1 - The International Biogeography Society

frontiers of biogeographyvol. 4, nº 1 - april 2012the scientific magazine of theInternational Biogeography SocietyISSN 1948-6596 – freely available at http://www.biogeography.org/

frontiers of biogeographythe scientific magazine of the International Biogeography Societyvolume 4, issue 1 - april 2012frontiers of biogeography is published by the International Biogeography Society (IBS), an international and interdisciplinary societycontributing to the advancement of all studies of the geography of naturefrontiers of biogeography is available at http://escholarship.org/uc/fb and the IBS website: http://www.biogeography.org/html/fb.htmlfrontiers of biogeography aims to be a forum for biogeographers and a way to disseminate research in biogeography to the generalpublic; our scope includes opinions, perspectives, and reviews, symposia proceedings, research letters, book reviews, research updates,interviews, and articles on how to teach, disseminate and/or apply biogeographical knowledge. Research letters and symposium proceedingsmay include novel analyses of original datasets (see editorial instructions). Manuscripts should be submitted via eScholarship athttp://escholarship.org/uc/fb. Editorial enquiries should be made to frontiersofbiogeography@gmail.com, unless stated otherwise.frontiers of biogeography uses a publication agreement based on the Creative Commons scheme to ensure that the authors retain fullintellectual property (IP) rights on their work, and that this is freely available for any non-commercial use. Under this agreement, the IBSretains only the copyright of the journal compilation under a Creative Commons Attribution Non-Commercial No Derivatives (CCANCND)license. The authors have full IP over their texts under an universal Creative Commons Attribute License (CCAL), being able to distributetheir work (including the original PDFs) actively from the day of publication, and passively from one year after (see the full license informationat the end of the issue).you can find information about the International Biogeography Society at http://www.biogeography.org/; for the latest job announcementsand 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).editorial boardeditor-in-chief:Joaquín Hortal – Museo Nacional de Ciencias Naturales (CSIC),Spain and Universidade Federal de Goiás, Brazilassociate editors:Antje Ahrends – Royal Botanic Garden Edinburgh, UKJan Beck – University of Basel, SwitzerlandJessica Blois – University of Wisconsin, Madison, USAChris Burridge – University of Tasmania, AustraliaMarcus V. Cianciaruso – Universidade Federal de Goiás, BrazilMarkus Eichhorn – University of Nottingham, UKRoy Erkens – Maastricht University, The NetherlandsCamilla Fløjgaard – Aarhus University, DenmarkDan Gavin – University of Oregon, USAMatthew J. Heard – Brown University, USADavid G. Jenkins – University of Central Florida, Orlando, USAFrank A. La Sorte – Cornell lab of Ornithology, USARichard Ladle – Universidade Federal de Alagoas, Brazil and OxfordUniversity, UKRichard Pearson – American Museum of Natural History, USAThiago F. Rangel – Universidade Federal de Goiás, BrazilWillem Renema – NCB Naturalis, The NetherlandsNúria Roura-Pascual – Universitat de Girona, SpainSpyros Sfenthourakis – University of Cyprus, CyprusISSN 1948-6596deputy editors-in-chief:Michael N Dawson – University of California, Merced, USARichard Field – University of Nottingham, UKeditorial assistant:Lauren Schiebelhut – University of California, Merced, USAadvisory board:Miguel B. Araújo – Museo Nacional de Ciencias Naturales (CSIC),Spain and Universidade de Évora, PortugalLawrence R. Heaney – Field Museum of Natural History, Chicago,USADavid G. Jenkins – University of Central Florida, Orlando, USARichard Ladle – Universidade Federal de Alagoas, Brazil and OxfordUniversity, UKMark V. Lomolino – State University of New York, USAIBS V. P. for Public Affairs & CommunicationsInternational Biogeography Society officers 2011-2012President: Lawrence R. HeaneyPresident Elect: Rosemary GillespieVP for Conferences: Daniel GavinVP for Public Affairs & Communications: Michael N DawsonVP for Development & Awards: George StevensSecretary: Richard FieldTreasurer: Lois F. AlexanderDirector-at-large: Catherine GrahamDirector-at-large: Kathy WillisStudent-at-large: Ana M. C. SantosFirst Past President: James H. BrownSecond Past President: Mark V. LomolinoThird Past President: Brett R. RiddleFourth Past President: Vicki FunkFifth Past President: Robert J. WhittakerUpcoming meeting host (ex officio): Kenneth FeeleyPast Graduate student representative (ex officio): Matthew Heardcover: Caterpillar with cocoons of an ectoparasitoid wasp from the Braconidae family. Picture by 8 th star, availableat Wikipedia Commons.

editorial ISSN 1948-6596editorialAdvancing Frontiers, with a prospectiveAbout this time last year, we ran a retrospectiveeditorial charting the growth of Frontiers of Biogeography,this scientific magazine published bythe International Biogeography Society (Dawsonet al. 2011). We outlined its development sincethe coming-together of the IBS Newsletter teamwith the vision of the original Frontiers of Biogeographybook—to provide a series of integrativeand interdisciplinary volumes published anddeveloped in association with the InternationalBiogeography Society (IBS; Lomolino and Heaney2004; Hortal and Dawson 2009). The publicationof this issue marks the next stage in developmentof the journal.For the past year, we have been workingwith staff at eScholarship 1 to deliver a professionalsociety journal that meets the highest standardsin publishing and research. The result is acombination of the benefits of an establishedcommercial journal and those of Open Access(remarkably including low or no cost to the authorand zero cost for readers) brought to you by theeditorial team you know well as your peers in theIBS. By joining with eScholarship, which has 10years’ experience in Open Access publishing and acommitment to advancing this model of disseminatingscience, we have made it easier for you topublish in this peer-reviewed journal. Meanwhile,the editorial team is freed to concentrate on advancingthe journal and helping publish your recentadvances in biogeography.With this new format, Frontiers of Biogeographyspecifically aims to complement our growingcore of Opinions, Perspectives, and Reviewswith publications in two additional categories:Research Letters, which are concise manuscriptspresenting original scientific research, and Proceedingsof conferences, symposia, workshopsand other scientific meetings 2 . Further benefits toyou are publications rapidly disseminated withinthe society and to the broader community, withindexing in Google Scholar (we also are workingon listing in the Directory of Open Access Journals 3and in proprietary databases), monthly updateswith statistics on views and downloads of yourarticles, and uninhibited access to your work byresearchers in any institution and country. Manyof these benefits have always been available withFrontiers of Biogeography, but they are attainedmore easily with eScholarship.Frontiers of Biogeography is now 11 issues,and not quite three years, old (the first issue waspublished in September 2009). During this period,we have maintained an average of over 35 pagesper issue (over 40 in Volume 3) of increasinglydiverse content, including opinions andperspectives, book reviews, thesis abstracts, newsand comment on the current literature, andupdates for the IBS membership. This has beenaccompanied by a small but steady increase in thenumber of citations to the journal (e.g. Fagin andHoagland 2011, Gibbons et al. 2011, Dawson2012, Smith and Lundholm 2012, Peterson andLieberman 2012). Such presence in themainstream literature continues to be pushedforward by our long-standing series of contributedopinion and perspective articles such as Ladle etal. (2011), Scheiner (2011), and Beckage et al.(2012); we expect this will increasingly becomethe case for Research Letters and Proceedings too.This growth would not have been possible withoutthe support of IBS members in submitting articles;it is also the result of the work of a dedicatedteam of Associate Editors 4 , including senior andearly-career scientists with a great enthusiasm forthe diversity of biogeographic study. It is ourprivilege to work with them and for you inadvancing Frontiers alongside the growth of the1 http://escholarship.org/2 http://escholarship.org/uc/search?entity=fb;view=submissionguidelines describes article types and provides guidancefor authors3 http://www.doaj.org/4 http://escholarship.org/uc/search?entity=fb;view=editorialboardfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society1

news and updatediscipline and its leading society, and we areexcited to provide a new venue for Frontiers thatreflects the long-standing values of science and isin the vanguard of publishing.Michael N Dawson 1 , Richard Field 2 andJoaquín Hortal 31 Deputy Editor‐in‐Chief, Frontiers of Biogeography andIBS Vice‐President for Public Affairs and Communications2 Deputy Editor‐in‐Chief, Frontiers of Biogeography andIBS Secretary3 Editor‐in‐Chief, Frontiers of BiogeographyReferencesBeckage, B., Gross, L., Platt, W., Godsoe, W. &Simberloff, D. (2012) Individual variation andweak neutrality as determinants of forestdiversity. Frontiers of Biogeography, 4, 145-155.Dawson, M.N (2012) Parallel phylogeographic structurein ecologically similar sympatric sister taxa.Molecular Ecology, 21, 987-1004.Dawson, M.N, Field, R. & Hortal, J. (2011) AdvancingFrontiers, with a retrospective. Frontiers ofBiogeography, 3, 1–2.Fagin, T. & Hoagland, B. (2011) Patterns from the past:Modeling Public Land Survey witness treedistributions with weights-of-evidence. PlantEcology, 212, 207-217.Gibbons, R.E., Barrio, J., Bravo, G.A. & Alza, L. (2011)Assessing the geographic range of Black-frontedGround-Tyrants (Muscisaxicola frontalis) usingextralimital and winter range occurrencerecords and ecological niche modeling. Journalof Field Ornithology, 82, 355-365.Hortal, J. & Dawson, M.N (2009) Frontiers ofBiogeography, a new frontier for the IBS.Frontiers of Biogeography, 1, 1.Ladle, R.J., Jepson, P., Malhado, A.C.M., Jennings, S. &Barua, M. (2011) The causes andbiogeographical significance of species’rediscovery. Frontiers of Biogeography, 3, 111-118.Lomolino, L.R. & Heaney, M.V. (2009) From thefoundations to the frontiers of biogeography.Frontiers of Biogeography, 1, 3–4.Peterson, A. & Lieberman, B. (2012) Species’geographic distributions through time: Playingcatch-up with changing climates. Evolution:Education and Outreach, doi:10.1007/s12052-012-0385-2.Scheiner, S.M. (2011) Musings on the Acropolis:Terminology for biogeography. Frontiers ofBiogeography, 3, 62-70.Smith, T.W. & Lundholm, J.T. (2012) Environmentalgeometry and heterogeneity–diversityrelationships in spatially explicit simulatedcommunities. Journal of Vegetation Science,doi:10.1111/j.1654-1103.2011.01380.x.2 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and update ISSN 1948-6596updateA cure for seeing double? Convergence and unification inbiogeography and ecologyMore perspectives are presented by Biogeographyand ecology: two views of one world (Jenkins andRicklefs 2011a) 1 than are enumerated in the title.This collection of ten papers targets, and revealsmuch about, the “converge[nce of biogeographyand ecology] at intermediate spatial and temporalscales”; yet “lingering differences” hint at thechallenges facing these disciplines during “the currenttrend towards their unification” (Ricklefs andJenkins 2011).Approximately twenty years ago, after decadesof separation, the disciplines of biogeographyand ecology found common ground spanningintermediate spatial and temporal scales, frommetapopulations to regional communities (Jenkinsand Ricklefs 2011b) or perhaps from assemblagesto species’ ranges (Guisan and Rahbek 2011, Hortal2011, Figure 1). Macroecology (Brown 1995)played a pivotal role, providing the raison d’êtreand methodology for assembling details on individualspecies into broad generalizable pictures ofcontinental diversity (Brown and Maurer 1989). Asecology reached out spatially and back in time toexplain, for example, patterns of mammal bodysize on four major continents (Smith and Lyons2011), so phylogenetic analyses began organizingspecies’ traits and then intra-specific ecologicalvariation in an evolutionary framework(Felsenstein 1985, Poulin et al. 2011). So too, thetools for distinguishing species genetically andphenotypically increased, permitting new questionsabout the relationships between taxonomic,phylogenetic, and functional diversity: how do themany forms of biodiversity vary spatially and temporally,how do different aspects of diversity relateacross scales, and what does this tell us aboutcommunity assembly and ecosystem function(Cavender-Bares et al. 2009, Davies and Buckley2011, Emerson et al. 2011, Weiher et al. 2011.Hortal et al. 2012)? The increasing rate of publicationon such issues (Cianciaruso 2011, Jenkins andRicklefs 2011b, Smith and Lyons 2011) suggestsinterdisciplinary understanding of the organizationof biodiversity at intermediate spatial andtemporal scales is a common goal of ecologistsand biogeographers helping to resolve, for example,the meaning and consequences of the niche(Chase and Myers 2011, Wiens 2011) and of diversity(Chiarucci et al. 2011).The image of biogeography and ecologyconjured up by this collection is one of manythreads at the beginnings of a braid. Each paper islightly intertwined with another on one of fourtopics—niche, macroecology and comparativeecology, community assembly, diversity—thenwound loosely around each other within thetheme (Ricklefs and Jenkins 2011). The strands areheld together by only a few connections, encouragedin advance of the symposium: on average, afew cross-citations and acknowledgement of commentson earlier manuscripts. Thus, at present,the natural convergence of biogeography andecology at intermediate spatial and temporalscales looks more like interdigitation than mutualassimilation of ideas and techniques. The challengeis, in part, overcoming the practical limitationsof measuring biodiversity across multiplelevels of organization and spatial scales of interest,and turning the perceptual biases dependenton grain size, spatial extent, and phylogeny intoopportunities for developing scale-free formulationsof biodiversity (Chiarucci et al. 2011) or scalingconcepts that help merge community ecologywith biogeography (Weiher et al. 2011). Overcomingthis challenge may require taking the organizationalmodel applied in this symposium one ortwo steps further, attracting individuals frommany backgrounds to engage as interdisciplinaryteams on common projects.Whether the “trend towards … unification”1 This August 2011 theme issue of Philosophical Transactions of the Royal Society B reports the proceedings of theInternational Biogeography Society symposium “Biogeography and Ecology: Two Lenses in One Telescope” – seehttp://www.biogeography.org/html/Meetings/2011/program.htmlfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society3

news and updateFigure 1. Studies of biogeography and ecology converge atintermediate spatial and temporal scales typical of themetapopulation (mp), metacommunity (mc) and regionalcommunity (rc), following Jenkins and Ricklefs (2011b).Other symbols represent the individual organism (i), population(p), assemblage (a), species’ range/s (sr), and biota(b). Positioning in relation to the space and time axes isintended to represent approximate scale, while acknowledgingthere may be considerable spatial and temporalvariation within and between places, times, and also amongtaxa. Examples of two hypothetical taxa (black text in theforeground and white in the background, representing avery simple system with respect to the taxon axis) are providedto illustrate that ecological and biogeographic processeslikely have spatio-temporal relationships that varywith species’ traits and biophysical interactions that do notscale linearly with organism size (e.g. Dawson & Hamner2008 [figure 6]). Identifying natural boundaries amongthese scales, or the extent to which scales overlap, is a challengingresearch agenda that may be enabled by coordinatedteams of biogeographers and ecologists (Chiarucci etal. 2011) working across scales, using new metrics such asphylobetadiversity (Emerson et al. 2011). These are fundamentalchallenges in how we represent, and therefore howwe think about (and vice versa), the world.of biogeography and ecology (Ricklefs and Jenkins2011) can continue may similarly depend uponthe approach. Recent efforts to develop unifiedtheory are a little younger than macroecology(Hubbell 1997, 2001, Vellend 2005, Rosindell andPhillimore 2011) and more strongly contested(e.g. Roughgarden 2009, Fukami 2010, Clark2012), perhaps in part because of the long historyof thought surrounding idiosyncrasies in biology(Gould 1989, McIntyre 1997, Lawton 1999). Yetthe truism that each extant species has an uniquehistory of lineage and place (Lomolino et al. 2006)should not overwhelm the evidence from manynatural examples of evolutionary convergences(Norris 1991, Van Valkenburgh 2007) and parallels(Elmer and Meyer 2011, Smith and Lyons 2011,Dawson 2012), nor from general theory (e.g. Mac-Arthur and Wilson 1967) and ‘rules’ (e.g. Lomolinoet al. 2006), that suggest natural laws (Ghiselin1994). By broadening the scales across whichsome natural phenomena traditionally have beenstudied (see Rosindell et al. 2011) we may encompassindividualism and unity within a single framework,thus avoiding the extremes of übercontingencyor naturalistic theism (Gould 1989,Morris 2005), and maintaining an healthy tensionthat lends itself to inquisition via the logical treeof strong inference (Platt 1964). The perspectivesprovided by comparative biogeography and macroecology,for example, escape the overwhelminglycomplicated contingency at the intermediatescales studied by community ecology (Lawton1999).Striking a balance between empiricism andtheory also is essential. Sixty-two years ago, AlbertEinstein published his conception for general unifiedfield theory (Einstein 1950), which at the timewas largely ignored but now coalesces mainstreamresearch in fundamental physics. Einsteinbelieved that extrapolation from phenomenologicalstudy depended too much on concepts veryclose to the measured experience (Van Dongen2010:63). In biogeography and ecology, the scalesat which the heterogeneity in interactions betweenorganisms and environment manifest—andestimates of α-, β-, and γ-diversity—are affectedby the grain, focus, and extent by which a particularassemblage is measured (Chiarucci et al. 2011).Because of this epistemological constraint, Einsteinbelieved that physics needed to “apply freespeculation to a much greater extent” (Van Dongen2010:93).4 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and updateEvans, M.R., Norris, K.J. & Benton, T.G. (2012) Predictiveecology: systems approaches. PhilosophicalTransactions of the Royal Society B, 367, 163–169.Felsenstein, J. (1985) Phylogenies and the comparativemethod. American Naturalist, 125, 1–15.Fukami, T. (2010) Why a grand unified theory is neitherfeasible nor desirable. Science, 330, 1049–1050.Ghiselin, M.T. (1997) Metaphysics and the origin ofspecies. State University of New York Press, Albany,New York.Gould, S.J. (1989) Wonderful life: the Burgess Shale andthe nature of history. W. W. Norton, New York,NY.Guisan, A. & Rahbek, C. (2011) SESAM – a new frameworkintegrating macroecological and speciesdistribution models for predicting spatiotemporalpatterns of species assemblages. Journalof Biogeography, 38, 1433–1444.Hortal, J. (2011) A modelling framework to open thegates of assemblage structure. Frontiers of Biogeography,3, 43.Hortal, J., De Marco Jr, P., Santos, A.M.C. & Diniz-Filho,J.A.F. (2012) Integrating biogeographical processesand local community assembly. Journal ofBiogeography, 39, 627–628.Hubbell, S.P. (1997) A unified theory of biogeographyand relative species abundance and its applicationto tropical rain forests and coral reefs. CoralReefs, 16, S9–S21.Hubbell, S.P. (2001) The unified neutral theory of biodiversityand biogeography. Monographs in PopulationBiology, 32. Princeton University Press,Princeton, NJ.Jenkins, D.G. & Ricklefs, R.E. (eds) (2011a) Theme issue:‘Biogeography and ecology: two views of oneworld’. Philosophical Transactions of the RoyalSociety B, 366, 2331–2448.Jenkins, D.G. & Ricklefs, R.E. (2011b) Biogeography andecology: two views of one world. PhilosophicalTransactions of the Royal Society B, 366, 2331–2335.Langerhans, R.B. & DeWitt, T.J. (2004) Shared andunique features of evolutionary diversification.American Naturalist, 164, 335–349.Lawton, J.H. (1999) Are there general laws in ecology?Oikos, 84, 177–192.Levin, S.A. (2010) The evolution of ecology. Chronicle ofHigher Education, 8 th August. http://chronicle.com/article/The -Evolution -of-Ecology/123762/ Accessed 21 March 2012.Lomolino, M.V., Sax, D.F., Riddle, B.R. & Brown, J.H.(2006) The island rule and a research agenda forstudying ecogeographical patterns. Journal ofBiogeography, 33, 1503–1510.MacArthur, R.H. & Wilson, E.O. (1967) The theory ofisland biogeography. Princeton University Press,Princeton, NJ.McIntyre, L. (1997) Gould on laws in biological science.Biology and Philosophy, 12, 357–367.Morris, S.C. (2005). Life's solution: inevitable humans ina lonely universe. Cambridge University Press,Cambridge, UK.Norris, R.D. (1991) Biased extinction and evolutionarytrends. Paleobiology, 17, 388–399.Platt, J.R. (1964) Strong inference. Science, 146, 347–353.Poulin, R., Krasnov, B.R., Mouillot, D. & Thieltges, D.W.(2011) The comparative ecology and biogeographyof parasites. Philosophical Transactions ofthe Royal Society B, 366, 2379–2390.Ricklefs, R.E. & Jenkins, D.G. (2011) Biogeography andecology: towards the integration of two disciplines.Philosophical Transactions of the RoyalSociety B, 366, 2438–2448.Rosindell, J. & Phillimore, A.B. (2011) A unified modelof island biogeography sheds light on the zoneof radiation. Ecology Letters, 14, 552–560.Rosindell, J., Hubbell, S.P. & Etienne, R.S. (2011) TheUnified Neutral Theory of Biodiversity and Biogeographyat age ten. Trends in Ecology andEvolution, 26, 340–348.Roughgarden, J. (2009) Is there a general theory ofcommunity ecology? Biology and Philosophy,24, 521–529.Smith, F.A. & Lyons, S.K. (2011) How big should a mammalbe? A macroecological look at mammalianbody size over space and time. PhilosophicalTransactions of the Royal Society B, 366, 2364–2378.Van Dongen, J. (2010) Einstein’s unification. CambridgeUniversity Press, Cambridge, UK.Van Valkenburgh, B. (2007) Déjà vu: the evolution offeeding morphologies in the Carnivora. Integrativeand Comparative Biology, 47, 147–163.Vellend, M. (2005) Species diversity and genetic diversity:parallel processes and correlated patterns.American Naturalist, 166, 199–215.Weiher, E., Freund, D., Bunton, T., Stefanski, A., Lee, T.& Bentivenga, S. (2011) Advances, challengesand a developing synthesis of ecological communityassembly theory. Philosophical Transactionsof the Royal Society B, 366, 2403–2413.Wiens, J.J. (2011) The niche, biogeography and speciesinteractions. Philosophical Transactions of theRoyal Society B, 366, 2336–2350.Edited by Richard Field6 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

ISSN 1948-6596news and updateworkshop summaryThe application of species distribution models in the megadiverseNeotropics poses a renewed set of research questionsSpecies distribution models: applications, challenges and perspectives – Belo Horizonte, Brazil,29 th –30 th August 2011The community of researchers and techniciansinterested in biogeography is large and growing inBrazil, with members coming from fields as diverseas ecology, evolution and conservation. Theincrement in the number of postgraduate programsin ecology and evolutionary biology islinked to many research questions about thecauses and dynamics of species’ ranges, as well asabout their consequences for short-term evolutionaryprocesses. As a result, the use (and abuse)of species distribution models (SDMs) as a tool inresearch and technical studies has grown rapidlyin recent years. Systems that integrate biodiversitydatabases (e.g. SpeciesLink 1 ) allow one to obtaindistributional information about many species;and easy-to-use SDM software (e.g. MaxEnt 2 ,Phillips et al. 2004, or openModeller 3 , Muñoz etal. 2011) is also available online. The easy availabilityof data and SDM tools provides a powerfulmeans for answering questions about the geographicdistribution of species. Potential usersinclude non-specialist researchers, untrained postgraduatestudents and government technicians.But is this helping to develop a sound and solidlygrounded knowledge of the distribution ofNeotropical diversity?In order to unite the Brazilian community ofSDM users and provide them with a better understandingof the technique, the PostgraduateCourse on Plant Biology of the Universidade Federalde Minas Gerais organized a workshop in BeloHorizonte last August. This workshop allowedmore than 100 students and technicians to meetand discuss with researchers working with SDMs.The main conclusions of the meeting were thatthere is a growing interest in using the techniqueto study species’ distributions and find unknownpopulations of rare species, and that there is aneed for a code of good practice in both field surveysand SDM applications. These conclusionshave been already discussed elsewhere (Kaminoet al. 2011). Here we develop further one of theproblems identified during the workshop: the lackof clear questions in many studies using SDMs; inother words, the mere application of species distributionmodelling as a fashionable techniqueoften ‘justifies’ a study.Reviews summarizing the most importantchallenges for SDMs (e.g. Araújo and Guisan 2006,Zimmermann et al. 2010, Peterson et al. 2011)typically present how users view the field andwhat problems they perceive in each step in themodelling process. However, things move fast inthis emerging field of research because, while thenumber of studies using SDMs increases steadily(see Fig. 1 in Lobo et al. 2010), the technique isalso used to address completely new questions.Thus, the challenges to their application are themselveschanging. We believe that the most importantof these challenges are theoretically ratherthan methodologically grounded, although werecognize that both kinds of problems overlap tosome extent. Here we outline the basic questionsthat we believe SDM users must take into accountwhile studying current species’ distributions.Soberón (2007) provides perhaps the beststarting point to understand the theoretical problemsof SDMs (see also Colwell and Rangel 2009;Soberón 2010). The first problem is the definitionof a clear research question. Here it is importantto discriminate between theoretical questionssuch as “Why is this species here?” – which aremore interesting in the long run – from those thatare eminently practical such as “Where is this species?”– which unfortunately seem to be the mostcommon. But even practical questions stimulated1 http://www.splink.org.br; last accessed 10/11/20112 http://www.cs.princeton.edu/~schapire/maxent/; last accessed 10/11/20113 http://openmodeller.sourceforge.net/; last accessed 10/11/2011frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society7

news and updateby, for instance, the design of conservation programsneed to be properly defined before SDMsare used, because they provide the basis for makingadequate methodological choices.Different questions may deserve differentmethods. One of the recurrent dilemmas for usersis choosing the most appropriate modelling technique.However, when the research question iswell defined, choosing one particular strategycomes more easily. The organization of SDM techniquesin categories of complexity 4 , together withacknowledging the advantages and drawbacks ofthe techniques in each category, may help tochoose the best modelling strategy for each researchquestion (Kamino et al. 2011). Since thereare different products, they must have differentuses. While simple models may help to find sitesthat are climatically similar to the known occurrencesof the species, or be suited to describing itsrealized niche, more complex models may be bestfor describing the actual distribution of the species(see Jiménez-Valverde et al. 2008).The definition of an adequate modellingstrategy also raises several issues. Surprisingly, thefirst is that there may not be a minimum numberof occurrences to be used in SDMs (other thanthey should be higher than zero). What is necessary,however, is that the question tackled isclearly understood and the relationship betweenthe number of occurrences and the SDM techniqueused is known. Datasets with very few occurrencesin combination with simple SDM methodsmay be useful for providing a description ofthe areas climatically similar to the recorded presences,allowing the planning of new field surveys(Siqueira et al. 2009) or a preliminary gap analysisto identify areas for conservation (Nóbrega andDe Marco Jr 2011). However, larger datasets areneeded for more complex methods and/or answeringmore sophisticated questions about thedistribution of the species.Determining how many predictors shouldbe used to develop the models and how manyparameters should be allowed while modellingdeserves special attention. Many complex SDMtechniques suffer from chronic overfitting – a diseasethat is easy to catch but hard to cure. Overfittedmodels may restrict too much the predicteddistributions to the geographic and environmentaldomain of the observed occurrences, impedingidentification of areas where the species could bepotentially present, or the reliable representationof the realized response of the species to the environment(Lobo 2008).Another theoretical problem of SDMs isthat the current distributions of the species areoften not in equilibrium with the environment(Araújo and Pearson 2005, Soberón 2007, 2010,Jiménez-Valverde et al. 2008). This may resultfrom barriers to dispersal, but also from climatechange, even in the absence of such barriers, ifthe dispersal ability of the species is not enough totrack the changes in its potential geographic distribution.Similarly, recently originated species areexpected to have restricted ranges and not be inequilibrium with the environment, if they lackedthe time to disperse to their whole potential distribution.Such lack of equilibrium poses two keyquestions: how to restrict model predictions tothe truly occupied areas (if that is the intention)and how to incorporate dispersal abilities intoSDM strategies. The former can be tackled eitherby defining appropriate pseudo-absences to includein the calibration dataset (Lobo et al. 2010)or by restricting the extent of the study to thearea that could have been colonized by the species(Anderson and Raza 2010). Incorporating dispersalabilities may be more difficult, but the incorporationof cellular automata or other spatiallyexplicit methods (e.g. Brook et al. 2009) will generatea new generation of SDMs that may allowstudy of the spatiotemporal dynamics of species’ranges.Finally, contrary to the general opinion ofmany users, the ability of species distributionmodels to measure the fundamental niche of aspecies is severely limited. First and foremost, thelack of equilibrium of the species’ range with the4 From simple (DOMAIN, BIOCLIM, Euclidean Environmental Distance) to moderate (Mahalanobis distance, GLM,GAM) and complex models (MaxEnt, Artificial Neural Networks, GARP) (T.F. Rangel unpublished, see also Jiménez-Valverde et al. 2008).8 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and updateenvironment may interfere with any attempt toestimate its potential distribution (i.e., the fundamentalniche in relation to climate). Gaining insightsabout the fundamental niche of a speciesbased on environmental information and somerecorded occurrences relies on prior knowledge ofits requirements and having an adequate theoreticalmodel for the niche itself. Additionally, manyresearchers tend to use variables only from widelyavailable datasets (such as WorldClim 5 ), instead ofseeking other predictors that may be less easy –but yet possible – to obtain (Kamino et al. 2011, S.Amaral unpublished). We would like to emphasizethat variables representing edaphic conditions areoften not available, yet these could be importantpredictors for some species and are rarely consideredin SDMs. These problems, in combinationwith the collinearity that is intrinsic to most spatiallyexplicit variables, may be impeding the identificationof predictors that are truly determiningthe geographic distribution of many species. Itfollows that it would be difficult to evaluate therelative importance of different environmentalpredictors in the absence of a good theoreticalmodel and some knowledge on the physiologyand ecology of the studied species. The interactionwith students and technicians during theworkshop showed us that a good understandingof the framework proposed by Soberón (2007,2010) helps in clarifying the concepts and theirmeanings in this respect. But further work and alot of didactics are needed to actually imprint thenext generation of SDM users with a solidly rootedknowledge about the technique they use, the implicationsof the methodological choices theymake and the power that these models actuallyhave to solve theoretical and applied questions inecology, evolution, biogeography and conservation.AcknowledgementsWe thank all attendees of the workshop for thediscussions. The workshop was organized by LHYKand JRS and supported by UFMG (Pró-Reitoria dePós-Graduação, Instituto Ciências Biológicas, Pós-Graduação em Biologia Vegetal, Genética, Zoologia,and Ecologia, Conservação e Manejo de VidaSilvestre), Conselho Regional de Biologia and Viamundi.LHYK was financed by CAPES Grant, PDMJrand JRS by continuous CNPq productivity grantsand JH by a CNPq Visiting Researcher Grant.Luciana H. Y. Kamino 1 , Paulo De Marco Jr 2 ,Thiago F. Rangel 2 , Silvana Amaral 3 ,Marinez F. de Siqueira 4 , Renato DeGiovanni 5 , João Renato Stehmann 1 andJoaquín Hortal 2,61 Depto. Botânica, Instituto Ciências Biológicas, UniversidadeFederal de Minas Gerais, Belo Horizonte, Brazil.lucianakamino@gmail.com; http://www.icb.ufmg.br/pgbot/2 Depto. Ecologia, Instituto de Ciências Biológicas, UniversidadeFederal de Goiás, Goiânia, Brazil3 Divisão de Processamento de Imagens, Instituto Nacionalde Pesquisas Espaciais, São José dos Campos, Brazil4 Unidade de Botânica Sistemática, Instituto de PesquisaJardim Botânico do Rio de Janeiro, Rio de Janeiro, Brazil5 Centro de Referência em Informação Ambiental, Campinas,Brazil6 Depto. Biodiversidad y Biología Evolutiva, Museo Nacionalde Ciencias Naturales (CSIC), Madrid, SpainReferencesAnderson, R.P. & Raza, A. (2010) The effect of the extentof the study region on GIS models of speciesgeographic distributions and estimates ofniche evolution: preliminary tests with montanerodents (genus Nephelomys) in Venezuela. Journalof Biogeography, 37, 1378–1393.Araújo, M.B. & Guisan, A. (2006) Five (or so) challengesfor species distribution modelling. Journal ofBiogeography, 33, 1677–1688.Araújo, M.B. & Pearson, R.G. (2005) Equilibrium of species'distributions with climate. Ecography, 28,693–695.Brook, B.W., Akçakaya, H.R., Keith, D.A., Mace, G.M.,Pearson, R.G. & Araújo, M.B. (2009) Integratingbioclimate with population models to improveforecasts of species extinctions under climatechange. Biology Letters, 5, 723–725.Colwell, R.K. & Rangel, T.F. (2009) Hutchinson's duality:The once and future niche. Proceedings of theNational Academy of Sciences USA, 106, 19651–19658.5 http://www.worldclim.org/; last accessed 10/11/2011frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society9

news and updateJiménez-Valverde, A., Lobo, J.M. & Hortal, J. (2008) Notas good as they seem: the importance of conceptsin species distribution modelling. Diversityand Distributions, 14, 885–890.Kamino, L.H.Y., Stehmann, J.R., Amaral, S., De Marco Jr,P., Rangel, T.F., de Siqueira, M.F., De Giovanni,R. & Hortal, J. (2011) Challenges and perspectivesfor species distribution modelling in theNeotropics. Biology Letters, in press.doi:10.1098/rsbl.2011.0942Lobo, J.M. (2008) More complex distribution models ormore representative data? Biodiversity Informatics,5, 14–19.Lobo, J.M., Jiménez-Valverde, A. & Hortal, J. (2010) Theuncertain nature of absences and their importancein species distribution modelling. Ecography,33, 103–114.Muñoz, M.E.S., Giovanni, R., Siqueira, M.F., Sutton, T.,Brewer, P., Pereira, R.S., Canhos, D.A.L. and Canhos,V.P. (2011) openModeller: a generic approachto species’ potential distribution modelling.Geoinformatica, 15, 111–135.Nóbrega, C.C. & De Marco Jr, P. (2011) Unprotectingthe rare species: a niche-based gap analysis forodonates in a core Cerrado area. Diversity andDistributions, 17, 491–505.Peterson, A.T., Soberón, J., Pearson, R.G., Anderson,R.P., Martínez-Meyer, E., Nakamura, M. &Araújo, M.B. (2011) Ecological niches and geographicdistributions. Princeton University Press,Princeton.Phillips, S.J., Dudík, M. and Schapire, R.E. (2004) Amaximum entropy approach to species distributionmodeling. In: Proceedings of the 21st InternationalConference on Machine Learning, ACM-Press, New York, pp. 655–662Siqueira, M. F., Durigan, G., De Marco Jr, P. and Peterson,A. T. (2009) Something from nothing: Usinglandscape similarity and ecological niche modelingto find rare plant species. Journal for NatureConservation, 17, 25–32.Soberón, J. (2007) Grinnellian and Eltonian niches andgeographic distributions of species. Ecology Letters,10, 1115–1123.Soberón, J. (2010) Niche and area of distribution modeling:A population ecology perspective. Ecography,33, 159–167.Zimmermann, N.E., Edwards, T.C., Graham, C.H., Pearman,P.B. & Svenning, J.-C. (2010) New trends inspecies distribution modelling. Ecography, 33,985–989.Edited by Richard FieldISSN 1948-6596Remember that being a member of IBS means you can get free online access to four biogeographyjournals: Journal of Biogeography, Ecography, Global Ecology and Biogeography andDiversity and Distributions. You can also obtain a 20% discount on the journals Oikos and Journalof Avian Biology.Additional information is available at http://www.biogeography.org/.10 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and updatesymposium summaryTowards new directions and collaborations in macroecology6 th Annual Meeting of the Specialist Group for Macroecology of the Ecological Society of Germany,Austria and Switzerland (GfÖ) – Frankfurt am Main, Germany, 29 th February–3 rd May 2012This year, Katrin Böhning-Gaese’s group(Biodiversity and Climate Research Center, BiK-F),with support from the Senckenberg Gesellschaftfür Naturforschung and the Ecological Society ofGermany, Austria and Switzerland (GfÖ) hostedthe annual meeting of the Macroecology SpecialistGroup. Over 100 people attended the conferenceand participated in a full schedule of eventsincluding stimulating talks and posters (6keynotes, 24 contributed talks, 18 posters), alively panel discussions on publication in ecologyand reflections of the keynote speakers on thenovelty of the conference and future researchdirections in macroecology 1 .The conference opened with a warmwelcome from Katrin Böhning-Gaese who set thestage for the meeting with an inspiring overviewof macroecology research at the BiK-F. Thispresentation was followed by Carsten Rahbekwho challenged the audience to “think big” andtake on long-unresolved challenges inmacroecology by embracing new tools (i.e.,genomics, ancient DNA), gathering more empiricaldata and extending collaboration among researchlabs and different types of scientists. (Macro)ecologists should not be too modest in composinglarge research projects!The major themes of the conferenceincluded: macroecological patterns and theirunderlying causes; niches, distributions,communities and phylogenies under globalchange; advances in modelling, which includedboth theoretical and statistical approaches aimedat including more biological realism in models;and extinctions, conservation and new frontiers.Macroecological topics ranged from considerationof ecologically mediated diversity limits whenevaluating diversification rates (Yael Kisel) toquantification of spatial and environmental effectsof beta diversity in China’s woody plants (ZhihengWang), and evaluation of the biogeographicpatterns/hypotheses of thermal melanism inEuropean dragonflies (Dirk Zeuss).Presentations and posters under thethemes of niches, distribution, communities andphylogenies under global change extendedcurrent state-of-the-art attempts to integratethese multiple types of data to address bigquestions in macroecology and biogeography. Forinstance, Sébastien Lavergne evaluated whetherpast rates of niche evolution influenced currentdemographic trends in European birds; RafaelWüest explored how species pool definitioninfluences inference about mechanisms (i.e.,environmental filtering or biotic interactions) thatstructure assemblage composition; Dieter ThomasTietze presented a poster evaluating differentmechanisms causing variation in diversitygradients in Himalayan birds; and Sarah Whitmeeteased apart phylogenetic relatedness andgeographic location to evaluate patterns of rangefilling in mammals.Advances in modelling included elegantexamples of new process-based Bayesian models(Florian Hartig), models integrating statistical andmechanistic models (Oliver Schweiger), combiningmultiple interacting species, speciation anddemography (Juliano Sarmento Cabral and MiguelB. Araújo) as well as macroecological simulationstudies that explored what patterns emerge whenspecific ecological and evolutionary processes areconsidered (David Orme). Finally, broad-scaleprocess-based dynamic global vegetation models(DGVMs) were presented by Thomas Hickler whodrew comparisons between these ecophysiologicaland macroecological models and calledfor further integration of the two approaches.The last overarching topic (extinctions,1 See program abstracts for complete details: http://www.bik-f.de/files/veranstaltungen/gfoe_macroecology/homepage_program_full.pdffrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society11

news and updateconservation and new frontiers) included talksabout the loss of cryptic genetic diversity withclimate change (Steffen Pauls), past populationtrends (and extinctions) estimated from ancientgenetics (David Nogués-Bravo) and a call for amacroecological approach to study ecotoxicology(Mikhail Beketov).Beside the scientific topics the hostsorganized a rather unusual but therefore highlywelcomed panel discussion about currentproblems within the publication circus. Thepodium was represented by editors-in-chief,associate editors and editors from a variety ofjournals. Topics such as how to acknowledge thelabour-intensive reviewer and editor work(“reviewer crisis”), the future of (non-) openaccesspublishing, methods to acknowledgeauthor contributions on multi-authored papers(an issue that has become more common withincreased collaboration), and H-factors and impactfactors (and other such metrics) were heatedlydiscussed (discussion continued afterwards at thenicely situated conference dinner). It was evidentthat the issues raised are alarming and need alarger platform for discussion and solutionsamong researchers as well as publishers.The major themes that emerged from thetalks and the discussions included the need formore interdisciplinary research and the realizationthat the macroecological approach can be usefullycombined with a growing number of disciplinesand types of data from a variety of spatial andtemporal scales. This conclusion was partlyprompted by presentations and discussions offields that have had limited interaction withmacroecology including: ecophysiologicalmodelling of past vegetation, ecotoxicology,phenotypic plasticity and microevolution. All feltthat exciting new frontiers lay at the intersectionof disparate disciplines and new collaborationsamong these disciplines should be fostered.Additionally, the sophistication of new types ofmodels to address questions in macroecology wasinspiring and prompted a loud and persistent callfor more empirical data. Such data are not onlynecessary to parameterize models but arerequired to address unanswered questions inmacroecology and biogeography. Further, itbecame clear that more collaboration amongthose gathering data and those using them issorely needed. Finally, there was a call to stayrelevant and use our science to address the ongoingbiodiversity crisis.Catherine Graham 1 and Marten Winter 21 Department of Ecology and Evolution, Stony Brook University,New York, USA. catherine.graham@stonybrook.edu;http://life.bio.sunysb.edu/ee/grahamlab/2UFZ - Helmholtz Centre for Environmental ResearchGmbH, Halle (Saale), Germany. marten.winter@ufz.de;http://www.ufz.de/index.php?en=7081Edited by Jan BeckISSN 1948-6596Your participation in frontiers of biogeography is encouraged. Please send us your articles, commentsand/or reviews, as well as pictures, drawings and/or cartoons. We are also open to suggestionson content and/or structure.Please check http://www.biogeography.org/html/fb.html for more information, or contact us atibs@mncn.csic.es and frontiersofbiogeography@gmail.com.12 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and updatebook reviewAvian history is written by the winnersAvian survivors: The history and biogeography of Palearctic birds, by Clive Finlayson, 2011, T & ADPoyser Ltd., 304 pp. £50 (Hardback) ISBN: 9780713688658; http://www.acblack.com/naturalhistory/This is a difficult book to review fairly, because it isnot a book to read through, nor is its intentioneasily discerned. It is a profound and deeplydetailed compilation, assessing the climatic,historical and biogeographical attributes of thebird fauna of the Western Palaearctic. Itsintention is to understand how this particularconstellation of species survived through theclimatic vicissitudes of the Tertiary Period, andespecially the last 2 million years of thePleistocene climatic fluctuations, to be combinedas the extant fauna.Finlayson was inspired by the remarkablydiverse faunas that appear in the caves ofGibraltar, at the extreme southern tip of theWestern Palaearctic. At different times theseincluded northern species such as Great Auk andVelvet Scoter, at others southern species such asLammergeier and Crag Martin. Those that stilloccur in the Palaearctic are the survivors of histitle. He starts with a short outline of the climatic,geographical and ecological changes that affectedthe region during the Tertiary, as Africa and Indiashifted north to collide with Europe and Asia, soclosing the Mediterranean, and causing the upliftof the mountain ranges across central Europe andAsia that changed the climate, especially thestrength and direction of rain-bearing winds.Grasses evolved in this period, and grasslands,replacing tropical forests, became morewidespread. His analysis of the times of origin ofbird genera that, at present, include migratoryspecies, especially Trans-Saharan migrants,suggests that many of them evolved in theMiocene, as the grasslands appeared. Popular,perhaps superficial, opinion has assumed that theglacial-interglacial cycles of the Pleistoceneprompted the evolution of the migratory habit,but this analysis strongly suggests that it wasalready well established.The majority of the book is a detailedexamination of each family, genus and speciesthat is currently represented in the Palaearctic. Itassesses their climatic limits, habitat requirementsand likely history. Much modern work on geneticphylogeny is used to assess the zoogeographicorigins of such taxa, when they migrated into thePalaearctic, and how their climatic requirementshave persisted, along with the taxa themselves.Each is assigned to its climatic and latitudinaltolerance ranges, and to its migratory group(sedentary, partial migrant, Trans-Saharanmigrant). For instance, Finlayson interprets thehistory of the Corvidae as probably evolving inAustralia, and migrating into the tropical forests ofSoutheast Asia, where a great diversity of basalgenera still occur; but one, the Choughs(Pyrrhocorax) migrated out along the greatmountain chain right across Asia and Europe inthe Miocene, as mountain uplift created theirdrier upland habitat. However, most corvidsremained essentially forest species, and migratedwest along the forest belts south (mostlydeciduous) or north (mostly coniferous) of themountains. Corvids reside mostly in more humidclimatic zones, and few show much migratorybehaviour. Fossil corvids are represented inEurope by Middle Miocene times; two moderngenera are present by the Early Pliocene(Pyrrhocorax, Pica, two more by Middle Pliocene(Garrulus, Nucifraga) and all by Early Pleistocene.Sometimes the detailed evidence on whichsuch histories are based is clearly indicated. Thefossil record is largely quoting the compilation byMlíkovský (2002), and the phylogenetic referencesare well cited. However, much of the story isinformed speculation, of probable histories andprobable spreads, based upon known climatic andgeographical changes, and the known broadecological requirements of the taxa. Given thenumber of taxa involved, this amounts to quite alot of speculation accumulated over the book as awhole. This is not to suggest that it is wrong, nordoes it deny its value. A lot of diverse informationfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society13

news and update ISSN 1948-6596has been interwoven to compile these stories. Thevalue of the book lies in the last three chapters,which analyse the current Palaearctic fauna as awhole in terms of its zoogeographic origins andhistory, climatic tolerances and migratorytendencies. This provides some fascinatinginsights. For instance, while much of thePalaearctic fauna originated in Southeast Asia, asubstantial proportion came from North America(and some Palaearctic lineages reciprocated), butvery few came from neighbouring sub-SaharanAfrica. The tables listing the species and groupsbelonging to each faunal component arefascinating. How few are the lineages that haveworldwide ranges, probably reflecting their originin the warm, humid, Eocene World (Table 19.1lists only 33 genera). Similarly, the dominance ofthe Palaearctic bird fauna by omnivorous speciescontrasts with the few specialist herbivores andlarge-prey predators; Finlayson makes theinteresting suggestion that the latter niches mightbe dominated instead by mammals. Manyomnivorous species show wide climatictolerances, and are often sedentary, while theinsectivores, another diverse group, manage toremain so because they are mostly migratory;many of them show restricted bioclimatictolerances. These alternative major strategies,migration or bioclimatic tolerance, are whatFinlayson divines as the major reasons for thecurrent diversity of the avifauna.This is not easy reading for the averageornithologist, but is a stimulating compilation forzoologists wanting an evolutionary perspective onthe fauna. They will find much to think about, andsome details to contest. Finlayson blithely acceptsthe phylogenetic argument that many bird lineagesoriginated in the late Cretaceous; this ignores thepaucity of palaeontological support, and I suspectinstead that the molecular clock ran fast for birds inthe early Palaeocene, as it surely did also formammals. In practice, this assumption makes littledifference to his overall analysis, and does notdetract from its value. As well as his inspiration fromthe ancient fauna of Gibraltar, he acknowledges theimportance of Moreau’s influential and magisterialanalysis, now 40 years old, in The Palaearctic-AfricanMigration Systems (1972). Finlayson has indeedcompiled a worthy sequel.ReferencesMlíkovský, J. (2002). Cenozoic Birds of the World. NinoxPress, Prague.Moreau, R. E. (1972). The Palaearctic-African MigrationSystems. Academic Press, London.Derek W. YaldenHonorary Reader in Zoology, School of Life Sciences,University of Manchester.derek.w.yalden@manchester.ac.ukEdited by Markus EichhornYou can find information about the International Biogeography Society at http://www.biogeography.org/, and contact with other biogeographers at the IBS blog (http://biogeography.blogspot.com/), the IBS facebook group (http://www.facebook.com/group.php?gid=6908354463) and the IBS twitter channel (https://twitter.com/biogeography).14 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

ook reviewEverything changes – especially on islandsnews and updateIsland Environments in a Changing World, by Lawrence R. Walker & Peter Bellingham, 2011,Cambridge University Press, 324 pp. £70 (Hardback) / £30 (Paperback) ISBN: 9780521519601 /9780521732475; http://www.cambridge.org/‘Fog in Channel – continent cut off’Are we not all islanders, after all, holding our ownoff-continent perspective of the world? Whetherit is the Earth in the cosmos or Africa between theoceans, or even a tiny islet where a lonelyshepherd attends his goats, humans areeverywhere surrounded by non-inhabitableterritories. It is thus not surprising that islandsexert a special allure to us all. At the same time,typical islands, i.e. relatively small bodies of landsurrounded by water, provide a huge set ofnatural laboratories for the study of numerousbiological and physical phenomena. This is mostlybecause they are relatively small, simple and welldelimitedopen systems. The story of MacArthurand Wilson’s (1967) clever choice of islands asmodel systems for their theory is definiteevidence in support of this.After an early start through Wallace’sseminal book, Island Life (Wallace 1881), theecological literature on islands grew tremendouslybut has been mainly of a rather technical flavour,focusing on island biogeography, evolutionarydynamics and invasions of alien species (see Lososand Ricklefs 2010). A recent attempt of almostmonumental dimensions, the Encyclopedia ofIslands (Gillespie and Clague 2009), tried to bringthe ecological view of islands closer to the layperson, but its very size and breadth of scope mayalso be its weak point. At the same time, there areseveral other books accessible to the public thatexamine island life from an evolutionaryperspective or at a local geographical scale (e.g.,Grant 1997, Quammen 1997, Samways et al.2010). The special properties of islands, i.e.finiteness and small size, act as timelineintensifiers, enabling a perception of theconstantly changing environment which may bemore difficult to grasp in the vast expanses ofcontinental areas.This Heraclitean view of nature is the maintheme of this recent book on island environmentsby two plant ecologists, L.R. Walker (Univ. ofNevada, USA) and P. Bellingham (LandcareResearch, New Zealand). These authors present aconcise account of almost all the importantaspects of insular environments, using simplelanguage without loss of scientific rigour. Nearly150 figures including diagrams and photos(unfortunately most in black and white, whilethose in colour plates could have been printed inbetter quality) are inserted in the text, assistingthe reader in capturing concepts, processes andvisualising examples. The scope of the book mighthave merited a larger volume, but the authorsaimed for a much shorter text, adopting a cleverstrategy that helps them keep the length withinthe readable limit of around 300 pages. Out of thethousands of insular systems around the globe,they select nine island groups that exhibit a goodsample of crucial insular features. These groupsexemplify all three categories of islands identifiedby biogeographers, namely oceanic, continentalfragments and continental shelf islands, as well asa variety of geographical zones, from tropical totemperate, even subarctic. This way, the authorsare able to examine a vast array of insularfeatures with concrete examples from each islandgroup. They also use the nine island groups toprovide a vivid account of major causes of ‘naturaldisturbances’, such as volcanic activity,earthquakes, erosion, fire, floods and so forth,affecting different island systems at varying rates.Processes affecting the biota of islands, mostimportantly dispersal and extinction, are offered achapter of their own, with a short discussion ongeneral richness and endemicity patterns. Hereagain, selected examples are inserted in textboxes giving a solid form to abstract concepts.Almost half of the book explores the roleplayed by humans in changing the environment ofislands, placed in a historical framework, from firsthuman colonisation, through gradually intensifiedeffects, up to the present day. The complexity ofhuman interactions with insular environments isrevealed through a balanced examination of theoften contradictory effects of activities liketourism and agriculture. Sustainability comes upagain as the holy grail of management andfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society15

news and update ISSN 1948-6596conservation planning, and the authors are clearcutin their final chapter about the future: ‘thefate of island ecosystems rests in the hands ofhumans’ (p.305).This is definitely a book that can reach thelay reader, decision-makers and administrativebodies, offering a clear view of the majorproblems that need to be amended without delay.The professional ecologist and the islandbiogeographer also have much to gain from thiscompilation of data and examples, as well as fromthe firmly dynamic view of insular environments.There are a few points that may raise someeyebrows though. The lack of citations in the textis one, although a list of selected reading isoffered at the end of each chapter. The bias infavour of a historical time scale is another, but thiscan be ascribed to the need for prioritising theaccount of intensive human effects on islands. Amore contentious point might be found in theselection of examples, which are not quiterepresentative of the actual diversity andheterogeneity found within island types.Continental shelf islands, in particular, with alltheir variety in physical and historicalcharacteristics, are exemplified by a single group,the British Isles, hardly the most typical exampleof its kind. This under-representation ofcontinental shelf islands leads to a slightlydistorted account of historical effects on islandenvironments. Factors such as agriculture, grazingand fire have played quite variable roles oncontinental shelf islands of different geographicalregions. A rich and relatively well-studied exampleis offered by the more than 10,000 Mediterraneanislands, with a strong interest for bothconservation and theoretical studies (Blondel etal. 2010). Furthermore, due to their recentisolation, biotic components of continental shelfislands are under the strong influence ofoversaturation and relaxation processes, or source–sink dynamics, which may blur our perception ofenvironmental change. The importance of oceanicislands, and to some degree also continentalfragments, in understanding island environmentsand in the construction of general models(Whittaker et al. 2008) is beyond dispute, but thetime has probably come for similar work oncontinental shelf islands too. The dynamics ofconstant change may be more complex there,rendering their study a tougher challenge.These critical remarks do not intend todiminish the contribution of Walker andBellingham to island ecology and biogeography.The authors have accomplished a greatachievement, explaining such a broad range ofconcepts, processes, patterns and problems in aneasy-to-follow text, keeping track of recentdevelopments in the fields of island biogeographyand conservation biology, in addition to global andlocal economics, history and culture. Their workwill become standard suggested reading forstudents of islands in all respective fields. Bookslike this are valuable triggers for further work inour struggle to understand and wisely manage ournatural environment, and can also help usestablish the often elusive paradigm of nature incontinuous flow. Islands will be our guide formany years to come.ReferencesBlondel, J., Aronson, J., Bodiou, J.-Y. & Boeuf, G. (2010)The Mediterranean Region: Biological Diversity inSpace and Time. Oxford University Press, Oxford.Gillespie, R.G. & Clague, D.A. (eds) (2009) Encyclopediaof Islands. University of California Press, Berkeleyand Los Angeles, CA.Grant, P. (ed.) (2007) Evolution on Islands. OxfordUniversity Press, Oxford.Losos, J.B. & Ricklefs, R.E. (eds) (2010) The Theory ofIsland Biogeography Revisited. PrincetonUniversity Press, Princeton, NJ.MacArthur, R.H. & Wilson, E.O. (1967) The Theory ofIsland Biogeography. Princeton University Press,Princeton, NJ.Quammen, P. (1997) The Song of the Dodo: IslandBiogeography in an Age of Extinctions. Scribner,New York, NY.Samways, M.J., Hitchins, P., Bourquin O. & Henwood, J.(2010) Tropical Island Recovery: Cousine Island,Seychelles. Wiley-Blackwell, Chichester.Wallace, A.R. (1881) Island Life. Harper & Bros, NewYork, NY.Whittaker, R.J. & Fernandez-Palacios, J.M. (2007) IslandBiogeography: Ecology, Evolution, andConservation. Oxford University Press, Oxford.Whittaker, R.J., Triantis, K.A. & Ladle, R.J. (2008) Ageneral dynamic theory of oceanic islandbiogeography. Journal of Biogeography, 35, 977–994.Spyros SfenthourakisDepartment of Biological Sciences, University of Cyprus.sfendour@ucy.ac.cyEdited by Markus Eichhorn16 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

ook reviewHeadwaters in the cloudsnews and updateTropical Montane Cloud Forests, by L. A. Bruijnzeel, F. N. Scatena & L. S. Hamilton (editors),2011, Cambridge University Press, 768 pp. £65 (Hardback) ISBN: 9780521760355; http://www.cambridge.org/Tropical Montane Cloud Forests (TMCFs) aregaining in scientific popularity since a firstinternational symposium on this ecosystem washeld in Puerto Rico in 1993. The promotion of themeeting by the UNESCO InternationalHydrological Programme illustrates a far-reachingeffect of cloud forests: they act as water collectorsfor tropical forelands. TMCFs also harbourextraordinarily many plant species, contributing tooutstanding positions in each of the five hottestspots of plant diversity.Why edit yet another comprehensive opuson TMCFs, after several previous fundamentalworks, despite their encompassing under 0.15% ofthe global terrestrial surface? The answer is clearto those who know this biome: there hardly existsa more fascinating environment than exuberant,moss-covered cloud forests, often called elfinforests due to their mystical appearance. Theyform highly complex ecosystems, which ondifferent continents show divergent biocœnosesbecause of their fragmentation and isolatedposition within distinct tropical mountains in theNeotropics and, to a lesser degree, in thePaleotropics, and in a few cases even on Pacificislands. This book deals with general features ofTMCFs (12 chapters) and contains examples fromMiddle America (21), South America (19),Southeast Asia (10), Africa (5) and Oceania /Australia (5). Most contributions result from aconference in Hawaii in 2004. A first glancereveals a nearly complete thematic spectrum.The book is subdivided into seven sectionswith a total of 72 chapters. The first part containsgeneral features of TMCFs. Altitudinaldistributions are presented in an introductorychapter, though integrative references tosurrounding belts are missing. A useful GIS-basedmodelling approach provides instructive data onTMCF resources and losses including tables ontheir dimensions and distributions. Interestingly,Indonesia and Congo rank first in national extent,with neotropical countries falling lower down. Aclimate chapter is based on a dataset of 477weather stations in cloud forest sites. Manygraphs present vast dot clouds of data fromstations between 200 and 5,000 m asl, whichraises the confusing suspicion that TMCFs occur inregions of extremely dissimilar climates. A shortbut informative chapter on changes in fogprecipitation should have been part of a latersection, as also applies to one on epiphytism.Comments on global and local soil variations, aswell as on nutrient cycling and limitation inTMCFs, provide convincing and compactestimations. Coloured maps of TMCF distributionhighlight their restricted extent and naturalfragmentation on a global scale.The subsequent and sadly brief section onregional aspects of floristic and faunistic diversitycontains fascinating information from all TMCFbearingcontinents. The range extends fromresearch on epiphyte-diversity on solitary trees(up to 4,806 individuals of 114 vascular plantspecies on one single fig tree!) to potential andactual distribution patterns of the mountain tapirand Andean bear. The only point of criticism isthat in a book on a biological realm this sectioncould have been broader.The third section on hydrometeorologycovers a broad remit since fog, rain and theirinterception are decisive triggers for theformation of TMCFs. Several parts display theimportance of potential evaporation andirradiation as driving forces for the variablecharacter of forests. Additionally, the degrees oflitter mineraliation and soil acidity become crucialcauses of ecological peculiarities. Thecontributions vary from rather specialistmethodical content (e.g. measuring interception,usage of stable isotopes for diagnoses ofprecipitation origins) to comments on the waterfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society17

news and update ISSN 1948-6596retention capacity of epiphytes. The tone ofseveral chapters conveys the impression thatTMCFs serve more as open-air laboratories forprocedural techniques rather than as interestingecosystems in their own right.After a section on nutrient dynamics,physiological features such as water use andphotosynthesis in TMCFs are addressed in the fifthsection. Corresponding implications of forestconversions into pastureland and the resultingdramatic alterations in microclimate and soilhydrology once again highlight the unique physicalproperties of these dense and dark stands. Thesixth section on climatic variability and changefurther illustrates effects caused by clear-cut foralternative land usage. The long-rangeimplications of lowland land use change throughvertical shifts in the condensation level, as well asresponses of epiphyte formations to stand climatechanges and their value as indicators, are stressedhere too. Global warming has implications forincreasing rates of fire with fatal consequences forTMCFs. An interesting aside, albeit barely relatedto the climate topic, is the last chapter onmodelling regeneration dynamics and successionrates on abandoned land.Chapters on cloud forest conservation,restoration, and management follow in the lastsection, including historical aspects ofdeforestation and regrowth, partly in the light ofsocial background. Sustainability concepts andlessons for the future are demonstrated byspecific case studies (e.g. epiphyte breeding, firemanagement). Two informative chapters areincluded on assessments needs and payments forthe environmental services provided by TMCFs. Afinal contribution compiles the state of knowledgeand a pathway towards sustainability. This broadoutlook groups core issues into five strategicthemes: ecosystem function, climate change,nature conservation, ecosystem management(and restoration), and species management. Atabular appendix including data on around 50TMCF sites appears isolated and out of place.Two thirds of the authors are employed ininstitutions of extratropical countries, whichillustrates a major problem in TMCF preservation:expert knowledge on functions, values andprotective measures for this endangeredecosystem is concentrated in Europe and the USArather than in the countries concerned. Thedeepening of specific insights provided by thebook might contribute to an improved transfer ofknowledge through scientific cooperation.Despite the efforts of the editors to createan integrated opus it suffers from the patchworkcharacter of symposium volumes, withcontributions of varying quality. Sporadicallyoverlong tables or figures of similar contentconsume much space. Several topics are missing,such as geomorphologic processes or impacts ofecological disturbance regimes. Also absent aregeneral descriptions of typical growth forms andleaf features. Nevertheless, the attractivelydesigned book offers a presentation of copiousthemes and a vast collection of references, andthus should be considered an important sourcefor experts to enlarge their knowledge of TMCFecosystems.Michael RichterInstitute of Geography, University of Erlangen, Germany(retired); sairecabur@web.deEdited by Markus Eichhorn18 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

news and updatebooks noted with interestMarine Protected Areas: A multidisciplinaryapproachJoachim Claudet (editor)2011, Cambridge University Press, 377 pp.AUS$145 (Hardback) / AUS$86.95(Paperback) / US$47.00 (e-book)ISBN: 9780521766050 / 9780521141086 /9781139153614http://www.cambridge.org/In recent years Marine Protected Areas (MPAs)have become a hot topic in fisheries policy. Theireffects scale from local to regional, while anenormous breadth of disciplines are involved intheir study, cutting across ecology, conservationbiology, socioeconomics and politics. An array ofcontributors cover all aspects within this singlevolume, including both theoretical and empiricalapproaches, alongside a strong practical focus. Forthose working in any one area related to MPAsthis will be a valuable source of information onthe linkages among them, and an inspiring insightinto the wider dimensions, right up to the globalscale.Agent-based and individual-based modeling:A practical introductionSteven F. Railsback and Volker Grimm2012, Princeton University Press, 329 pp.$99.50 (Hardback) / $55.00 (Paperback)ISBN: 9780691136738 / 9780691136745http://press.princeton.edu/Many scientific fields have discovered thatmodeling the actions of individual entities canprofoundly alter our interpretation ofphenomena. Biologists, however, have beenrelatively slow to take advantage of enhancedcomputing power and unlock the potential ofthese techniques. This book removes any excuse.Based on a course run by the authors, who bothcome from an ecological background, and buildingon an earlier, more conceptual book, this aims toprovide the necessary tools to students andresearchers. Practical exercises involve the freesoftware NetLogo with updates andsupplementary material available online. Complexbiological systems are suddenly appearing moretractable.Editorial policy for book reviewsFrontiers of Biogeography will publish in-depth reviews of recently published books (typically less than oneyear old) on biogeography or of interest to biogeographers, alongside a ‘Noted with Interest’ section providingbrief details of new publications. Authors, editors or third parties are invited to suggest books for reviewto the Book Review Editor, Dr Markus Eichhorn, School of Biology, University Park, Nottingham NG72RD, United Kingdom; telephone ++44 (0)115 951 3214; e-mail markus.eichhorn@nottingham.ac.uk. Wewelcome offers to review books for Frontiers of Biogeography, but will not accept an offer to review a specificbook. Anyone wishing to review books should send a brief curriculum vitae, description of competencies,and a statement of reviewing interests to the Book Review Editor. Reviews should be in an essay style, expressingan opinion about the value of the book, its focus and breadth, setting it in the context of recentdevelopments within the field of study. Textbook reviews should consider their utility as resources for teachingand learning. Avoid describing the book chapter by chapter or listing typographical errors. The lengthshould normally be 1000 words (1500 words for joint reviews of related texts) including a maximum 10 references.Authors may suggest a short heading for the review, followed by the title of the book(s), the authors/editors,publisher, publication date, price, hbk/pbk, pages, ISBN and website (where available). Figuresor tables will not ordinarily be included. Authors of reviews must verify that they have not offered (and willnot offer) a review of the same book to another journal, and must declare any potential conflict of interestthat might interfere with their objectivity. This may form a basis for editorial decisions and such disclosuresmay be published. Book reviews will usually go through a light editorial review, though in some circumstancesalso will be considered by one or more referees.frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society19

news and update ISSN 1948-6596thesis abstractEcology and biogeography of island parasitoid faunasAna M. C. Santos 1,2,3PhD Thesis, Silwood Park Campus, Division of Biology, Imperial College London, UK1 Departamento de Ecologia, Instituto de Ciências Biológicas, Universidade Federal de Goiás, 74001-970Goiânia, GO, Brazil; 2 Centro de Biologia Ambiental, Faculdade de Ciências da Universidade de Lisboa, C25th Floor, 1749-016 Lisbon, Portugal; 3 Departamento de Biodiversidad y Biología Evolutiva, MuseoNacional de Ciencias Naturales (CSIC), C/ José Gutiérrez Abascal 2, 28006 Madrid, Spainana.margarida.c.santos@googlemail.com; http://www.wix.com/guidasanto1/anamcsantosAbstract. Species on islands tend to use a wider range of resources than their mainland counterparts. Inthis thesis I investigated whether island parasitoid communities have proportionally more idiobiont species(which tend to have a wider host range; i.e. are more generalist) than their mainland source, andwhich factors determine island community structure. These questions were approached using data on thedistribution of Ichneumonoidea species worldwide and data from a survey conducted in the Macaronesianislands and mainland. Prior to the global analyses, I assessed whether islands and archipelagos follow thesame species–area relationship, and identified which islands have comparable inventories. Globally, islandshave proportionally more idiobionts than continental areas, and the species pool for colonization isthe most important determinant of island community structure. Specimens collected in the Macaronesianregion were tentatively identified using a protocol based on host dissection and DNA barcoding. At thisscale, mainland faunas have proportionally more koinobiont species and island communities have agreater proportion of idiobionts.Keywords: community structure, DNA barcoding, host-parasitoid interactions, host range, island biogeography,island species–area relationship, Macaronesia, species poolIslands constitute natural laboratories for thestudy of evolutionary and ecological processesbecause of their discrete and isolated nature,small size and simplified biotas. Island faunas, especiallythose of oceanic islands, tend to be species-poorand disharmonic, meaning that thereare often fewer species on an oceanic island thanon a same-sized area of mainland, and that thestructure of their communities is different fromtheir continental counterparts (e.g. some trophicguilds can be absent from island communities).The phenomenon of ecological release, typical inmany island populations, occurs when a speciescolonizing an island encounters a new environmentin which competitors and/or predators aremissing (Whittaker and Fernández-Palacios 2007).One of the consequences of this process is theexpansion to empty or invasible niche space, leadingto niche expansion and/or niche shifts, withspecies from island assemblages often using awider range of resources than their counterpartsfrom the source mainland. Therefore it is not surprisingthat many oceanic islands have a high representationof generalist species when comparedto the mainland (e.g. Olesen and Valido 2003). Inaddition, some evidence suggests that generalistspecies may simply have an a priori advantageduring the colonization process (Piechnik et al.2008).Parasitoids are generally defined as insectswhose larvae develop to adulthood by feeding inor on the body of an arthropod host, eventuallycausing its death (Quicke 1997; see more detailsof their biology in Santos & Quicke 2011). Parasitoidsare usually divided into two different groupsthat reflect different life-history strategies: koinobionts,which allow the host to continue its developmentafter oviposition of the parasitoid, andidiobionts, which do not (Askew and Shaw 1986).Many life-history traits appear to be correlatedwith this dichotomy (e.g. Hawkins 1994, Quicke1997; see Table 1). Host range, i.e. the group of20 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Ana M. C. SantosTable 1. Life-history traits associated with the dichotomy idiobionts vs. koinobionts. However, there is a wide spectrumfilled between idealized idiobiont and koinobiont strategies, and there are exceptions to virtually any generalizationabout them.IdiobiontsEctoparasitismPermanently paralyse their hostsDo not allow the host to continue its developmentLittle interaction with their hosts’ immune systemsAttack a wider variety of hostsHost generalistsShort larval development timeLonger adult lifeTend to attack later host stages (except for the eggs)Hosts live in concealed situationsSynovigenic (produce few, large eggs that develop overthe life of the adult wasp)KoinobiontsEndoparasitismDo not paralyse their hosts (or only temporarily)Allow the host to continue its developmentInteract with their hosts’ immune systemsAttack a more restricted number of hostsHost specialistsLong larval development timeShort adult lifeHosts are in relatively young developmental stagesHosts are exposedPro-ovigenic (produce many small eggs, that are all fullydeveloped the moment the wasp hatches)potential hosts that a parasitoid species can attacksuccessfully after exhibiting a certain pattern ofsearching behavior that allows it to find hostsregularly (Shaw 1994), is also related to this dichotomy,with idiobionts typically having a widerhost range and koinobionts being consideredmore specialist (e.g. Askew and Shaw 1986, Hawkins1994, Shaw 1994). This pattern is largely dueto the fact that koinobionts interact more intimatelywith their hosts’ immune systems, whileidiobionts either do not have to deal at all withtheir hosts’ immune system or attack hosts withreduced immunological systems, such as eggs orpupae (e.g. Shaw 1994, Quicke 1997).Despite their ecological importance in mostterrestrial systems (LaSalle and Gauld 1993), verylittle is known about parasitoids’ diversity, distributionand biology, particularly on islands. Regardingthe community assembly of island parasitoids,one could expect that island faunas are biasedtowards generalist species (i.e. idiobionts),since hosts on islands may be unusual or novelcompared with those on the mainland. However,the few previous studies that looked into changesin host ranges at large geographical scales gavecontradictory information; some showed that generalistsare better in adapting to new habitatsand new hosts (e.g. Cornell and Hawkins 1993),while others indicated that host range cannot bepredicted from biological or ecological traits of theparasitoids (e.g. Hawkins and Marino 1997). Onestudy even found that the early-phase parasitoidcolonists of Anak Krakatau island were dominatedby taxa that are koinobionts of Lepidoptera, althoughno comparison with mainland communitieswas attempted (Maetô and Thornton 1993).The general aim of my PhD thesis was tostudy the geographic patterns of generalism andspecialism in island parasitoid faunas, with themain hypothesis being that island parasitoid faunasare biased towards idiobiont species whencompared to the mainland faunas. This hypothesiswas analysed using an interdisciplinary approachthat integrated two different geographical scales:a global scale using data on the distribution of Ichneumonoidea(Hymenoptera) species worldwide,and a regional scale using data collected in someislands and continental areas of the Macaronesianregion.At the global scale, we evaluated whetherisland parasitoid faunas are biased towards idiobiontspecies when compared with the correspondingspecies pool, and examined which factors,of those usually thought to control theassembly of island faunas, also have an affect onthe ratio between idiobionts and koinobionts onthe islands. To investigate this we used apublished database on the distribution offrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society21

island parasitoid community structureBraconidae and Ichneumonidae worldwide(Taxapad; Yu et al. 2005). In this database the informationon parasitoid inventories is often presentedfor archipelagos rather than for single islands.Therefore, we first examined whether thereis a consistency in the processes building up thebiotas of single islands and entire archipelagos,assessing whether archipelagos follow the samespecies–area relationship as their constituent islands(Santos et al. 2010a). Here we found thatarchipelagos do often follow the same islandspecies–area relationship (ISAR) of theirconstituent islands, and that the archipelagicpoint (corresponding to the total area andrichness of the island group) is congruent with itsISAR. Among other things, such consistencyimplies that both islands and archipelagos can beused as distinct units themselves in large-scalebiogeographical and macroecological studies. Thearchipelagic residual (calculated as the residual ofthe prediction provided by the ISAR using the totalarea of the archipelago, standardized by totalrichness) indicated that the ISAR underpredictsarchipelagic richness in the least isolatedarchipelagos. Also, the magnitude of thedeparture from the ISAR was related tonestedness; the more nested the biota of thearchipelago, the lower the archipelagic residual.Departures from the ISAR are thus expected insystems that are either highly nested or notnested at all; in highly nested systems, thepredicted number of species for the total area ofthe archipelago will be higher than the observedspecies richness, while in non-nested systems theobserved archipelagic species richness should behigher than that predicted by the ISAR.Most studies on large-scale diversitygradients are based on biodiversity databases thatcompile information on the distribution of speciesgathered from an often heterogeneous range ofdifferent inventories and methodologies (Hortal etal. 2007). These data are not free from errors; it iswell known that our knowledge of thegeographical distribution of biodiversity is, ingeneral, taxonomically and geographically biased(e.g. Jones et al. 2009). Therefore, prior to furtheranalysis it is advisable to evaluate data quality,and assess the consistency of the results amongdifferent kinds of territorial units (that can differin size and nature). Since the database we used isa compilation of all available knowledge on thedistribution of the studied families, it was expectedthat the quality and completeness of thedata on Braconidae and Ichneumonidae faunas isalso uneven. Therefore, we evaluated the biasesand problems to ensure that the data usedincluded comparable units with no majorshortfalls. To achieve this, we developed a simplescoring method that did not rely on measures ofsampling effort (e.g. number of survey records,individuals or traps), and that would allowassessment of which islands have comparableinventories. The protocol we proposed is based inthree criteria: (i) completeness at high taxonomiclevels, which accounts for the effort made indescribing and inventorying species from differenthigh-level taxa and indicates any potential biastowards particular taxa; (ii) congruence with wellestablishedecological relationships, whichassumes that obvious outliers in the species–arearelationship are unlikely to have been adequatelyinventoried; and (iii) publication effort received,which determines whether a significant amount ofinventory effort was devoted to the territorialunit, using the number of pages in the workscompiled in the database as a proxy for samplingeffort (see Santos et al. 2010b).Finally, the islands with comparableinventories were used to examine whether islandfaunas are biased towards idiobionts, and toevaluate the relationships between different environmental,physical and regional factors and therelative proportions of idiobionts and koinobiontsin both Braconidae and Ichneumonidae. All specieswere classified as either idiobionts or koinobiontsaccording to the known biology of the subfamiliesthey belong to. This trait is well conservedat this taxonomic level (Askew & Shaw 1986,Gauld 1988) and so this classification gives a fairlygood estimate of host range (Hawkins et al. 1992).Results indicated that, for pooled data, islandshave a higher proportion of idiobionts than themainland. This does not seem to be due to therecurrent presence of certain subfamilies on is-22 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Ana M. C. Santoslands, as of the six subfamilies that are more representedon islands (i.e. that have more than 25%of their species present on islands) only two includeidiobiont species. However, in pairwise comparisonsmost islands have a similar proportion ofidiobionts to that found in the adjacent mainlandspecies pool, and the composition of this colonizationpool seems to be a key factor determining thestructure of island parasitoid communities. Thereis also a latitudinal gradient in the level of generalismof island faunas (measured as the proportionbetween idiobiont and koinobiont species), whichmay be the outcome of some environmental factorsand island characteristics, such as temperature,altitude and island species richness in thecase of the braconids, or region, island type andprecipitation in the case of the ichneumonids. Islandsof the Indomalayan region are particularlydominated by idiobiont species, which might bedue to the fact that these islands are home tolarge tropical rainforests 1 , where plant architecturalcomplexity (particularly of trees) is probablyresponsible for increasing idiobiont species richness(e.g. Hawkins et al. 1990, Hawkins 1994).Such results highlight the complexity of factorsshaping the diversity and structure of parasitoidcommunities (Santos et al. 2011a).At the regional scale, we studied how thediversity, parasitism rates and attack strategy ofthe parasitoid communities associated with a particularhost system vary between the islands andadjacent mainland areas of the Macaronesian region.In particular, we studied the community ofparasitoids attacking the tortricid moth Acroclitasubsequana that feeds on spurges (Euphorbiaspp., Euphorbiaceae). Traditionally, host–parasitoid interactions are ascertained throughrearing, which is labour intensive, time consumingand requires experience. Moreover, this techniqueis very difficult to apply when studying geographicalvariations of host–parasitoid interactionsat large scales, as it would require severalfield stations and the support of a large team. Toovercome this problem, we investigated geographicalvariations in host–parasitoid interactionsusing a new protocol based on host dissection andDNA barcoding (Santos et al. 2011b). Althoughthis protocol is somewhat time consuming andrequires the use of a fully equipped laboratory, itnevertheless has several advantages when comparedwith standard rearing methods: (i) it doesnot require a taxonomic specialist for identifications;(ii) a small team of researchers, or even asingle person (as in the case of this thesis) is sufficient;and (iii) it can be used for studies spreadacross several different regions. This protocol allowseach sequence (i.e. specimen) to be assignedto a Molecular Taxonomic Operational Unit(MOTU) that is usually defined as a cluster of sequenceswith pairwise distances below a certainthreshold. However, it might not always be possibleto correctly identify each MOTU to the specieslevel because available sequence databases arefar from complete, and are not yet reliableenough to be used for the identification of poorlystudied and hyperdiverse taxa such as parasiticHymenoptera and Microlepidoptera. Still, dependingon the goal of the particular research project,this might not be a problem because MOTUs canbe used as surrogate units of diversity, enabling usto produce parasitoid food-webs and quantifyhost–parasitoid interactions.Once all parasitoids found were assigned toMOTUs, we tested whether species richness andparasitism rates differ between islands andmainland, and whether island parasitoid faunasare biased towards idiobiont species (Santos et al.2011c). Once again, species were classified as eitheridiobionts or koinobionts in accordance tothe known biology on the subfamilies they belongto. The results showed that, overall, parasitoidspecies richness and parasitism rate were similaron islands and mainland. However, mainland speciesrichness was lower than expected from a randommodel, with communities being dominatedby koinobiont species; on the other hand, islandparasitoid communities were dominated by idiobionts.Also, islands had higher parasitism rates byidiobionts than expected from a random model,and mainland areas showed the highest koinobi-1 See the global land-cover map at http://eoimages.gsfc.nasa.gov/images/news/NasaNews/ReleaseImages/LCC/Images/lcc_global_2048.jpgfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society23

island parasitoid community structureont parasitism rates. These results suggest thatisland parasitoid communities are biased in favourof idiobionts (that tend to be more generalist),when compared with mainland communities. Theprocesses behind such patterns still need to beexplored, but they might be related to (i) the possibilitythat generalist parasitoids are better dispersers;(ii) the expectation that they are less constrainedby “sequential dependencies” (Holt et al.1999), not being dependent on the presence of aparticular resource; and (iii) the likelihood thatisland parasitoids have fewer competitors, favouringgeneralists.There are obvious differences in the natureof the data between the two scales studied in thisthesis. Data from the global scale came from agreat variety of habitats and hosts, and refer tothe available parasitoid species checklists. On theother hand, data from the regional scale originatedfrom a particular, environmentally quitehomogeneous area, and from the community ofparasitoids associated with just one host system.Nevertheless, results from both scales indicate abias towards idiobiont species on islands. At theglobal scale, pooled island parasitoid faunas hadcomparatively more idiobiont species than theentire pool of mainland areas, while at the regionalscale, island hosts suffered higher attackrates by generalist parasitoids. However, in theglobal analyses only a small number of islands departedsignificantly from the structure of theircolonization pool. In contrast, the structure of theisland parasitoid communities studied at the regionalscale differs from that of its colonizationpool. A hypothesis linking together the resultsfrom both scales is that the ecological processesthat determine island community structure regionallyscale up and result in higher proportionsof generalists at larger scales. Therefore, differencesin the results from the two approaches exploredhere may be due to differences in scale.Following this argument, one could consider thatfactors acting at the biogeographical scale maskthe effect of the ecological processes that lead toa general trend towards higher parasitoid generalismon islands, making the trend less apparentworldwide. Here, the structure of the species poolwould be the main determinant of the structureof island communities worldwide, but other factorswould also play a role in determining islandparasitoid faunas – causing some island parasitoidcommunities to deviate from their overall tendencyto higher proportions of generalists (Santos& Quicke 2011).AcknowledgementsI would like to thank my two supervisors, DonaldQuicke and Paulo Borges for their support andadvice. I also want to acknowledge Joaquín Hortal,Guillaume Besnard, Colin Fontaine, Owen Jones,Kostas Triantis and Rob Whittaker for theirvaluable contributions to my work. AMCS wassupported by two Portuguese FCT grants (SFRH/BD/21496/2005; SFRH/BPD/70709/2010) cofundedby the European Social Fund POPH-QRENprogram, and by a Brazilian CNPq juniorpostdoctoral fellowship (159763/2010-0).ReferencesAskew, R.R. & Shaw, M.R. (1986) Parasitoidcommunities: their size, structure anddevelopment. In: Insect parasitoids (ed. by J.Waage and D. Greathead), pp.225–264.Academic Press, London.Cornell, H.V. & Hawkins, B.A. (1993) Accumulation ofnative parasitoid species on introducedherbivores: a comparison of hosts as natives andhosts as invaders. The American Naturalist, 141,847–865.Gauld, I.D. (1988) Evolutionary patterns of hostutilization by ichneumonoid parasitoids(Hymenoptera: Ichneumonidae andBraconidae). Biological Journal of the LinneanSociety, 35, 351–377.Hawkins, B.A. (1994) Pattern and process in host–parasitoid interactions. Cambridge UniversityPress, Cambridge.Hawkins, B.A. & Marino, P.C. (1997) The colonization ofnative phytophagous insects in North Americaby exotic parasitoids. Oecologia, 112, 556–571.Hawkins, B.A., Askew, R.R. & Shaw MR (1990)Influences of host feeding-niche and foodplanttype on generalist and specialist parasitoids.Ecological Entomology 15, 275–280.Hawkins, B.A., Shaw, M.R. & Askew, R.R. (1992)Relations among assemblage size, hostspecialization, and climatic variability in North-American parasitoid communities. AmericanNaturalist, 139, 58–79.24 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Ana M. C. SantosHolt, R.D., Lawton, J.H., Polis, G.A. & Martinez, N.D.(1999) Trophic rank and the species–arearelationship. Ecology, 80, 1495–1504.Hortal, J., Lobo, J.M. & Jiménez-Valverde, A. (2007)Limitations of biodiversity databases: case studyon seed-plant diversity in Tenerife, CanaryIslands. Conservation Biology, 21, 853–863.Jones, O.R., Purvis, A., Baumgart, E. & Quicke, D.L.J.(2009) Using taxonomic revision data toestimate the geographic and taxonomicdistribution of undescribed species richness int h e B r a c o n i d a e ( H y m e n o p t e r a :Ichneumonoidea). Insect Conservation andDiversity, 2, 204–212.LaSalle, J. & Gauld, I.D. (1993) Hymenoptera andbiodiversity. CAB International, Wallingford.Maetô, K. & Thornton, I.W.B. (1993) A preliminaryappraisal of the braconid (Hymenoptera) faunaof the Krakatau Islands, Indonesia, in 1984–1986, with comments on the colonizing abilitiesof parasitoid modes. Japanese Journal ofEntomology, 61, 787–801.Olesen, J.M. & Valido, A. (2003) Lizards as pollinatorsand seed dispersers: an island phenomenon.Trends in Ecology and Evolution, 18, 177–181.Piechnik, D.A., Lawler, S.P. & Martinez, N.D. (2008)Food-web assembly during a classicbiogeographic study: species’ “trophic breadth”corresponds to colonization order. Oikos, 117,665–674.Quicke DLJ (1997) Parasitic wasps. Chapman and Hall,London.Santos, A.M.C., Whittaker, R.J., Triantis, K.A., Borges,P.A.V., Jones, O.R., Quicke, D.L.J. & Hortal, J.(2010a) Are species–area relationships fromentire archipelagos congruent with those oftheir constituent islands? Global Ecology &Biogeography, 19, 527–540.Santos, A.M.C., Jones, O.R., Quicke, D.L.J. & Hortal, J.(2010b) Assessing the reliability of biodiversitydatabases: identifying evenly inventoried islandp a r a s i t o i d f a u n a s ( H y m e n o p t e r a :Ichneumonoidea) worldwide. InsectConservation and Diversity, 3, 72–82.Santos, A.M.C. & Quicke, D.L.J. (2011) Large-scalediversity patterns of parasitoid insects.Entomological Science, 14, 371–382.Santos, A.M.C., Quicke, D.L.J., Borges, P.A.V. & Hortal,J. (2011a) Species pool structure determines thedegree of generalism of island paraistoid faunas.Journal of Biogeography, 38, 1657–1667.Santos, A.M.C., Besnard, G. & Quicke, D.L.J. (2011b)Applying DNA barcoding for the study ofgeographical variation in host–parasitoidinteractions. Molecular Ecology Resources, 11,46–59.Santos, A.M.C., Fontaine, C., Quicke, D.L.J., Borges,P.A.V. & Hortal, J. (2011c) Are island andmainland biotas different? Richness and level ofgeneralism in parasitoids of a microlepidopteranin Macaronesia. Oikos, 120, 1256–1262.Shaw, M.R. (1994) Parasitoid host range. In: ParasitoidCommunity Ecology (ed. By B.A. & W. Sheenan),pp. 111–144. Oxford Univesity Press, New York.Whittaker, R.J. & Fernández-Palacios, J.M. (2007) Islandbiogeography: ecology, evolution, andconservation. Oxford University Press, Oxford.Yu, D.S., van Achterberg, K.B. & Horstmann, K.I. (2005)World Ichneumonoidea 2004: taxonomy,biology, morphology and distribution. CD-Rom.Taxapad, Vancouver, Canada.Edited by Jan BeckYou can find information about the International Biogeography Society at http://www.biogeography.org/, and contact with other biogeographers at the IBS blog (http://biogeography.blogspot.com/), the IBS facebook group (http://www.facebook.com/group.php?gid=6908354463) and the IBS twitter channel (https://twitter.com/biogeography).frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society25

opinion and perspectivesperspectiveLosing time? Incorporating a deeper temporalperspective into modern ecologyFelisa A. Smith 1 and Alison G. Boyer 2ISSN 1948-65961 Department of Biology, University of New Mexico, Albuquerque, NM 87131, USA.fasmith@unm.edu; http://biology.unm.edu/fasmith/2 Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville,TN 37996, USA. alison.boyer@utk.edu; http://eeb.bio.utk.edu/boyer/index.htmlAbstract. Ecologists readily acknowledge that a temporal perspective is essential for untanglingecological complexity, yet most studies remain of relatively short duration. Despite a number ofexcellent essays on the topic, only recently have ecologists begun to explicitly incorporate a historicalcomponent. Here we provide several concrete examples drawn largely from our own work that clearlyillustrate how the adoption of a longer temporal perspective produces results significantly at odds withthose obtained when relying solely on modern data. We focus on projects in the areas of conservation,global change and macroecology because such work often relies on broad-scale or synthetic data thatmay be heavily influenced by historic or prehistoric anthropogenic factors. Our analysis suggests thatconsiderable care should be taken when extrapolating from studies of extant systems. Few, if any,modern systems have been unaffected by anthropogenic influences. We encourage the furtherintegration between paleoecologists and ecologists, who have been historically segregated intodifferent departments, scientific societies and scientific cultures.Keywords: climate change, conservation, macroecology, paleoecology, palaeoecology, woodratThe pronghorn (Antilocapra americana) is aquintessential symbol of the Great Plains. As thefastest land mammal in the Americas, it can reachclose to ~100 km/h and can sustain speeds of 45km/h for long distances (Byers 1997). Much of itsphysiology, morphology and life history reflect anoptimization for being swift; pronghorn haveoversized hearts and lungs, a 320° field of vision,hollow hair and overlong gestation for their size(Byers 1997). Understanding the selectivepressures that led to such specialized adaptationsis difficult without the knowledge that thepronghorn co-evolved with a suite of now extinctpredators, including the American cheetah(Micracinonyx trumani) (Byers 1997, Barlow2001). As the only surviving member of the oncespeciose North American family Antilocapridae,the pronghorn no longer has effective naturalpredators. Consequently, many of its social,morphological and physiological traits have littleapparent modern selective value (Byers 1997).Ecologists recognize the anachronisticnature of animals like the pronghorn, but more asa curiosity rather than as a concrete example ofthe substantial alteration of ecosystems thatoccurred in the late Quaternary. Although workinvestigating the ecology and life-historycharacteristics of tropical and temperate plantshas proposed that numerous adaptations fordispersal or regrowth arose in response toforaging by now-extinct megafauna (Janzen andMartin 1982, Wing and Tiffney 1987, Barlow2001), in general, the implications of theprehistoric loss of megafauna in the latePleistocene have been overlooked. Yet, theseanimals undoubtedly played key roles in terms ofecosystem structure and function; their abruptdisappearance some 11,000 years ago must haveprofoundly influenced ecosystem dynamics(Martin 1967, Donlan et al. 2005). How manyother life-history, ecological or distributionalfeatures of extant animals and plants are due insome part to now-extinct components of theecosystem?26 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Felisa A. Smith and Alison G. BoyerAs ecologists increasingly turn from‘explaining the present’ to ‘anticipating thefuture’, there has been renewed interest inbringing a historical perspective into ecology(Botkin et al. 2007, Gavin et al. 2007, Williams andJackson 2007). While earlier workers illustratedthe insights a wider temporal window yields (e.g.,Schoonmaker and Foster 1991, Herrera 1992,Delcourt and Delcourt 1998), the contemporaryfocus on anthropogenic climate change hasgalvanized efforts. After all, the late Quaternaryprovides abundant examples for examining theinfluence of changing abiotic conditions onorganism distribution, ecology and evolution(Clark et al. 2001, Botkin et al. 2007, Gavin et al.2007, Williams and Jackson 2007). Evidence fromfine-scale paleoclimate reconstructions (e.g.,pollen, cross-dated tree-ring chronologies, icecores, and other indicators) suggests abruptclimate shifts occurred with regularity in the past(Schoonmaker and Foster 1991, Allen andAnderson 1993, Dansgarrd et al. 1993, Bond andLotti 1995, Alley 2000). Some, such as the YoungerDryas, were significant events, with temperaturewarming of as much as 5–10°C reportedlyoccurring within a decade (Alley et al. 1993, Alley2000), a rate of change higher than that expectedunder most scenarios of anthropogenic climatechange (IPCC 2007). Virtually all species extanttoday were present and successfully coped withthe Younger Dryas. Thus, it has been recognized asa particularly useful analog for studying the likelyeffects of anthropogenic climate change. Indeed,the most recent IPCC report now contains sectionson paleoclimate.The de facto standard traditionally used byecologists to set ecological baselines is to replicateexperiments across space. The implicit assumptionis that if the spatial extent is sufficient itencompasses the possible range of naturalvariation. However, more than 20 years ago it wasrecognized that conceptual problems occur ifecologists use “short-term experiments to addresslong-term questions” (Tilman 1989, pg 139).Although the incorporation of a broader spatialperspective clearly increases the natural range ofvariation expressed in both abiotic and bioticconditions, space is not necessarily an adequatesubstitute for time. Simply put, ecological historymatters. This disparity may occur because of nonanalogclimatic conditions found in the past,leading to assemblages of mammal or vegetativecommunities not found together today (e.g.,Huntley 1990, Schoonmaker and Foster 1991,Overpeck et al. 1992, Graham et al. 1996, Williamset al. 2001, Williams and Jackson 2007), orbecause the type or magnitude of change in thepast dwarfs that represented along a modernspatial gradient (Jackson 2007).Here, we provide several concrete examplesof how the adoption of a deeper temporalperspective can sometimes provide morecomplete and often divergent insights intomodern ecology. These are drawn largely fromour own work or that of close colleagues becauseit was otherwise difficult to obtain the originaldata that would allow us to redo the analyses. Ourfirst examples demonstrate how limitingmacroecological studies to extant taxa may yieldskewed interpretations of broad-scale geographicpatterns. The second set of examplesdemonstrates the added utility that a longertemporal perspective can provide for conservationbiology. Both paleontologists and conservationbiologists are interested in extinction, andunderstanding past events could aid in theunderstanding of current risk for many taxa.Finally, we focus on studies of the likely effect ofanthropogenic climate change on organisms andecosystems. Climate scientists have traditionallyused forward-projected models with baselinesestablished using modern conditions. Given thelikelihood of non-analog climatic regimes in thefuture, models that are parameterized based onlyon modern conditions are likely to fail toaccurately predict ecological responses to novelclimates. Here, a historical perspective can beparticularly useful. Our intent is to demonstratethe need to integrate both paleontological andecological approaches in developing a synopticunderstanding of ecological systems.frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society27

losing time?Ecological examplesBody size distributions in macroecologyMacroecology was developed in the late 1980s asan effort to understand patterns underlying thelocal abundance, distribution and diversity ofspecies (Brown and Maurer 1989). As acomplementary approach to experimentalecological research, it has been remarkablysuccessful at illuminating large-scale spatial andstatistical patterns (Smith et al. 2008 andreferences therein). Body size is often used as avariable of interest in macroecological studiesbecause it is tightly related to many fundamentalphysiological, ecological and evolutionarycharacteristics, and moreover, is relatively easy tocharacterize even for fossil forms (Peters 1983,Damuth and MacFadden 1990). But what if theunderlying body-size distribution used in ananalysis is biased or incomplete? The use of bodysize as a dependent variable could potentially leadto misleading results if portions of the biota areselectively missing.In 1991, Brown and Nicoletto reported thatthe shapes of mammalian body-size distributionsin North America change with spatial scale. Thecontinental-level body-size distribution wasunimodal and right skewed, but as spatial scaledecreased, regional distributions becameprogressively flatter until they were nearlyuniform for local communities. Brown andNicoletto’s results had important implications interms of community assembly and structure andthe paper has been highly cited. Because speciesfound at local scales were not a random subsampleof the regional scale (differing in median,mean, skew and range of size), they argued therewere ‘rules’ influencing the assembly ofcommunities and a limit to the number of speciesof each body size that could co-exist locally.Distributions became peaked as sites wereaggregated over space because of highertaxonomic turnover in smaller-bodied species.Numerous authors have debated the validity ofboth the patterns and underlying mechanismssince this seminal paper was published; forexample, mammalian communities in SouthAmerican tropical forests reportedly show morepeaked distributions than those in other habitats(Marquet and Cofré 1999, Bakker and Kelt 2000)and the body-size distributions of bats are not flatat the local level across a wide range of latitudes(Willig et al. 2008).To what extent were the macroecologicalpatterns described by Brown and Nicoletto (1991)influenced by the use of extant North Americanmammals? The contemporary distributions ofboth North and South American mammals wereheavily impacted by the human-mediated latePleistocene megafaunal extinction (Martin 1967,Surovell and Waguespack 2009). This event wasextremely size biased; although ‘only’ 12.8% ofthe North American mammal fauna wereextirpated, they were mostly the largest speciespresent (Lyons et al. 2004). To address thesensitivity of these macroecological patterns, wereanalyzed the results reported by Brown andNicoletto (1991) at the local, regional andcontinental level, using a global database of bodysize in late Quaternary mammals (Smith et al.2003). As an example, we present the mammalianbody-size distribution of a county in New Mexiconested within the western grasslands biome ofNorth America. For each of these areas wedetermined the likely presence of extinct speciesbased on FAUNMAP range reconstructions andlocal fossil evidence (Harris 1970, Graham et al.1996, Wilson and Ruff 1999, NPS 2007).Our analysis suggests little sensitivity toinclusion of extinct species at the local level (Fig.1); although the range of mammalian body size atsites was underrepresented (e.g., the local areasupported larger animals than the maximumpresent today), the shape of the distribution wasnot significantly different. At coarser spatialscales, however, both the range of body size andthe shape of the distribution changed significantly(Kolmogorov–Smirnov two-sample tests, regional:P < 0.05, continental: P < 0.01). The body sizedistribution at both the regional and continentallevels contains a second mode of larger-bodiedmammals (Fig. 1). Recent work by Lyons et al.(2004) suggests that such multimodality is typicalof all continents when extinct late-Pleistocenemegafauna are included.28 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Felisa A. Smith and Alison G. BoyerFigure 1. Body size frequency distributions of North American mammal species before (black shading) and after (greyshading) late-Pleistocene extinctions. Distributions are shown at three different scales: the continent of North America(top), regional (a western grassland biome; middle), and local (Eddy County, New Mexico; bottom). While significantdifferences between past and present body size distributions are observed at the continental and regionalscales, patterns were statistically indistinguishable at the local scale.What does this mean in terms of ecosystemfunction? Macroecologists routinely use patternsin the body-mass distributions of animals as abasis for understanding the structure, assemblyand persistence of ecological communities. Manyhypotheses have been proposed to explain bodysizedistributions at various levels, including onesbased on energetics, ecology, phylogeny,biogeography and habitat or texturaldiscontinuities. However, a longer temporalperspective reveals the influence of the sizebiasedextinction on the shape of contemporarybody-size distributions. Clearly large animals havepivotal roles in ecosystems (e.g., Owen-Smith1987, Pringle et al. 2007), and integrativepaleoecological studies are beginning to revealthe profound effects of megafaunal extinction onvegetation structure, composition, and dynamics,both in North America (Gill et al. 2009), andelsewhere (Hansen and Galetti 2009, Johnson2009).Scaling of landmass area and body sizeSpace use in animals is strongly linked to body sizeand has been a focal point of muchmacroecological research (Brown and Maurer1989, Brown 1995, Gaston 2003, Jetz et al. 2004).Marquet and Taper (1998), and later Burness et al.(2001), observed that the size of the largestmammal on a given landmass increases with landarea. To explain this pattern, they noted that largemammals are characterized by both lowpopulation densities and large home ranges.Recent studies of mammalian (Okie and Brownfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society29

losing time?2009) and avian (Boyer and Jetz 2010) body sizeon islands also found robust scaling relationshipsbetween maximum size and island area. Theseeffects imply that to persist large species requirelarge areas to sustain viable population sizes, andimportant concept in reserve design andconservation of large-bodied mammals (Kelt andVan Vuren 2001). However, Jetz et al. (2004)found a high degree of home range overlap inlarge mammal species, suggesting that populationdensity rather than home range size is the bettermeasure to use in quantifying individual areaneeds for conservation purposes. Since manyislands experienced extinctions during the latePleistocene (Alcover et al. 1998), and theseextinctions may have affected the local body-sizedistribution (Lyons et al. 2004, Boyer and Jetz2010), we re-examined the scaling of maximumsize with land area before the influence of humanmediatedextinctions.We gathered data on the largest mammalspecies found today and in the late Pleistocene on30 islands and landmasses around the world.Mammal data were limited to herbivorous andomnivorous species, owing to differences in thescaling of population density and space usebetween carnivores and herbivores (Peters 1983,Jetz et al. 2004). Island area was based on present-day measurements. Because island mammalswould have experienced a dynamic land area dueto eustatic sea-level changes during the late-Pleistocene, and because the extinct taxa in ourdataset also differ in their dates of lastappearance, we found it difficult to assign a singlelate-Pleistocene value for land area to each island.However, because sea levels in most areas wereover 100m lower than present levels during thelast glacial maximum (Fleming et al. 1998), theland area of many islands would have beensubstantially larger during the late-Pleistoceneand some islands were connected to nearbycontinents by exposed land bridges. To control forthese issues, we excluded all land-bridge islandsand islands where extinction occurred when sealevels were substantially lower than current levels(ca. 7000 years before present, Fleming et al.1998). For comparison to the island data, we alsoincluded late-Pleistocene and modern body massFigure 2. Relationship between maximum body size and area of landmass for extant (open circles, N = 28) and latePleistocene (closed circles, N = 30) mammals. Modern data are missing for two islands, Barbuda and East Falkland,due to the extinction of all terrestrial, non-carnivorous, mammals. Both body size and area were log 10 -transformedprior to analysis. Slopes were indistinguishable between the two time periods, but the intercept for late-Pleistocenemammals was significantly larger (ANOVA; p < 0.01) than for extant species, suggesting that islands supported largeranimals in the past.30 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Felisa A. Smith and Alison G. Boyervalues and modern land area for six continentallandmasses (Australia, New Guinea, SouthAmerica, North America, Africa, and Eurasia).Body size (g) and area (km²) were log 10 -transformed prior to analysis. We compared thesize of the largest mammal species on each islandbefore and after the extinction, and plotted thesebody size maxima against land area (Fig. 2).The strength and direction of the scalingbetween maximum body size and area in extantspecies was not significantly altered by theinclusion of extinct species, but the intercept wassignificantly higher when late-Pleistocene specieswere included (ANOVA, P < 0.01, df=1,54, F=7.4)than for extant species (Fig. 2). This translates toan order of magnitude decrease in the size of thelargest mammal supported by a given land areasince the late Pleistocene. For example, accordingto the modern data, the largest mammal that anisland of 1000 km 2 would be expected to supportis about 431g, however, in the late-Pleistocenefauna, this maximum size was 2291g.In order to compare the scaling of body sizewith area to other studies, we computed the slopeof the relationship with body mass as theindependent variable (late-Pleistocene: slope =0.86, extant: slope = 0.71). Late-Pleistocenefaunas were statistically consistent with previousstudies of both the scaling of maximum body sizewith area in extinction-structured communities(slope = 0.79, Marquet and Taper 1998) and thebody-size scaling of population density inmammals (slope = 0.76, Jetz et al. 2004),suggesting that late-Pleistocene island faunasaccurately represent the ecologically constrainedscaling of maximum body size with area. It isbeyond the scope of this study to determine theecological mechanisms that allowed larger speciesto inhabit smaller areas in the past than thatobserved today; differences in scaling interceptmay relate to the allocation of land area andresurces for use by humans and their commensals(Boyer and Jetz 2010). However, it is clear thatmacroecological studies based solely on extantspecies, especially those conducted in areasknown to be affected by recent extinctions, mayoffer an incomplete picture of these ecosystems.Temporal perspective in conservation studiesA central goal of conservation biology is tounderstand the extinction process in order tomitigate current and future anthropogenicbiodiversity losses. Extinction risk analyses, likemany other predictive ecological studies, areoften based on current distributions of extantspecies (Cardillo 2003, Jones et al. 2003). Longertemporal records, however, can provide analternative perspective on how conservationistsview extinctions. From this perspective, near-timefossil data can be invaluable for identifyinggeneral patterns of extinction risk (McKinney1997, Willis et al. 2007, Boyer 2010). The inclusionof fossil data has several advantages: unlike dataon extant endangered species, fossil data providedirect information on the extinction process itself.Rarity and extinction do not always result fromthe same processes. Second, fossil data representan independent dataset of extinction probabilityon which to build predictive models. This avoidsthe circularity inherent in building a model andtesting it on the same dataset (such as the IUCNRed List). Not only can paleoecological dataprovide a comparison of prehistoric (>500 years)and historic (past 500 years) extinctions, but suchdata may also aid in determining baselineconditions for conservation and restoration, andhelp predict future extinction risk.As a conservation-oriented case study, weturn to the Holocene extinction of birds on Pacificislands. In the Hawaiian islands, the arrival ofPolynesian colonists about 1200 years agocorresponded with the extinction of about 50% ofindigenous land bird species (56/111 species;Olson and James 1982, 1991). In comparison,historic losses of Hawaiian birds amount to about40% of the historically observed species (23/55species; Boyer 2008). To put global Holoceneavian extinctions in context, Pimm et al. (2006)estimated extinction rates for pre-European andhistoric timescales, and compared these to thegeneric background of ~1 extinction per millionspecies per year (E/MSY). Historic rates werearound 26 E/MSY, but after accounting for pre-European extinctions, the estimate rose to ~100E/MSY. The vast majority of recorded extinctionsfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society31

losing time?before the 20 th century were on islands. If themost vulnerable species were lost quickly afterhuman colonization, we may expect rates onislands to slow down over time (Pimm et al. 1994).However, human impacts on island environmentshave intensified through time, so how doprehistoric and historic extinctions compare inHawaii?In the Hawaiian islands, the influence ofhumans on natural environments differedbetween the two time periods. Consequently,prehistoric and historic extinction waves may havehad different causes resulting in contrastingpatterns of extinction risk (see Boyer 2008).Prehistoric extinctions showed a strong biastoward larger body sizes and flightless and ground-nesting species, even after accounting for fossilpreservation bias. Many small, specialized speciesalso disappeared, implicating a wide suite ofhuman activities including hunting anddestruction of habitat. In contrast, the highestextinction rates in the historic period were inmedium-sized nectarivorous and insectivorousbirds. Although the most vulnerable species mayhave disappeared first, changing human activitiesled to continued extinctions through time.Currently endangered species are only themost recent victims of a human-causedbiodiversity crisis that began thousands of yearsago (Steadman 1995). Despite the crucialinformation the past can provide, paleoecologicaldata are not always incorporated in studies ofextinction risk (Blackburn et al. 2004, Trevino et al.2007). To illustrate the difference this mightmake, we examined correlates of extinction riskfor Hawaiian birds using decision tree models(Boyer 2008). We compared the results of twomodels: one including only extinctions thatoccurred since European colonization of Hawaii(ca. 1800 AD), and one incorporating all knownprehistoric and historic extinctions (cumulative).The addition of older data to the model produceda substantial increase in explanatory value (7.3 Δ%DE; Fig. 3). When the recent extinctions alonewere considered, extinction predictors includedbody mass and diet, but the cumulative treeexpanded the list to include endemism andflightlessness as significant risk factors as well.While these traits may have been most importantduring the prehistoric extinction, they remainimportant for modern birds. As well as identifyingtraits associated with past extinctions, the twomodels were used to predict extinction risk forextant Hawaiian birds. Predictions from thecumulative model were a better match to currentIUCN Red List status for extant Hawaiian birdsthan predictions from the historic model, but thedifference between the two models was notsignificant (ANOVA, cumulative r²=0.15, recentr²=0.09; P >0.85; df = 4, 56; F=0.33). ExtantHawaiian birds have already been through astrong extinction filter (Pimm et al. 2006) andthese past extinctions have relevance for theconservation of the remaining species.Although human environmental impacts onbirds and their habitats have changed over time,modern endangered birds within the Pacificregion share many of the same ecologicalcharacteristics as victims of previous extinctions(Boyer 2010). It seems logical that conservationand restoration policies should incorporateFigure 3. Classification tree model of extinction riskamong Hawaiian birds. Extinction probability increases tothe right of each branch point. Terminal nodes providenumber of species and estimated probability of extinction.See text for details. %DE = percent deviance explained.32 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

Felisa A. Smith and Alison G. Boyerpaleoecological information about bird species’ranges and island ecosystems. Fossil evidencesuggests that many species currently limited to asingle island were much more widespread beforehuman contact (Steadman 2006). Thus, Steadmanand Martin (2003) argued that future extinctionsmay be partially offset by selectively translocatingbirds to islands where they once occurred. TheMarquesas Lorikeet (Vini ultramarina) and thePolynesian Megapode (Megapodius pritchardii)have been reintroduced to well-forested islands intheir former range; in New Zealand, similar islandsanctuaries have been quite successful (BirdlifeInternational 2004). Steadman and Martin (2003)provided examples of five more species that couldbenefit from such translocations. Proactiveconservation strategies present opportunities forpaleoecology to step outside its traditionallyretrospective role.Ecological history and climate changeA better understanding of ecological history canenhance our understanding of how organismsrespond to climate change in several ways. Finescaledpaleoclimate reconstructions of the lateQuaternary indicate that climate variability overthe past 100 years does not adequately representthe full range of climate changes that occur inecosystems and, further, that we tend tounderestimate the degree of climatic ‘teleconnections’(Schoonmaker and Foster 1991, MacDonaldet al. 2008). Given that many researchers haveemployed forward projected models to predictfuture climate and ecosystem responses, suchmodels parameterized based only on modernconditions are likely to be misleading (Botkin et al.2007, Williams and Jackson 2007). For someorganisms the paleorecord provides us withmultiple examples of responses to climate shiftsof varying magnitude and frequency. For example,pollen records can provide well-resolvedinformation on regional shifts in abundance, anddistributional movements of plants over the lateQuaternary (Schoonmaker and Foster 1991, Davisand Shaw 2001, Williams et al. 2001, Williams etal. 2002). Similarly, there are detailed records foranimals (Graham 1986, Graham and Grimm 1990,Graham et al. 1996, Hadly 1996). Of particularinterest is the woodrat paleomidden record,which allows fine-grained study of morphologicaladaptation to climate shifts of varying intensity.Woodrats (genus Neotoma) are smallrodent herbivores found throughout much ofNorth America. They are unique in creatingmiddens or debris piles consisting of plantfragments, fecal pellets and other materials heldtogether by evaporated urine (“amberat”). Whensheltered in a rock crevice or cave, the contentsform an indurated conglomerate, which can bepreserved for thousands of years (Betancourt etal. 1990). Microscopic identification andradiocarbon dating of the materials yieldsestimates of diet and vegetation over time.Moreover, the width of the pellets is highlycorrelated with body mass, thus allowingestimates of morphological change of populationsover time; ancient DNA can also be extracted(Smith et al. 1995, Smith and Betancourt 2006).Middens are ubiquitous across rocky arid regionsof the western United States; a well-sampledmountain region may yield upwards of 50–100discretely dated samples spanning 20,000 years ormore. Thus, paleomidden analysis yields a finegrainedcharacterization of both morphologicaland genetic responses of woodrat populations toclimate fluctuations over thousands of years.We have used the woodrat paleomiddenrecord to investigate the response to late Quaternaryclimate change over the western UnitedStates. We find that in most instances woodratsreadily adapted in situ, although there were intervalswhen temperature alterations apparently exceededspecies’ thermal tolerances (Smith andBetancourt 2006, Smith et al. 2009). Overall,woodrats follow Bergmann’s rule: within a regionthe body size of woodrat populations was largerduring cold temporal intervals and smaller duringwarmer episodes. As might be expected, sites locatednear modern range boundaries where animalsapproach physiological and ecological limitsdemonstrate more complicated responses. Atrange boundaries, elevation matters. Populationsat higher elevations adapt, while those lowelevationsites may become extirpated, dependingfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society33

Felisa A. Smith and Alison G. Boyermass; during colder episodes they were larger,and during warmer intervals, animals weresmaller (Fig. 4b). Their presence may have beentied into a much more widespread historicaldistribution of juniper (Juniperus spp.); wedocument a downward displacement of ~1,000 mrelative to juniper’s modern extent in theAmargosa Range. These results suggest a coolerand more mesic habitat association persisting forlonger and at lower elevations than previouslyreported. As climate warmed during theHolocene, N. cinerea adapted and retreatedupslope; populations were eventually completelyextirpated on the east side of Death Valley,despite the presence of what would appear to beenough high-elevation habitat (Fig. 4a; Smith et al.2009). Moreover, the range retraction of thelarger and behaviorally dominant N. cinerea led toa range expansion of N. lepida, which eventuallyreached the limits of its cold tolerance at ~1800m,an upper elevational limit maintained into moderntimes. Of particular interest is the remarkable andrapid dwarfing of body mass of N. cinereapopulations from the full glacial (~21,000 calendarybp) to the Holocene (Fig. 4b); for much of thistime, they occupied the same elevational range,but adapted to climate changes in situ. Similarpatterns are seen in other parts of the range (e.g.,Smith et al. 1995, Smith and Betancourt 2006).Note that a modern ecologist would detect thepresence of only one species (N. lepida) in thearea today, occupying an elevational range from -84 to 1800m, with a maximum body mass of~250g. Analysis of present distributions provides alimited perspective when trying to evaluate thepotential response of these species toanthropogenic warming; clearly considerable insitu evolutionary adaptation occurred along withextensive distributional/elevational migrations.Our paleomidden work highlights just howdynamic and sensitive body size and range are tothermal shifts, and suggests that both are likelyresponses to future anthropogenic climate shifts.Finding timeAs Herrera (1992) stated, “Ecologists study thintemporal slices of historically dynamic systems.”Certainly, the examples we have provided allunderscore this point. The conclusions drawn byeach study were markedly different without theincorporation of a longer temporal perspective inthe analysis. In each of these instances, the“missing perspective” was biased in a way thatcompromised the results. Flightlessness, forexample, did not come out as a factorpredisposing insular birds to extinction in ouranalysis based on modern data, because theflightless birds had already gone extinct. Yet, ifsuch an analysis was extended to predictextinction risk and direct conservation efforts inanother archipelago, the absence of flight abilityas a factor in the analysis could have verydamaging real-world results. Similarly, given thatmacroecological studies are often dependent onlarge-scale distributional information, how muchvalidity do we give studies aiming to tease apartfactors influencing the structure and function ofecological systems if major components of thesystem are missing?Our message is not novel; a number ofworkers have emphasized the importance ofincorporating a longer-term perspective inmodern ecology (Schoonmaker and Foster 1991,Herrera 1992, Delcourt and Delcourt 1998, Botkinet al. 2007). And progress is ensuing. Certainly,ecologists increasingly recognize the importanceof long-term studies; a number of important fieldprojects have now been running for many decades(e.g., Paine 1994, Brown et al. 2001). Yet, asTilman (1989) noted, non-linear dynamics andnew equilibrium states can complicateinterpretations from even the best-designed andlongest-running ecological studies. Here, we haveprovided examples where the temporal scalerequired was much longer than that achieved byany ecological study. Our purpose in doing so wasnot to criticize modern ecology, but rather toclearly illustrate why paleoecology is relevant. Byproviding concrete examples, we hope we haveclearly demonstrated the utility of incorporating ahistorical perspective. There are many resourcesavailable to do so; numerous comprehensivedatabases provide paleo distribution andabundance information for pollen, mammals, andfrontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society35

losing time?other fossils, making it possible to imbed modernecological work into a deeper context.The extent to which time matters clearlydepends on the questions ecologists ask. In somecases, a millennial-scale temporal perspective maynot be relevant. But for many of the most pressingecological issues facing society, an appreciation ofpast history is imperative. Along with earlierworkers (e.g., Schoonmaker and Foster 1991,Herrera 1992, Delcourt and Delcourt 1998, Botkinet al. 2007), we encourage better integrationbetween paleoecologists and ecologists, who areoften interested in the same questions. Effectivecommunication between these disciplines remainscomplicated by the traditional structuring ofuniversities and funding agencies into the physicaland natural sciences, which have physically andphilosophically segregated paleoecology fromecological or evolutionary disciplines. Yet,progress on some of the most topical issuesrequires that we integrate across both macro- andmicro-evolutionary and ecological theory andcombine both theoretical and empiricalperspectives. In much the same way thatunderstanding the specialized running abilities ofpronghorn antelope requires an understanding ofthe context in which they evolved, there may bemany more ecological features of extant animalsand plants that are due in some part to nowextinctcomponents of ecosystems.AcknowledgementsWe thank members of the Smith and Brown labsfor many helpful discussions; S.K. Lyons kindlyprovided data and assistance. This is contributionnumber 5 from the NSF sponsored ResearchCoordination Network: IntegratingMacroecological Pattern and Process acrossSpace. Funding provided by NSF grants BIO-0541625 and BIO-DEB-0344620 to FAS. AGB wassupported by a Smithsonian PostdoctoralFellowship and a NSF Postdoctoral ResearchFellowship in Biological Informatics (DBI-0805669).ReferencesAlcover, J.A., Sans, A. & Palmer, M. (1998) The extentof extinctions of mammals on islands. Journal ofBiogeography, 25, 913–918.Allen, B.D. & Anderson. R. Y. (1993) Evidence fromwestern North America for rapid shifts inclimate during the Last Glacial Maximum.Science, 260, 1920–1923.Alley, R.B. (2000) The Younger Dryas cold interval asviewed from central Greenland. QuaternaryScience Review, 19, 213–226.Alley, R.B., Meese, D.A., Shuman, C.A., Gow, A.J.,Taylor, K.C., Grootes, P.M., White, J.W.C., Ram,M., Waddington, E.D., Mayewski, P.A., &Zielinski, G.A. (1993) Abrupt accumulationincrease at the Younger Dryas termination in theGISP2 ice core. Nature, 362, 527–529.Bakker, V.J. & Kelt, D.A. (2000) Scale-dependentpatterns in body size distributions of neotropicalmammals. Ecology, 81, 3530–3547.Barlow, C.C. (2001) The ghosts of evolution:nonsensical fruit, missing partners, and otherecological anachronisms. Basic Books, New York.Betancourt, J.L., van Devender, T.R. & Martin, P.S.(1990) Packrat middens. The last 40,000 years ofbiotic change. University of Arizona Press,Tucson.Birdlife International. (2004) New Zealand to createmore island sanctuaries. [WWW document] URLhttp, //www.birdlife.org/news/news/2004/05/nz_island_sanctuaries.html.Blackburn, T.M., Cassey, P., Duncan, R.P., Evans, K.L. &Gaston, K.J. (2004) Avian extinction andmammalian introductions on oceanic islands.Science, 305, 1955–1958.Bond, G. & Lotti, R. (1995) Iceberg discharges into theNorth Atlantic on millennial time scales duringthe last glaciation. Science, 267, 1005–1010.Botkin, D.B., Saxe, H., Araújo, M.B., Betts, R., Bradshaw,R.H.W., Cedhagen, T., Chesson, P., Dawson, T.P.,Etterson, J.R., Faith, D.P., Ferrier, S., Guisan, A.,Hansen, A.S., Hilbert, D.W., Loehle, C., Margules,C., New, M., Sobel, M.J. & Stockwell, D.R.B.(2007) Forecasting the effects of global warmingon biodiversity. BioScience, 57, 227–236.Boyer, A.G. (2008) Extinction patterns in the avifaunaof the Hawaiian islands. Diversity andDistributions, 14, 509–517.Boyer, A.G. (2010) Consistent Ecological SelectivityThrough Time in Pacific Island Avian Extinctions.Conservation Biology, 24, 511-519.Boyer, A.G. & Jetz, W. (2010) Biogeography of BodySize in Pacific Island Birds. Ecography, 33, 369-379.36 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

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membership cornerfrom the societyYour IBS peers – An overviewOver the past 10 years, the InternationalBiogeography Society (IBS) has grown frommodest numbers to a wonderfully diverse andexpanding membership. As of February 29 th , 2012,the IBS has 634 active members. About 29% of themembership base is made up of students, and theremaining are academics, professionals and postdocs.In terms of geographic make-up, the societyis largely European (43%) and North American(38%) based but spans an increasingly widenumber of countries. Central and South Americamakes up 9% of the remaining membership, theSouth Pacific 6%, Asia 3%, and Africa 1%.One hundred fifty-three new membersjoined the IBS in 2011 – split almost exactly 50/50between student and non-student members, with3% taking advantage of the developing countryreduced rates. The majority of new memberscame from Europe (44%) and the US (29%) butincluded memberships from a total of 35 differentcountries. Since the IBS’ inception in 2001, a totalof 67 countries have been represented bymembers of the society, which is impressive. Oneof the key topics being worked on by the IBSBoard is increasing geographical diversity to makethis truly a more international society. If you havesuggestions on ways to reach out to underrepresentedareas, please drop us a line atbiogeography.ibs@gmail.com – we alwayswelcome the input!ISSN 1948-6596The IBS changed its membership feestructure in 2011, offering decreased membershipfees for people from developing nations,maintaining fee levels for students at the samelow level, and providing a tiered fee structurebased on income beyond student memberships.There have been a few issues that have come upwith some people’s renewals. If you’re due torenew during the next half-year and run into anyproblems, please don’t hesitate to contact me.And thank you for considering the new structurewhen renewing or signing up – your membershipfees help the IBS continue to be a thriving society!With the upcoming conference in Florida inJanuary of 2013, we expect to see an influx of upto 200 more members later in the year. Thanks forhelping to get the word out to your colleaguesabout the society and upcoming event – it’sshaping up to be a fantastic conference!Karen FallerIBS Manager of Membership Services; karenfaller@gmail.comEdited by Matthew HeardDid you know that any member of the IBS may raise an issue or appeal a decision of the governingBoard of Directors by placing a matter before the Board of Directors for discussion?If there is a matter you would like discussed at the next Board meeting, write to the society'sSecretary (check current list of officers at http://www.biogeography.org/).40 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

membership cornerfrom the societyUpdate on the 6 th Biennial Conference of the InternationalBiogeography SocietyPlanning is well underway for the Sixth BiennialConference of the International BiogeographySociety, which will be held at the KovensConvention Center located on Biscayne Baycampus of Florida International University inMiami Florida on January 9 th to 13 th , 2012. Januaryclimate in South Florida is mild, with temperaturesranging from 17 to 26 °C and only a small chanceof rain. The main conference hotel is locateddirectly on the beach.The preliminary program includes fourplenary symposia: (1) Predicting species and biodiversityin a warmer world: are we doing a goodjob? (2) Beyond Bergmann: new perspectives onthe biogeography of traits, (3) Islandbiogeography: new syntheses, and (4)Convergence of conservation paleontology andbiogeography. Plenary speakers will include,among others, James Brown (University of NewMexico), Jonathan Losos (Harvard University),Elizabeth Hadley (Stanford University), AntoineGuisan (University of Lausanne), and LaurenBuckley (University of North Carolina).Over 70 contributed talks will address avariety of biogeographic themes, from marinebiogeography to phylogeography. The spaciousconference center will accommodate 300 posters.Graduate students will be able to engage seniorscientists during specially organized lunch-timesessions.Pre-conference workshops (Jan 9 th ) indevelopment include half-day sessions on thetopics of (1) merging ecophysiological data intospecies distribution models, (2) popular sciencewriting, (3) advice for early-career and non-nativeEnglish speakers on preparing work forpublication, (4) Bayesian modeling, and a full-daysession on (5) biodiversity informatics.Field trips are being planned for thebeginning (Jan 9 th ) and end (Jan 13 th ) of theconference. These tours, mostly for small groupsof less than 20 people, may range from airboatrides, hikes in Everglades National Park, canoeingthe historic Oleta River, birding tour of southFlorida, sea kayaking, snorkeling, and tours of theFairchild Tropical Botanic Garden.Abstract submission and registration willopen mid-2012 1 .Daniel GavinVice President for Conferences, International BiogeographySociety. dgavin@uoregon.eduEdited by Matthew Heard1. Further details are posted at http://www.biogeography.org/html/Meetings/2013/index.html.frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society41

membership cornerfrom the membersContribute to “The Alfred Russel Wallace Page” WebsiteSeveral years ago, I solicited what I termed“commentaries” for a feature on my “AlfredRussel Wallace Page” website 1 , which includestranscriptions of Wallace publications. Almost allof Wallace’s 1000+ published works can now befound there 2 , and I am hoping that IBS membersand other parties might be interested incontributing one or more such “commentaries” tothis array. These “commentaries” typically havebeen 250 to 350 words in length, and wouldfeature your slant on why a particular Wallacearticle remains of historical and/or currentinterest. A good example may be found at the endof the page for “The Theory of Glacial Motion” 3 .It is unimportant that you may not consideryourself a “Wallace expert,” and in fact many ofthose who have contributed in the past are not. Ifyou do not wish to wade through the entire list ofWallace’s publications to choose a subject writing,I can suggest a handful of articles (many quiteshort) that might be of most interest to you.The upcoming year, 2013 is the onehundredth anniversary of Wallace’s death, andsome dozen book projects and conferences are inpreparation or planning. If you might beinterested in contributing to my site (and in turnto the anniversary celebration), please contact mefor more details at the e-mail below. Thanks verymuch for your time and attention!Charles H. Smith, FLSWestern Kentucky University, USA. charles.smith@wku.eduEdited by Matthew Heard1. http://people.wku.edu/charles.smith/index1.html2. http://people.wku.edu/charles.smith/wallace/writings.htm3. http://people.wku.edu/charles.smith/wallace/S184.htm42 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

membership cornerJob announcementsThree 2-year postdoc positionsAarhus University, DenmarkThe Department of Bioscience, AarhusUniversity invites applications for three 2-yearpostdoc positions at its new Center for InformaticsResearch on Complexity in Ecology (CIRCE, lead byprof. Jens-Christian Svenning).Ecosystems are highly complex in structureand function, and global change makes it criticalto understand how this complexity affectsecosystem responses to environmental trends andshifts. At the same time informatics-basedmethodologies (cyberinfrastructure, GIS, statisticsetc.) provide unprecedented possibilities forbreakthroughs. CIRCE studies the importance ofcomplexity for how ecosystems function andrespond to environmental change, focusing onthree major complexity factors (speciesinteractions, dispersal, and environmentalvariability) to assess their general importance,how they interact, and the mechanisms involved.CIRCE employs an informatics approach, analyzinglarge ecological data sets using advancedstatistical modeling.Postdoc project on “Ecological complexity in Arcticpopulation and ecosystem dynamics”Postdoc project on “Ecological complexity in birdcommunities across contintents”Postdoc project on “Ecological complexity in lakesacross the globe”One of the benefits open to IBS members is the opportunity to have job openings posted on theIBS blog (http://biogeography.blogspot.com/). If you have a position you would like to have advertised,please contact Karen Faller (faller@wisc.edu) or Michael Dawson(mdawson@ucmerced.edu) with details.frontiers of biogeography 4.1, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society43

membership cornerupcoming events21 st Workshop of the European VegetationSurvey (EVS)24–27 May 2012 – Vienna, Austriahttp://evs2012.vinca.at/Web of life in a changing world5 June 2012 – Montpellier, Francehttp://www.weblife.univ-montp2.fr/International Symposium on Invasive Plantsand Global Change13–17 June 2012 – Urumqi, Xinjiang, Chinahttp://www.egi.cas.cn/2012/49 th Annual Meeting of the Association forTropical Biology and Conservation18–22 June 2012 – Bonito, Mato Grosso do Sul, Brazilhttp://www.tropicalbio.org/12 th International Congress on the Zoogeographyand Ecology of Greece and AdjacentRegions18–22 June 2012 – Athens, Greecehttp://www.zoologiki.gr/12iczegar/BES Macroecology Group Inaugural MeetingWhat is Macroecology?20 June 2012 – London, UKhttp://www.britishecologicalsociety.org/meetings/VertNet biodiversity informatics trainingworkshop24–30 June 2012 – Boulder, USAhttp://vertnet.org/about/BITW.phpEvolution 2012First joint congress on evolutionary biology6–10 July 2012 – Ottawa, Canadahttp://www.confersense.ca/Evolution2012/index.htm97th ESA Annual MeetingLife on Earth: Preserving, utilizing, and sustaining ourecosystems5–10 August 2012 – Portland, USAhttp://esa.org/meetings/3 rd European Congress of Conservation BiologyConservation on the edge28 August – 1 September 2012 – Glasgow, UKhttp://www.eccb2012.org/8 th EuropeanConference on Ecological Restoration9–14 September 2012 – Česke Budějovice, Czech Republichttp://www.ecer2012.eu/NEOBIOTA 2012 — 7th Conference on BiologicalInvasionsHalting biological invasions in Europe: from data todecisions12–14 September 2012 – Pontevedra, Spainhttp://neobiota2012.blogspot.com/6th Evolutionary Biology Meeting18–21 September 2012 – Marseilles, Francehttp://sites.univ-provence.fr/evol-cgr/BES Annual Meeting 201218–20 December 2012 – Birmingham, UKhttp://www.britishecologicalsociety.org/meetings/6th International Conference of the IBS9–13 January 2013 – Florida, USAhttp://www.biogeography.org/INTECOL 2013Ecology: Into the next 100 years18–23 August 2013 – London, UKhttp://www.intecol2013.org/If you want to announce a meeting, event or job offer that could be of interest for (some) biogeographers,or you want to make a call for manuscripts or talks, please contact us atibs@mncn.csic.es and frontiersofbiogeography@gmail.com.44 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.1, 2012

table of contentsfrontiers of biogeographythe scientific magazine of the International Biogeography Societyvolume 4, issue 1 - april 2012editorialISSN 1948-6596editorial: Advancing Frontiers, with a prospective, by M.N Dawson et al. 1news and updateupdate: A cure for seeing double? Convergence and unification in biogeography and ecology, by M.N Dawson & J. Hortal 3workshop summary: The application of species distribution models in the megadiverse Neotropics poses a renewed set ofresearch questions, by L.H.Y. Kamino et al. 7symposium summary: Towards new directions and collaborations in macroecology, by C. Graham & M. Winter 11book review: Avian history is written by the winners, by D.W. Yalden 13book review: Everything changes – especially on islands, by S. Sfenthourakis 15book review: Headwaters in the clouds, by M. Richter 17books noted with interest, by M. Eichhorn 19thesis abstract: Ecology and biogeography of island parasitoid faunas, by A.M.C. Santos 20opinion and perspectivesperspective: Losing time? Incorporating a deeper temporal perspective into modern ecology, by F.A. Smith & A.G. Boyer 26membership cornerfrom the society: Your IBS peers – An overview, by K. Faller 40from the society: Update on the 6 th Biennial Conference of the International Biogeography Society, by D. Gavin 41from the members: Contribute to “The Alfred Russel Wallace Page” Website, by C.H. Smith 42Job announcements 43Upcoming events 44frontiers of biogeography copyright noticeCopyright © 2011 International Biogeography Society (IBS) under a Creative Commons Attribution Non-Commercial No Derivatives (CCANCND)license. All rights reserved. It is strictly forbidden to alter the journal contents in any manner without the express written permission of the IBS. Itis also strictly forbidden to make copies of whole issues of this journal for any commercial purpose without the express written permission of theIBS. The IBS holds the right for the passive distribution (i.e. through its publication on the Internet) of any part or the whole issue of the journalduring one year after its publication. Such right is explicitly transferred to eScholarship trhough an agreement with the journal and the IBS. Anyactive distribution of any part or the whole issue of the journal is explicitly permitted since the date of publication, and any passive distribution isexplicitly permitted after one year of the date of publication. Any individual and/or institution can download, read and/or print a copy of anyarticle or the whole journal for non-commercial educational or non-commercial research purposes at any time. This includes an express permissionto use articles for non-commercial educational purposes by making any number of copies for course packs or course reserve collections.Academic institutions/libraries may also store copies of articles and loan them to third parties. All copies of articles must preserve their copyrightnotice without modification. All articles are copyrighted by their authors under a universal Creative Commons Attribute License (CCAL), which isalso explicitly included in the agreement made betwen the journal and eScholarship. This license permits unrestricted use, distribution, and reproductionin any medium, provided the original author and source are credited. All authors endorse, permit and license the IBS to grant any thirdparty the copying and use privileges specified above without additional consideration or payment to them or to the IBS. These endorsements, inwriting, are on file in the office of the IBS. Consult authors for permission to use any portion of their work in derivative works, compilations or todistribute their work in any commercial manner.From the IBS constitution: "Bylaw 10. Publications. All titles, copyrights, royalties or similar interests in tape recordings, books or other materialsprepared for the International Biogeography Society Inc activities will be held solely by the International Biogeography Society Inc and in the nameof the International Biogeography Society Inc.". And "Article 8. Publications. The publications of the Society shall include journals, newsletters, andsuch other publications as the Governing Board of Directors may authorize."We gratefully acknowledge Evolutionary Ecology, Ltd. and Mike Rosenzweig in particular for the advice on copyright matters.