveredspecies. Morerecently,biogeographershavestarted touseinformationonspeciesrediscoveriestotest theoriesofpopulationdeclineandrangecollapse underanthropogenicdisturbance(Fisher2011a,b; FisherandBlomberg2011).Theunderlyingideais both simple and elegant: the location of a redis covered species relative to its historical range re flects the pattern of range collapse. Thus, if an thropogenic pressures (e.g. unsustainable exploi tation) are strongest at the periphery (Channel and Lomolino 2000) the rediscovery will most likely be made near the centre of the historic range. Diana Fishers (2011a) study of 67 species ofrediscoveredmammalsfoundanumberofclear trends, although these tended to be dependent upontheecologyofthespecies.Forexample,one ofthestrongestpatternsobservedwasthatredis coveriesweregenerallymadeathigherelevations than the original record (excluding mountaintop and coastally restricted species). This provides some support for the hypothesis that higher ele vations can sometimes provide ecological refugia (TownsandDaugherty 1994)andfitswiththefre quently observed pattern of habitat destruction andpopulationextinctionprogressingfromlowto highaltitudes(Triantisetal.2010). However, like extinction, rediscovery is challenging to define. This should not be surpris ingsincerediscoveryandextinctionareconceptu ally intertwined; extinction is the permanent ab sence of current and future records while redis 112 covery reflects the temporary absence of such records. Moreover, rediscovery is the proof re quiredtorefuteahypothesisofextinction.Given the close conceptual linkage between the con ceptsofrediscoveryandextinctionitisinteresting that,untilrecently,therehavebeensofewstud ies linking patterns of rediscovery to contempo rarytheoriesofpopulationdeclineandextinction. Oneimpedimenttosuchresearchisthelackofa systematicapproachtospeciesrediscoveriesthat allow scientists to identify cases of rediscovery that have biogeographical or conservation signifi cance, and which can be subject to meaningful analysis. Here, we propose a conceptual frame work for understanding and analyzing species re discovery, based on the social, institutional and ecologicalfactorsthatcreatedthetemporalgapin occurrence data. We believe that formalizing the conceptofrediscoveryinthiswayhasthepoten tialtocreatenewmeasuresofthestateofknowl edgeoftheworldsrarestspecies,provideaquan tifiable metric to support existing endangerment categorizations, and would help to maintain the culture of biogeographical exploration that con tributestothedatasetsthatunderpinglobalcon servationtargetsetting,advocacyandmonitoring.
Conceptualframework
Thelossofaspeciesanditssubsequentrediscov erycanbeconceptualizedasaresultoftheinter playamongfoursocioecologicalaspectsofredis covery (schematically illustrated in Figure 1): (1) the state of knowledge of species loss and redis covery;(2)thepresenceofpeopleand/ororgani zations with the interest, motivation, resources, skillsandtechnologytofindtargetspecies;(3)the accessibility of the areas, habitats or sites where the species are thought to survive; and (4) the easewithwhichaspeciescanbelocatedwhenit ispresentwithinahabitat.Itshouldbenotedthat although these factors potentially apply to all losttaxa,owingtoissuesofhistoricaldataqual ity, funding and the culture of scientific explora tion,rediscoveryresearchhasfocusedalmostex clusively on herptiles, birds and mammals (cf. Scheffersetal.2011).
RichardJ.Ladleetal.
Knowledgeoflostspecies
Enormousadvanceshavebeenmadeoverthelast 40yearsinenumeratingwhichspeciesareappar entlylost.Forexample,BirdLifeInternationalhas made significant investments in compiling new andauthoritativeassessmentsofthreatenedspe cies using information from a variety of sources includingamateuranduniversityledresearchex peditions and major reviews of existing museum specimens.Inparticular,fromthemid1980stwo majorregionalRedListreviewswerecompiledfor the Americas (Collar et al. 1992) and Asia (Collar et al. 2001), the findings of which were then fed backtotheBirdLifenetworkofpioneeringprofes sional and amateur ornithologists (Tobias et al. 2006,Butchart2007). The knowledge of what is lost is compli cated, as rediscoveries can logically be split into four categories that reflect different degrees of uncertainty (and authority) about the continued existence of a target species (Table 1). An addi tionalcategorycouldpotentiallybeaddedtothis typology to account for cases where an unre corded subspecies is elevated to full species status. For example, the Sangihe Shrikethrush (Colluricincla sanghirensis) was rediscovered in 1985butitsstatusasafullspecieswasonlyestab lished in 1999 (Rozendaal and Lambert 1999). Changes in taxonomic status may have profound impactsonsurveyeffort:accordingtoRasmussen etal.(2000),thedemotionoftheSangiheWhite eye(Zosteropsnehrkorni)tosubspecificstatusby
Stresemann (1931) had the effect of making the species of only marginal, regional interest and as a consequence for many years [it] received littleattention(p.69). Fromtheperspectiveofinvestigatingrange changes, confounding different categories of re discovery could seriously influence research find ings.Forexample,wemightexpectthatallother thingsbeingequal,specieswhosehabitatorrange has not been surveyed for a significant period of timeandforwhichtherearenostrongreasonsto assumehavebecomeextinct(Table1,category4), areaslikelytoberediscoveredattheedgeorcen tre of their historic range as are betterknown species.Moreover,allfourcategoriesofrediscov ery may contain species that were only known fromasmallnumberofmuseumspecimensthe rediscoveryofwhichmaytellsusmoreaboutthe history of biogeographical exploration than the ecology of decline and extinction. Indeed, Schef fers et al. (2011) found that the majority of re cently claimed amphibian, bird and mammal re discoveries represent first documentations since their original scientific description. It should also be noted that such rare species may have re mainedunrecordedbecauseofintrinsicbiological characteristics (e.g. nocturnal habits, cryptic colouration, etc.) rather than a lack of sampling effortandthatthesefactorsneedto becarefully untangled in any analysisof patterns of rediscov ery (see McCarthy 2008; Fisher and Blomberg 2011).
Figure 1.Thefourmajordimensions ofspeciesrediscovery(seetext). frontiersofbiogeography3.3,20112011theauthors;journalcompilation2011TheInternationalBiogeographySociety
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rediscoveriesinbiogeography
Type
1.
Rediscoveryof
Example
aspeciesdeclaredextinctbyanauthori ThePohnpeiStarling(Aplonispelzelni)wasdeclared tativesource extinctbytheIUCN(1990)andrediscoveredin1995 (Buden1996) aspeciesconsideredprobablyextinct by The Sao Tome Grosbeak (Neospiza concolor) was anauthoritativesource described as probably extinct by Greenway (1967) andrediscoveredin1991(Sergeantetal.1992) a species believed to be still extant but According to the NGO BirdLife International the forwhichsubstantivesearchesoverdec Madagascar Serpent Eagle (Eutriorchis astur) was adeshavedrawnablank. notdefinitelyrecordedbetween1930and1993de spiteconsiderablesearcheffortwithinitshabitat. aspecieswhosehabitatorrangehadnot beensurveyedforasignificantperiodof time, but for which there is no real rea sontoassumehasbecomeextinct TheChestnutbelliedFlowerpiercer(Diglossaglorio sissima)wasunrecordedfor38years:since2003it has been recorded from three locations (Tobias et al.2006)
2.
3.
4.
Table1.Acrudetypologyofspeciesrediscoverybasedondecreasinglevelofcertaintythattherediscoveredspecies wasextinct.
Perhaps the most important type of redis coveryforconservationiswhereapreviouslywell knownspeciesundergoesapopulationdecline,is lostfrombiogeographicalknowledge,andisthen rediscovered.ApossibleexampleistheAustralian Pygmy Bluetongue Lizard Tiliqua adelaidensis. This rather secretive lizard was relatively well known up to its disappearance in 1959; its redis covery in 1992 (in the stomach of a snake) con firmed that the species now has a dramatically reduced geographical range (Milne and Bull 2000, p. 296). The rediscovery of the Ivorybilled Woodpecker(Campephilusprincipalis)(Fitzpatrick etal.2005)wouldbeanevenbetterexample,ex cept that this rediscovery is increasingly looking like a case of mistaken identity (Dalton 2005, 2010, Stokstad 2007). The apparent scarcity of such rediscoveries (cf. Scheffers et al. 2011) stronglysuggeststhataspecies that undergoesa well documented decline and disappearance is likelytobeextinct.However,formallytestingthis hypothesis would require good information on populationtrendsofrediscoveredspeciespriorto theiroriginaldisappearancedatathatrarelyex istforoldercasesofspeciesloss. A final aspect of the knowledge needed to findlostspeciesisthereliabilityofbiogeographic information on where to search for the species. 114
Thus, the Blackhooded Antwren (Formicivora erythronotos)wasknownonlyfroma19thCentury type specimen, for which the type locality was probablyincorrect,andwhichwasalsoputinthe wrong genus. Balchon (2007) suggests that this ledtoresearcherslookinginthewrongplace,for thewrongsortofbirdandlisteningforinappropri ate vocalizations. Thus, lost species can some timesturnupthousandsofkilometresawayfrom wheretheywerelastseen,orincompletelydiffer ent habitats. For example, the Largebilled Reed Warbler (Acrocephalus orinus) was previously known from just a single specimen collected in 1867intheSutlejValley,HimachalPradesh,India. However,alivingspecimenwastrappedinMarch 2006 at Laem Phak Bia, Phatchaburi Province, southwestThailand,over3000kmfromthetype locality(Roundetal.2007).Therenewedinterest in this species led to the unearthing of ten new museum specimens (Svensson et al. 2008) and, shortlyafterwards,tothediscoveryofabreeding population in northeast Afghanistan (Timmins et al.2010).
Institutional,scientificandtechnicalcapacity
Even when a species is identified as possibly still extant, the institutional and technical capacity to find it may not exist. Such capacity, at a global
RichardJ.Ladleetal. level,hasvariedconsiderablyovertimeandspace inresponsetovariousculturalandecologicalfac tors.Mostnotably,themainstreamingofbiodiver sity into international development following the 1992EarthSummitcreatedmanynewsourcesof fundsandemploymentopportunitiesforscientists inlessdevelopedcountries.Withrespecttobirds, this increase in local capacity coincided with the creationofBirdLifeInternationalin1993.BirdLife emerged from the International Council for Bird Preservation (founded in 1922) when its leaders devisedthecompellingpropositionofformingan international partnership, under a single name, with smaller, national, birdorientated conserva tionorganizations(JepsonandLadle2010).More generally, increased funding of expeditions by in ternational NGOs has probably been the driving forcebehindtheincreasingfrequencyofrediscov eriesofvarioustaxa(Scheffersetal.2011). Other trends within science and conserva tionalsohelpdeterminethecapacityandmotiva tion that enables rediscoveries, especially the in troduction of new technology. For example, ad vances in molecular biology have made it much easier to genetically compare preserved type specimensinmuseumswithcontemporarymate rialcollecteddirectlyoracquiredfromhuntersor from rural markets. This has opened the way for completely new ways of rediscovering lost spe cies,whereafragmentofhairorafaecalsample may be sufficient to prove the continuing exis tenceofaspeciesthathasstillnotbeenphysically observed. Anexcellentexampleofsuchatechnology aided discovery is provided by Pitra et al. (2006), whorecentlyannouncedthecontinuingexistence of the giant sable antelope (Hippotragus niger variani), a subspecies unique to Angola that was feared extinct after almost three decades of civil war. They compared the mitochondrial DNA se quences derived from old museum specimens with samples extracted from dung samples re cently collected in the field. Such remotely col lectedDNAevidencecanalsobeusedtodiscount presumed discoveries or rediscoveries. For exam ple,Hennacheetal. (2003)usedarangeoftech niques, including captive hybridization experi mentsandanalysisofmitochondrialDNAandmi crosatellites, to conclusively demonstrate the hy brid origin of the imperial pheasant (Lophura im perialis). This mysterious bird had first been cap turedin1924whenasinglepairhadbeenshipped to the private aviary of Jean Delacour in France and was not seen again until one was trapped in 1990(Hennacheetal.2003). Itisnotonlyadvancesinmolecularbiology thatarefacilitatingrediscoveries.Thereadyavail abilityofsophisticatedaudiovisualequipmenthas beenespeciallyimportantintheevolutionofbird surveying. Two such technological advances, the increased availability of less expensive sound recording and playback equipment in the late 1990s and the more recent internetbased bird sound archives, have dramatically increased the capacity of both amateurs and professionals to locate and identify rare and cryptic bird species. Moreover,advancesinthequalityofcamerasand lenses, especially digital cameras and video re corders,havealsobeenimportantindocumenting and providing definitive proof of the existence of very rare species. For example, the New Zealand StormPetrel(Pealeornismaoriana)wasidentified from the details on a digital image taken in 2003 (Stephenson et al. 2008). It had previously been known only from putative fossil material, and from three specimens collected in the 19th Cen tury,150yearsbeforeitsrediscovery.
Accessibility
Even if a species is extant and potential habitats havebeenlocated,thespeciesmaynotbefound. Accesstosuitablehabitatmaybelimitedbecause of political instability/restrictions, or simply the remotenessofpotentialsites.Althoughintheera of cheap international air travel this is arguably lessimportant,itmayhaveplayedacriticalrolein restricting the intensity of surveys and therefore the rate of rediscoveries in many parts of the globe.Examplesofrediscoveriesthatwereproba blydelayed,andpossiblyevencaused,bypolitical instability include that of the Largebilled Reed Warbler in Afghanistan (see above) and the GabelaHelmetshrike(Prionopsgabela),rediscov eredin2003inAngola(Ryanetal2004). 115
rediscoveriesinbiogeography Acloselyrelatedfactorisalackofcommu nicationwithremoteandisolatedruralcommuni tieswhomayalreadyhaveknowledgeofthecon tinuedexistenceofaputativelyextinctspecies,or of a species new to science. Thus, a productive routetoincreasingrediscoveries(andnewspecies discoveries)mightbethroughbettercommunica tion with remote tribes and communities whose knowledge of local biodiversity may extend con siderably beyond that of conservationists. How ever, Fisher and Blomberg (2011) found that hu man population overlap did not predict rediscov ery rate in mammals, possibly because expedi tionsandsurveysmayintentionallyfocusonmore remoteareas. prisingly, between 1912 and 1990 there were no records of the Night Parrot until one was hit by traffic(Bolesetal.1994).
Rediscoveriesreconsidered
Given the very loose usage of the term rediscovery and the varying factors, social and ecological, that contribute to rediscoveries, both biogeographyandconservationmaybenefitfrom adopting a stricter policy of usage. One strategy wouldbetostrictlyconfinethetermrediscovery to species categorized as extinct in the IUCN sys tem(Maceetal.2008)oraspossiblyextinct,or lostbyauthoritativesources(Table1,categories 1,2and3).Itshouldbenotedthatmanyspecies that are considered possibly extinct are listed as critically endangered in the IUCN system. For example, Fisher (2011a) restricts her analysis to rediscoveredmammalspeciesthathadbeenpre viouslyreportedasgloballyextinctorpossiblyex tinct. It should be noted, however, that this ap proachwillnotcompletelyeliminateallthecases of species that are missing through low levels of surveying. An alternative strategy could be to classify rediscovery purely in terms of the length of time withoutaformalrecord.Ifthiswereadopted,the onlyissuewouldbeanappropriatetimeframefor a given taxon. For example, De Roland et al. (2007)feltjustifiedinclaimingtherediscoveryof the Madagascar Pochard (Athya innotata) just 15 yearsafterthelastconfirmedsightingconceiva blythesameindividual. Using a simple timebased criterion would provide a single, objective definition of rediscov erywhateverthecauseofthegapinzoological records.Conservationbodiescouldpotentiallyuse thisdefinitiontoperiodicallyproducelistsofspe ciesthatmaystillbeextantand,byextension,are inneedofrediscovery.Thesecouldbecategorized according tothetimesinceaspecieswaslastre corded (e.g. <25 years ago, 2549 years ago, 50 100 years ago, >100 years ago, etc.). One advan tage of such a system would be to maintain and extend the practice of biogeographical expedi tions to remote areas. It would also help guard againsttheoveruseormisrepresentationofredis
Ecologicalfactors
The final aspect of rediscovery is the ecological characteristics of the putatively extinct species that may make verification of its continued exis tence problematic. For example, if the species is veryrareand/ordispersed,thenitmaybedifficult to locate an individual/population within an area of potentially suitable habitat. Even if the survey team is in the same area as the target species, it may still not be encountered because of pheno typic and ecological traits (e.g. cryptic coloration, lackofvocalizations,skulkingbehaviour,etc.)that reduce the probability of detection (Scheffers et al.2011).However,theevidenceforthiseffectis variable:FisherandBlomberg(2011)foundthatin mammals many ecological characteristics such as cryptic coloration and arboreal and nocturnal be haviour were not significantly associated with re discovery although smaller rediscovered mam malshadbeenmissingforlongerperiodsoftime (Fisher2011b). A possible example of ecology driving the lackofrecordsistheNightParrot,aspeciesthatis known from 23 specimens and many sightings of varying reliability from a wide geographic area of inland Australia (McDougall et al 2009). From what little information exists, the Night Parrot is crepuscular or nocturnal, cryptic, and when ap proachedwillonlyflushatclosequarters,thenfly lowovershortdistancesbeforeplungingbackinto cover(ForshawandCooper2002).Perhapsunsur 116
RichardJ.Ladleetal. coveriesinthemedia(Ladleetal.2009).Itwould offeraviablealternativetotheuseoftermssuch as possibly extinct (Butchart et al. 2006) and data deficient, and would ensure better quality ofdataforfuturebiogeographicalstudies.
Boles,W.E.,Longmore,N.W.,&Thompson,M.C.(1994) A recent specimen of the Night Parrot Geopsit tacusoccidentalis.Emu,94,3740. Buden,D.W.(1996)RediscoveryofthePohnpeiMoun tain Starling (Aplonis pelzelni). Auk, 113, 229 230. Butchart, S.H.M., Stattersfield, A.J. & Brooks, T.M. (2006)Goingorgone:definingPossiblyExtinct species to give a truer picture of recent extinc tions. Bulletin of the British Ornithology Club, 126a,724. Butchart,S.H.M(2007)Birdstofind:areviewoflost, obscure and poorly known African bird species. BulletinoftheAfricanBirdClub,14,138157. Channell, R. & Lomolino, M.V. (2000) Trajectories to extinction: spatial dynamics of the contraction ofgeographicalranges.JournalofBiogeography, 27,169179. Collar, N.J., Gonzaga, L.P., Krabbe, N.,Madrono Nieto, A., Naranjo, L.G., Parker III, T.A. & Wege, D.C. (1992) Threatened birds of the Americas. 3rd edn.SmithsonianInstitutionPress,Washington. Collar,N.J.,Andreev,A.V.,Chan,S.,Crosby,M.J.,Subra manya, S. & Tobias, J.A. (2001). Threatened BirdsofAsia.BirdlifeInternational,Cambridge. Conservation International (2010) The search for lost frogs. Available from http:// www.conservation.org/campaigns/lost_frogs. Accessed24January2012. Crowley, B. (2011) Extinction and rediscovery: where thewildthingsare.JournalofBiogeography,38, 16331634. Dalton, R. (2005) Sighting of extinct bird may have been a case of mistaken identity. Nature, 436, 447. Dalton,R.(2010)Stilllookingforthatwoodpecker.Na ture463,718719. De Roland, L.R., Sam, T.S., Rakotondratsima, M.P.H. & Thorstrom, R. (2007) Rediscovery of the Mada gascar Pochard Aythya innotata in Northern Madagascar.BulletinoftheAfricanBirdClub14, 171174. Fisher, D.O. (2011a) Trajectories from extinction: where are missing mammals rediscovered? GlobalEcology&Biogeography,20,415425. Fisher,D.O.(2011b)Cost,effortandoutcomeofmam mal rediscovery: neglect of small species. Bio logicalConservation,144,17121718. Fisher,D.O.&Blomberg,S.P.(2011)Correlatesofredis covery and the detectability of extinction in mammals. Proceedings of the Royal Society B: BiologicalSciences,278,10901097. Fitzpatrick, J.W., Lammertink, M., Luneau Jr., et al. (2005) Ivorybilled woodpecker (Campephilus principalis) persists in continental North Amer ica.Science,308,14601462.
Conclusions
The rediscovery of a species that was thought to beextinctcangenerateglobalinterestandrepre sents a real opportunity for conservationists to reassertcorevaluesandraisefundsthatmayhelp protect poorly known habitats. Moreover, redis coveries provide a unique source of information abouttherarestandleastknownspecies(forcer tain taxa) that can be used to investigate bio geographic theories about range loss and extinc tion. Both of these important agendas would benefitfromagreatersystematizationofthecon cept of rediscovery, acknowledging the varying causes (both social and ecological) of gaps in the temporalrecordsofrarespecies. Insummary,thestudyofrediscoveriespro vides a wonderful opportunity to assess both the subtle ecological and biogeogeographical charac teristicsofexceptionallyrarespeciesofwellstud iedtaxasuchasamphibians,birdsandmammals, andthefascinatinghistoricalandculturaltrendsin zoologicalsurveyingandexploration.Considerable effortsarebeingmadetountangletheseinteract ing factors (Fisher 2011a,b; Fisher and Blomberg 2011,Scheffersetal.2011),whiletherecenttar geting of lost species by international conserva tion NGOs is generating considerable amounts of valuablenewdata.Nevertheless,thelackofredis covered species that were previously well known and which had undergone a well documented process of population decline, fragmentation and local extinction (Scheffers et al. 2011) remains a worryingtrendforglobalconservation.
References
Balchon,C.(2007)Backfromthedead!Apotpourriof recent rediscoveries in the Neotropics. NeotropicalBirding,2,411. BirdLife International (2009) Quest launched to find lost birds. Available from http:// www.birdlife.org/news/news/2009/08/ lost_and_found.Accessed25March2010.
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Forshaw, J.M. & Cooper, W.T. (2002) Australian par rots.3rdedn.AlexanderEditions,Robina. Greenway,J.C.(1967)Extinctandvanishingbirdsofthe world.2ndEdition.Dover,NewYork. Hennache, A., Rasmussen, P., Lucchini, V., Rimondi, S. & Randi,E. (2003) Hybridorigin of the imperial pheasant Lophura imperialis (Delacour and Jabouille, 1924) demonstrated by morphology, hybrid experiments, and DNA analyses. Biologi calJournaloftheLinneanSociety,80,573600. Jepson,P.J.&Ladle,R.J.(2010)Conservation:abegin nersguide.OneWorld,Oxford. Ladle, R.J. & Jepson, P.R. (2008) Toward a biocultural theory of avoided extinction. Conservation Let ters,1,111118. Ladle, R.J., Jepson, P., Jennings, S. & Malhado, A.C.M. (2009) Caution with claims that a species has beenrediscovered.Nature,461,723. Lomolino, M.V., Riddle, B.R. & Brown, J.H. (2006) Bio geography.3rdedn.Sinauer,Sunderland,MA. Mace,G.M.,Collar,N.J.,Gaston,K.J.,HiltonTaylor,C., Akakaya, H.R., LeaderWilliams, N., Milner Gulland,E.J.&Stuart,S.N.(2008)Quantification of extinction risk: IUCN's system for classifying threatened species. Conservation Biology, 22, 14241442. McDougall,A.,Porter,G.,MostertM.,CupittR.,Cupitt S., Joseph, L., Murphy S., Janetzki H., Gallagher A.&BurbidgeA.(2009)AnotherpieceinanAus tralian ornithological puzzle a second Night Parrot is found dead in Queensland. Emu, 109, 198203. McCarthy,M.A.(1998)Identifyingdecliningandthreat enedspecieswithmuseumdata.BiologicalCon servation,83,917. Milne,T.&Bull,C.M.(2000)Burrowchoicebyindividu als of different sizes in the endangered pygmy blue tongue lizard Tiliqua adelaidensis. Biologi calConservation,95,295301. Pitra,C.,VazPinto,P.,OKeeffe,B.W.J.,WillowsMunro, S., Jansen van Vuuren, B. & Robinson, T.J. (2006) DNAled rediscovery of the giant sable antelopeinAngola.EuropeanJournalofWildlife Research,52,145152. Rasmussen,P.C.,Wardill,J.C.,Lambert,F.R.,&Riley,J. (2000) On the specific status of the Sangihe Whiteeye Zosterops nehrkorni, and the taxon omyoftheBlackcrownedWhiteeyeZ.atrifrons complex.Forktail,16,6980. Riddle,B.,Ladle,R.J.,Lourie,S.&Whittaker,R.J.(2011) Basic biogeography: estimating biodiversity and mappingnature.In:Ladle,R.J.&Whittaker,R.J. (Editors) Conservation Biogeography. Oxford UniversityPress,Oxford,pp.4792. Round,P.D.,Hansson,B.,Pearson,D.J.,Kennerley,P.R. & Bensch, S. (2007). Lost and found: the enig matic largebilled reed warbler Acrocephalus orinus rediscovered after 139 years. Journal of AvianBiology,38,133138. Rozendaal, F.G. & Lambert, F.R. (1999) The taxonomic and conservation status of Pinarolestes sanghirensisOustalet1881.Forktail,15,113. Ryan, P.G.,Sinclair, I., Cohen, C.,Mills, M.S.L., Spottis woode,C.&Cassidy,R.(2004)Theconservation status and vocalisations of threatened birds from the scarp forest of the Western Angola Endemic Bird Area. Bird Conservation Interna tional,14,247260. Scheffers, B.R., Yong, D.L., Harris, J.B.C., Giam, X. & Sodhi, N.S. (2011) The Worlds rediscovered species: back from the brink? PLoS ONE 6, e22531. Sergeant, D.E., Gullick, T., Turner, D.A. & Sinclair, J.C. (1992) The rediscovery of the So Tom Gros beak Neospiza concolor in southwestern So Tom. Bird Conservation International, 2, 157 159. Stephenson, B.M., Flood, R., Thomas, B. & Saville, S. (2008) Rediscovery of the New Zealand storm petrel (Pealeornis maoriana Mathews 1932): two sightings that revised our knowledge of stormpetrels.Notornis,55,7783. Stresemann, E. (1931) Die Zosteropiden der indo australischen Region. Mitteilungen aus dem ZoologischenMuseumBerlin,17,201238. Stokstad, E. (2007) Gambling on the ghost bird. Sci ence,317,888892. Svensson,L.,PrsJones,R.,Rasmussen,P.C.&Olsson, U. (2008) Discovery of ten new specimens of largebilled reed warbler Acrocephalus orinus, and new insights into its distributional range. JournalofAvianBiology,39,605610. Timmins, R.J., Mostafawi, N., Rajabi, A.M., Noori, H., Ostrowski, S., Olsson, U., Svensson, L. & Poole, C.M. (2010) The discovery of Largebilled Reed Warblers Acrocephalus orinus in northeastern Afghanistan.BirdingASIA,12,4245. Tobias,J.A.,Butchart,S.H.M.&Collar,N.J.(2006).Lost and found: a gap analysis for the Neotropical avifauna.NeotropicalBirding,1,422. Towns,D.R.&Daugherty,C.H.(1994)Patternsofrange contractionsandextinctionsintheNewZealand herpetofauna following human colonization. NewZealandJournalofZoology,21,325339. Triantis, K.A., Borges, P.A.V., Ladle, R.J., et al. (2010) Extinction debt on oceanic islands. Ecography, 33,110.
EditedbyJanBeck
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GettingtoknowIBSEarlyCareerMembers
The International Biogeography Society (IBS), foundedjust10yearsago,isfastgrowingbothin termsofmembersandactivitiesoffered(Fieldand Heaney 2011). Students and earlycareer bio geographers are also becoming increasingly in volvedwithintheIBS.From2002to2010,thepro portionofnewmemberswhoarestudentsjoining theIBSeachyearhasincreasedfrom23%to48%. Currently, student members comprise 35% of IBSs 740 members. The IBS, aware of the rising importanceoftheseyoungermembers,hasbeen tryingtoincreasethebenefitsavailableforthem. In addition to the student travel grants, poster awards and discussion groups held at the IBS meetings, the IBS is trying to foster interaction amongstudentsandpostdocs,whichrecentlycul minated in the first IBS Early Career conference that was held at Oxford University from 23 to 25 September 2011 (http://www.biogeography.org/ html/Meetings/index.html). With the intention of getting to know its earlycareer members (herein ECM) and learning theiropinionsontheservicesprovidedbytheIBS and on how these can be improved, the IBS in vitedECMtoparticipateinasurveythatwasheld in June 2011. Of the 48 ECM that completed this survey,11%wereJuniorPostdocs,75%werePhD students,8%wereMastersstudents,and6%were undergraduate students. Around 17% were aged between2025years,49%were2630years,23% were 3135 years, and 11% were more than 35 years young; 56% were female and 44% were male. Although most ECM are currently affiliated either with North American or European institu tions (50% and 33% respectively; total of 42 an swers), they represent a total of 24 nationalities; 26% are from North America, 17% from Central and South America, 15% from Northern Europe, 28% from Southern Europe, and the other 12% from Australia/New Zealand, the Middle East, Af ricaandAsia.ECMworkonaverybroadrangeof topics, from species distribution patterns (the most mentioned topic), to evolutionary biogeog raphy,dispersalandcolonization,biogeographyof species traits, island biogeography, phylogeogra phy,globalchangebiology,marinebiogeography, orpaleobiogeography,amongothers.Theirbroad interests are also reflected in the fact that most ECM are also affiliated with societies focusing on diverse topics, including ecology, evolution, con servation,paleontology,geography,botany,mam malogy, entomology, etc. These are indeed very encouraging results that show the IBS is reaching youngresearchersfromawidevarietyofresearch topicsandgeographiclocations. In general terms, the IBS is meeting ECM needs (25% responded that the IBS is doing this very well, 60% fairly well). However, there is roomforimprovement(15%respondednotvery well), and several suggestions were made; re sponsestoopenendedquestionsemphasizedthe need for more offyear meetings (regional meet ings,workshops,etc.),morejobs/grantannounce ments, more travel grants, online teaching re sources, more talks at the IBS meetings by younger researchers and more opportunities to meet other researchers. The IBS is already work ing towards improving the services it provides to allitsmembers,andnewactionsarebeingmade toadoptsuggestions. ThefirstactionwastosupporttheIBSEarly Career conference (for students and biogeogra pherswhohavefinishedtheirPhDsinthepastfive years). Almost ninety young researchers partici pated and had the chance to present their work, and to interact with each other and with the IBS board members. This conference was organized into ten different sessions that covered several aspects of macroecology, island biogeography, phylogeography,paleobiogeography,evolutionary biogeographyandconservationbiogeography. Second, we are also working towards in creasingregularcommunicationamongIBSmem bers. One way of doing this is through online so cial networks, such as Facebook, and other web based platforms (e.g. the IBS blog; http:// biogeography.blogspot.com/). Currently, the IBS hasaFacebookgroupwith~590members,where 119
membershipcorner anyonecanpostannouncements,shareideasand publications of general interest, start discussions and interact with other members. Most ECM are in fact Facebook users (80%; only 7% are Twitter users),butonly8%ofthesemembersreadtheIBS Facebook page on a weekly basis, and 44% actu ally never read it (31% read it once per month, and 17% every 36 months). Regarding the IBS blog,againonlyasmallnumberofpeoplereadit onaweeklybasis(6%),withmostpeoplereading it once per month (38%; 31% read it every 36 monthsand25%neverreadit).Anotherplatform theIBShasforcommunicatingwithitsmembers, and to foster communication between its mem bers, is the online journal Frontiers of Biogeogra phy(http://www.biogeography.org/html/fb.html). This journal has a section especially devoted for thispurposethemembershipcornerofwhich most ECM were not aware (66%). Thirtysix per centofECMreadeveryissue,while31%read23 issues per year (27% read it rarely and only 6% never read it). Main sections of interest to the ECM are (i) minireviews on a particular taxon, biogeographic topic, or question, (ii) thesis ab stracts, and (iii) symposium/congress summaries. In fact, 88% showed interest in submitting a manuscripttoanyofthesesections. Oneofthemostimportantactivitiesorgan ized by the IBS is the biennial meeting. The next onewillbeheldatFloridaInternationalUniversity in Miami, Florida, in January 2013 (http://www. biogeography.org/html/Meetings/2013). Most ECM are planning to attend this meeting (79%) andwouldprefertogiveatalk(51%;23%prefera poster presentation and 26% have no particular preference). One of IBS concerns is to maximize compatibility between high quality talks and fair representation of researchers from different countries, gender, and career stages. There was almost an even split among ECM on favoring a similar number of talks by established and younger researchers, and having more talks by senior researchers plus some younger ones (40% and 43%, respectively; 11% would prefer to have mainlyseniorresearchersand6%showednopref erence). There was no overwhelming support for studentonly sessions in future meetings (55% found it important), but most respondents showed some willingness to extend their stay in order to attend this type of event (83%). In the previous meetings, students (particularly those who have been awarded with a student travel grant) have been invited to attend discussion groups, where senior biogeographers lead the discussion on several subjects, from career and publishing advice to specific research topics. Those who have attended these student discus sion groups in past meetings (41%) found them helpful(63%).Suggestionsfordiscussiontopicsin futuremeetings,otherthanthosealreadycovered in these discussion groups, included advanced analysis in biogeography and partnerships and international activities among researchers. There was some support for future offyear meetings (33% found it useful; 61% said it was somewhat useful, and over 90% said they would at least try toattend),especiallyifthesearefocusedonspe cificresearchtopicsandmethodologies(31%and 29%,respectively;therewasatiebetweenmeet ingsonspecificgeographicrealmsandonabroad scope within biogeography 20% each). Some respondents also called for workshops and semi nars, online courses, crosssociety ventures to boost interaction between similarly oriented aca demics and excursions into biogeographically in teresting regions covering a broad range of taxa. There was also a significant interest in having a showcaseatthenextIBSmeetingoffundingagen ciesfromdifferentcountries(70%),withmostre spondentsbeingwillingtoprovideinformationon thismatter(55%). The longterm success of any growing soci etydependsontheinvolvementandinterestofits youngest members. Were fortunate that many ECM have shown willingness to get involved in promotingcommunicationbetweenIBSmembers,
DidyouknowthatanymemberoftheIBSmayraiseanissueorappealadecisionofthegover ningBoardofDirectorsbyplacingamatterbeforetheBoardofDirectorsfordiscussion? IfthereisamatteryouwouldlikediscussedatthenextBoardmeeting,writetothesociety's Secretary(checkcurrentlistofofficersathttp://www.biogeography.org/).

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membershipcorner to help organizing offyear activities, and to sub mitmanuscriptstoFrontiersofBiogeography.The IBSwantstohearandsharemoreoftheearlyca reer members opinions and ideas; this article is intended as both thanks and encouragement for your active involvement, especially in the readily accessibleplatformssuchasFrontiersofBiogeog raphy and Facebook. Finally, we would like to thank all the members who participated in this survey, and particularly those who have shown interestindevotingsomeoftheirtimetothesoci ety.Welookforwardtoworkingwithandforyou inthecomingyears.
AnaM.C.Santos
IBSStudentatLarge;DepartamentodeEcologia, InstitutodeCinciasBiolgicas,UniversidadeFederal deGois,Brazil. email:ana.margarida.c.santos@googlemail.com
References
Field,R.&Heaney,L.R.(2011)Lookingtothefutureof theIBS:the2011IBSmembershipsurvey.Fron tiersofBiogeography,3,7173.
EditedbyMatthewHeard
fromthesociety
Callforproposalsforhosting7thBiennialConferenceoftheIBS
Weareseekingproposalsforhostingthe7thbien nialconferenceoftheInternationalBiogeography SocietytobeheldinearlyJanuary2015.Proposals should be submitted by individuals who are inte restedinchairingthelocal(host)committee.The dutiesofthelocalhostincludeconductingcontra ct negotiations with the venue and the hotel as wellasalllocallogisticsincludingfieldtriporgani zation and production of the abstract bo ok. Minimum requirements of the venue are 1) oneauditoriumwithacapacityof450550people (2days),2)threeorfoursmallerroomswithaca pacity of 75150 people (1 day), and 3) various smaller meeting rooms. The IBS is interested in holdingthebiennialconferenceinlocationsfairly convenient with respect to the majority of its membership base in North America and Euro pe.Locationsofpast(andupcoming)conferences can be seen here: http://www.biogeography.org/ html/Meetings/index.html. Please include the following information in theproposal: 1. Locationofthemeeting(city)andthehostins titutionororganization. 2. Whatwouldbethebenefitofhostingthecon ferenceatthislocation? 3. Actual site of the meeting and the capacity of theauditorium. 4. Space for poster sessionsgeneral size and lo cationrelativetotheauditorium. 5. Approximate cost for threeday use of the ve nue. A specific quote is not needed, but evi denceofthepricecompetitivenessiscrucial. 6. Transportation infrastructure, including travel fromairport. 7. Attractions in the vicinity of the conference site,includingfieldtrippotential. 8. Whowouldpotentiallyserveonthelocalorga nizingcommittee? Proposals from prospective hosts of the biennial conference must be received before 20 January 2012. Please send proposals by email to Daniel Gavin, IBS VicePresident for Conferences atdgavin@uoregon.edu.
DanGavin
IBSVicePresidentforConferences; DepartmentofGeography,UniversityofOregon,USA. email:dgavin@uoregon.edu
Erratum 11 Feb 2011:Theoriginalarticlepublishedon09Feb2011 incorrectlyidentifiedDanGavinasthe"IBSStudentatLarge".
Ifyouwanttoannounceameeting,eventorjobofferthatcouldbeofinterestfor(some)bio geographers, or you want to make a call for manuscripts or talks, please contact us at ibs@mncn.csic.esandfrontiersofbiogeography@gmail.com.
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Jobannouncements
Three Professorships and One TenureTrack Lectureship
UniversityofCalifornia,Merced,USA TheSchoolofNaturalSciencesattheUniversityof California,Mercedseeksapplicantsforfourfacul typositions: Ecology (Full orAssociatewithtenu re, or Assistant tenuretrack), Systems Biology (Assistant tenuretrack), and Biostatistics (Assistanttenuretrack),andonetenuretrackBio logy Lecturer. For the Ecology position, we seek outstanding individuals with research interests in anyecologicalfieldusingexperimental,field,com putational, and/or theoretical approaches and working at population to global scales. The Sys temsBiologypositionincludesresearchareasthat usecomprehensivedatasetsandmultipletypesof analysistorelateoverallbiologicalfunctiontoun derlying biochemical or biophysical processes for predictive understanding. The Biostatistics rese arch areas of interest include statistical methods for experimental design, epidemiology, medical informatics,evolutionarybiology,sequencebioin formatics, genomics, evolution of microbial sys tems and pathogens, and systems biology. The Lecturer position closely parallels a tenuretrack Assistant Professor but with an emphasis on un dergraduate education. All applicants must be able to teach effectively at both undergraduate andgraduatelevels.Formoreinformationandto apply go to: http://jobs.ucmerced.edu/n/ academic/listings.jsf;jsessionid=95FADBAFFF4C13 F912A3B023DA4F1F80?seriesId=1 Interested applicants should submit mate rials online. Applications will be considered star ting05December2011(Biostatistics,SystemsBio logy professorships), or 16 December 2011 (Ecology professorship and Biology Lecturer). UC MercedisanAA/EOPemployer.
upcomingevents
VIPCAMolecularEcology
47February2012Vienna,Austria http://www.vipca.at/MOLECOL/
VertNetbiodiversityinformaticstraining workshop
2430June2012Boulder,USA http://vertnet.org/about/BITW.php
AnnualConferenceoftheSocietyforTropical Ecology(gt)
Islandsinlandandseascape:Thechallengesoffrag mentation 2225February2012Erlangen,Germany http://www.gtoeconference.de/
97thESAAnnualMeeting
LifeonEarth:Preserving,Utilizing,andSustainingour Ecosystems 510August2012Portland,USA http://esa.org/meetings/
6th Annual Meeting of the Specialist Group onMacroecologyoftheEcologicalSocietyof Germany,AustriaandSwitzerland(Gf)

29February2March2012Frankfurt,Germany http://www.bikf.de/
3rd European Congress of Conservation Biol ogy

Conservationontheedge 28August1September2012Glasgow,UK http://www.eccb2012.org/
21st Workshop of the European Vegetation Survey(EVS)

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6thInternationalConferenceoftheIBS
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table of contents
thescientificmagazineoftheInternationalBiogeographySociety
volume3,issue3November2011
newsandupdate update:Speciesareacurvesandtheestimationofextinctionrates,byJ.Beck update:Extinctorextant?Woodpeckersandrhinoceros,byR.Ladle update:Climatewars,byJ.Beck update:Emergingresearchopportunitiesinglobalurbanecology,byF.A.LaSorte update:Beyondtaxonomicalspace:largescaleecologymeetsfunctionalandphylogeneticdiversity,byM.V. Cianciaruso bookreview:Amangrovecompendium,byU.Berger
ISSN19486596
81 83 84 85 87 91
bookreview:Acomprehensivefoundationfortheapplicationofbiogeographytoconservation,byT.Newbold 93 bookreview:Anewencyclopediaforbiologicalinvasions,byR.A.Francis bookreview:ApiscinehistoryoftheNeotropics,byA.E.Magurran booksnotedwithinterest thesisabstract:ApplyingspeciesdistributionmodelingfortheconservationofIberianprotectedinvertebra tes,byR.M.Chefaoui opinionandperspectives opinion:PoliticalerosiondismantlestheconservationnetworkexistingintheCanaryIslands,byJ.M.Fernn dezPalacios&L.deNascimento perspective:Thecausesandbiogeographicalsignificanceofspeciesrediscovery,byR.J.Ladleetal. membershipcorner fromthesociety:GettingtoknowIBSEarlyCareerMembers,byA.M.C.Santos fromthesociety:Callforproposalsforhosting7thBiennialConferenceoftheIBS,byD.Gavin Jobannouncements Upcomingmeetings
frontiersofbiogeographycopyrightnotice Copyright2011InternationalBiogeographySociety(IBS)undera CreativeCommonsAttributionNonCommercialNoDerivatives(CCANCND) license.Allrightsreserved.ItisstrictlyforbiddentoalterthejournalcontentsinanymannerwithouttheexpresswrittenpermissionoftheIBS.It isalsostrictlyforbiddentomakecopiesofwholeissuesofthisjournalforanycommercialpurposewithouttheexpresswrittenpermissionofthe IBS.TheIBSholdstherightforthepassivedistribution(i.e.throughitspublicationontheInternet)ofanypartorthewholeissueofthejournal duringoneyearafteritspublication.Anyactivedistributionofanypartorthewholeissueofthejournalisexplicitlypermittedsincethedateof publication, and any passive distribution is explicitly permitted after one year of the date of publication. Any individual and/or institution can download,readand/orprintacopyofanyarticleorthewholejournalfornoncommercialeducationalornoncommercialresearchpurposesat anytime.Thisincludesanexpresspermissiontousearticlesfornoncommercialeducationalpurposesbymakinganynumberofcopiesforcourse packsorcoursereservecollections.Academicinstitutions/librariesmayalsostorecopiesofarticlesandloanthemtothirdparties.Allcopiesof articlesmustpreservetheircopyrightnoticewithoutmodification.AllarticlesarecopyrightedbytheirauthorsunderauniversalCreativeCom monsAttributeLicense(CCAL).Thislicensepermitsunrestricteduse,distribution,andreproductioninanymedium,providedtheoriginalauthor andsourcearecredited.Allauthorsendorse,permitandlicensetheIBStograntanythirdpartythecopyinganduseprivilegesspecifiedabove withoutadditionalconsiderationorpaymenttothemortotheIBS.Theseendorsements,inwriting,areonfileintheofficeoftheIBS.Consult authorsforpermissiontouseanyportionoftheirworkinderivativeworks,compilationsortodistributetheirworkinanycommercialmanner. FromtheIBSconstitution:"Bylaw10.Publications.Alltitles,copyrights,royaltiesorsimilarinterestsintaperecordings,booksorothermaterials preparedfortheInternationalBiogeographySocietyIncactivitieswillbeheldsolelybytheInternationalBiogeographySocietyIncandinthename oftheInternationalBiogeographySocietyInc.".And"Article8.Publications.ThepublicationsoftheSocietyshallincludejournals,newsletters,and suchotherpublicationsastheGoverningBoardofDirectorsmayauthorize." WegratefullyacknowledgeEvolutionaryEcology,Ltd.andMikeRosenzweiginparticularfortheadviceoncopyrightmatters.
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A comprehensive foundation for the application of biogeogra phytoconservation

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Field guide Afghanistan: Flora and Communityecology PeterJ.Morin vegetation

2011,2ndedition,WileyBlackwell,407pp. 90(Hardback),34.99(Paperback) ISBN9781444338218/9781405124119 http://www.wiley.com/

opinion and perspectives

1. Seedifferentreactionsathttp://www.nodescatalogacion.com,http://www.wwf.es,http://www.greenpeace.org, http://www.atan.org, http://www.ecologistasenaccion.org, http://especiesamenazadascanarias.blogspot.com, http://ecooceanos.blogspot.com,http://www.seo.org,.

Figure 1.Thefourmajordimensions ofspeciesrediscovery(seetext). frontiersofbiogeography3.3,20112011theauthors;journalcompilation2011TheInternationalBiogeographySociety

DidyouknowthatanymemberoftheIBSmayraiseanissueorappealadecisionofthegover ningBoardofDirectorsbyplacingamatterbeforetheBoardofDirectorsfordiscussion? IfthereisamatteryouwouldlikediscussedatthenextBoardmeeting,writetothesociety's Secretary(checkcurrentlistofofficersathttp://www.biogeography.org/).

6th Annual Meeting of the Specialist Group onMacroecologyoftheEcologicalSocietyof Germany,AustriaandSwitzerland(Gf)

3rd European Congress of Conservation Biol ogy

21st Workshop of the European Vegetation Survey(EVS)

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