Frontiers of Biogeography Vol

frontiers of biogeography vol. 3, nº 3 ‐ november 2011

the scientific magazine of the International Biogeography Society

ISSN 1948‐6596 – freely available at http://www.biogeography.org/

frontiers of biogeography the scientific magazine of the International Biogeography Society volume 3, issue 3 ‐ November 2011 ISSN 1948‐6596 frontiers of biogeography is published by the International Biogeography Society (IBS), an international and interdisciplinary society contributing to the advancement of all studies of the geography of nature frontiers of biogeography is available online at the IBS website: http://www.biogeography.org/html/fb.html frontiers of biogeography aims to be a forum for biogeographers and a way to disseminate research in biogeography to the general public; our scope includes opinions, perspectives, and reviews, symposia proceedings, letters to the editor, book reviews, research upda‐ tes, interviews, and articles on how to teach, disseminate and/or apply biogeographical knowledge. Letters to the editor and symposium proceedings may include novel analyses of original datasets (see editorial instructions). Manuscripts should be submitted to frontiersof‐ [email protected]. Editorial enquiries should be made to the Editor‐in‐Chief at [email protected]. frontiers of biogeography uses a publication agreement based on the Creative Commons scheme to ensure that the authors retain full intellectual property (IP) rights on their work, and that this is freely available for any non‐commercial use. Under this agreement, the IBS retains 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 distribute their work (including the original PDFs) actively from the day of publication, and passively from one year after (see the full license infor‐ mation at the end of the issue). you can find information about the International Biogeography Society at http://www.biogeography.org/; for the latest job announce‐ ments and other news please visit also the IBS blog (http://biogeography.blogspot.com/), and the IBS facebook group (http:// www.facebook.com/group.php?gid=6908354463). editorial board editor‐in‐chief: deputy editors‐in‐chief: Joaquín Hortal – Museo Nacional de Ciencias Naturales (CSIC), Michael N Dawson – University of California, Merced, USA Spain and Universidade Federal de Goiás, Brazil Richard Field – University of Nottingham, UK associate editors: editorial assistant: Antje Ahrends – Royal Botanic Garden Edinburgh, UK Lauren Schiebelhut – University of California, Merced, USA Jan Beck – University of Basel, Switzerland Jessica Blois – University of Wisconsin, Madison, USA advisory board: Chris Burridge – University of Tasmania, Australia Miguel B. Araújo – Museo Nacional de Ciencias Naturales (CSIC), Marcus V. Cianciaruso – Universidade Federal de Goiás, Brazil Spain and Universidade de Évora, Portugal Markus Eichhorn – University of Nottingham, UK Lawrence R. Heaney – Field Museum of Natural History, Chicago, Roy Erkens – Universiteit Utrecht, The Netherlands USA Camilla Fløjgaard – Aarhus University, Denmark David G. Jenkins – University of Central Florida, Orlando, USA Dan Gavin – University of Oregon, USA Richard Ladle – Universidade Federal de Alagoas, Brazil and Oxford Matthew J. Heard – Brown University, USA University, UK David G. Jenkins – University of Central Florida, Orlando, USA Mark V. Lomolino – State University of New York, USA Frank A. La Sorte – Cornell lab of Ornithology, USA IBS V. P. for Public Affairs & Communications Richard Ladle – Universidade Federal de Alagoas, Brazil and Oxford University, UK Richard Pearson – American Museum of Natural History, USA Thiago F. Rangel – Universidade Federal de Goiás, Brazil Willem Renema – NCB Naturalis, The Netherlands Núria Roura‐Pascual – Universitat de Girona and Centre Tecnològic Forestal de Catalunya, Spain Spyros Sfenthourakis – University of Patras, Greece

International Biogeography Society officers 2011‐2012

President: Lawrence R. Heaney First Past President: James H. Brown President Elect: Rosemary Gillespie Second Past President: Mark V. Lomolino VP for Conferences: Daniel Gavin Third Past President: Brett R. Riddle VP for Public Affairs & Communications: Michael N Dawson Fourth Past President: Vicki Funk VP for Development & Awards: George Stevens Fifth Past President: Robert J. Whittaker Secretary: Richard Field Treasurer: Lois F. Alexander Director‐at‐large: Catherine Graham Upcoming meeting host (ex officio): Kenneth Feeley Director‐at‐large: Kathy Willis Past Graduate student representative (ex officio): Matthew Heard Student‐at‐large: Ana M. C. Santos

cover: Flowering red buglosses (Echium wildpretii, also named tajinastes rojos in Spanish) in front of Mount Teide (Tenerife, Canary Islands). Photograph by Ana M. C. Santos. news and update ISSN 1948‐6596 update Species–area curves and the estimation of extinction rates

The species–area relationship (SAR) is one of the not surprising that SAR‐based estimates of extinc‐ longest‐known, most intuitive and empirically best tion have been welcome despite critical studies ‐proven patterns of biodiversity (Arrhenius 1921). that often found lower extinction rates than pre‐ Various authors determined theoretically that the dicted (e.g., Kinzig & Harte 2000). It was argued, SAR can be approximated as a power‐law function reasonably, that on top of imminent extinction in (i.e., S = cAz where S is species richness, A is area some species, others will be doomed to future and c and z are constants; Preston 1962, May extinction because of reductions in their popula‐ 1975, Harte et al. 1999), with z ≈ 0.25 in continen‐ tion size, and that this ‘extinction debt’ explains tal areas but higher when dispersal barriers are apparent misfits. Other sources of uncertainty of involved (e.g., ‘island species–area relationship’). the SAR‐based estimates are the (often false) as‐ Empirical data suggested lower z in continental sumption of a completely inhospitable matrix be‐ areas (0.13‐0.18) and values up to 0.35 for island tween remaining habitat patches (Koh & Ghazoul systems (Rosenzweig 1995). Dengler (2009) re‐ 2010) or the use of default slope values (z) in the cently came to the conclusion that the power law absence of system‐specific fitted data. fits empirical data best in most cases (see also He & Hubbell (2011) pointed out that a Dengler & Odeland 2010). Various authors ob‐ backward interpolation of SARs is a flawed con‐ served further systematic variations of z, such as cept of measuring extinction rates (see also Kinzig when considering spatial scale or sampling design & Harte 2000). This is because the area gain (Plotkin et al. 2001, Scheiner 2006, Tjørve 2006, needed to encounter the first individual of a new Dengler 2009). Kinzig & Harte (2000) pointed out species (which shapes the SAR) is always smaller the difference between SAR and the endemics– than the area loss needed to remove the last indi‐ area curve (EAR), which considers only species vidual. To show this, they formulated both as spa‐ endemic to a part of the region under analysis. So tially explicit sampling processes (SAR for first en‐ what could He & Hubbell (2011) report that was counters, EAR for last encounters). They con‐ so novel and generally relevant about SARs to cluded that SAR‐derived estimates of imminent merit recent publication in Nature? extinction will always be too high, unless individu‐ Since area seems always to affect biodiver‐ als are randomly distributed (i.e., no aggregated sity, no matter what taxon, system or scale, SARs occurrence of individuals within a species), which have frequently been used to estimate species is an unrealistic assumption. He & Hubbell (2011) richness loss resulting from anthropogenic habitat also showed that the EAR is a good predictor of destruction, i.e. extinction rates in a conservation empirical extinction rates even if no spatial aggre‐ context. The loss of a certain amount of area leads gation is modelled, which offers an alternative to fewer species existing in a region – at least (but a more challenging one) for estimating imme‐ some regional extinctions occur – and the shape diate extinction of endemics from area loss. of the SAR has typically been used to retrieve He & Hubbell (2011) clearly acknowledged quantitative estimates of how many species will that there is an anthropogenic extinction crisis go (regionally) extinct. and that habitat loss causes extinction. Further‐ Providing empirical evidence for the extinc‐ more, they did not claim that small population tion of a species is challenging and estimating ex‐ sizes of remaining species could not lead to fur‐ tinction rates across a community even more so ther, lagged extinction (in He & Hubbell’s view, (Ladle et al. 2011, this issue). Yet this is needed for EARs model only imminent extinction – and so do many conservation applications, such as schemes SARs, but wrongly). Despite this, He & Hubbell for offsetting biodiversity loss (Curran et al. 2011) (2011) already anticipated that pointing out this or, not least, for political argument. It is therefore error in estimating extinctions would not be frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 81 news and update greeted with enthusiasm among conservationists, Jan Beck and the correspondence on the paper (Evans et al. University of Basel, Dept. Environmental Science 2011, Brooks 2011; see also online comments at (Biogeography section), Basel, Switzerland. http://www.nature.com/nature/journal/v474/ e‐mail: [email protected]; n7351/full/474284b.html) seems to confirm that. http://www.biogeography.unibas.ch/beck The paper is viewed as irresponsibly undermining conservation efforts by allowing anti‐conservation References groups to claim that things are not as bad as previ‐ Arrhenius, O. (1921) Species and area. Journal of Ecol‐ ously asserted (fossil fuel lobbying in the climate ogy, 9, 95–99. change discussion is cited as example of this tac‐ Brooks, T.M. (2011) Extinctions: consider all species. tic). Conserving nature is not only about science, Nature, 474, 284. but it is to a large degree politics – and correcting Curran, M., De Baan, L., de Schryver, A.M., van Zelm, an error leads to better science but might weaken R., Hellweg, S., Koellner, T., Sonnemann, G. & Huijbregts, M.A.J. (2011) Toward meaningful political success. I think scientists must correct end points of biodiversity in life cycle assess‐ themselves and not hold on to preconceived ment. Environmental Science and Technology, ideas, even if it creates such dilemmas. 45, 70–79. However, He & Hubbell (2011) studied area Dengler, J. (2009) Which function describes the species effects as a sampling problem in continental re‐ –area relationship best? A review and empirical evaluation. Journal of Biogeography, 36, 728– gions, which is probably appropriate for capturing 744. immediate extinction in many conservation set‐ Dengler, J. & Oldeland, J. (2010) Effects of sampling tings which occur at the regional or landscape protocol on the shapes of species richness scale. It remains to be understood and tested curves. Journal of Biogeography, 37, 1698–1705. whether their conclusions – that (a) EAR estimates Evans, M., Possingham, H. & Wilson, K. (2011) Extinc‐ extinction better than SAR (cf. Kinzig & Harte tions: conserve not collate. Nature, 474, 284. Harte, J., Kinzig, A. & Green, J. (1999) Self‐similarity in 2000, Pereira et al. 2012) and (b) z differs system‐ the distribution and abundance of species. Sci‐ atically between SAR and EAR (which is presented ence, 284, 334–336. confusingly) – are generalities. Thus it remains to He, F. & Hubbell, S.P. (2011) Species–area relationships be seen whether SARs always overestimate ex‐ always overestimate extinction rates from habi‐ tinction, as He and Hubbell (2011) claimed. A fur‐ tat loss. Nature, 473, 368–371. ther task will be to quantitatively estimate how Kinzig, A. & Harte, J. (2000) Implications of endemics– area relationships for estimates of species ex‐ many more species may go extinct after a time tinctions. Ecology, 81, 3305–3311. lag: how large the extinction debt really is (see Koh, L.P. & Ghazoul, J. (2010) A matrix‐calibrated spe‐ also Pereira et al., in press). In this context, it may cies–area model for predicting biodiversity be worthwhile to thoroughly investigate under losses due to land‐use change. Conservation which circumstances, if any, the consequences of Biology, 24, 994–1001. Ladle, T.J., Jepson, P., Malhado, A.C.M., Jennings, S. & area lost to habitat destruction could be under‐ Barua, M. (2011) The causes and biogeographi‐ stood solely on the basis of island biogeographic cal significance of species’ rediscovery. Frontiers mechanisms (Rosenzweig 2001) – that is, species of Biogeography, 3, 111–118. richness as equilibrium between immigration + May, R.M. (1975) Patterns of species abundance and speciation and extinction. The spatial and tempo‐ distribution. In Cody M.C. & Diamond J.M. ral scales of analysis, among other factors, may be (eds.), Ecology and evolution of communities, pp. 81–120; Belknap Press, Cambridge (Mass.). relevant for this. Under such circumstances, SARs Pereira, H.M., Borda‐de‐Agua, L. & Martins, I.S. (2012) may estimate the new equilibrium state, account‐ Geometry and scale in species–area relation‐ ing for imminent and time‐lagged extinctions. ships. Nature, in press. Plotkin, J.B., Potts, M.D., Yu, D.W., et al. (2000) Predict‐ ing species diversity in tropical forests. Proceed‐ ings of the National Academy of Sciences USA, 97, 10850–10854.

Preston, F.W. (1962) The canonical distribution of com‐ Scheiner, S.M. (2003) Six types of species–area curves. monness and rarity: Part I. Ecology, 43, 185– Global Ecology and Biogeography, 12, 441–447. 215. Tjørve, E. (2006) Shapes and functions of species–area Rosenzweig, M.L. (1995) Species diversity in space and curves: a review of possible models. Journal of time. Cambridge University Press, Cambridge. Biogeography, 30, 827–835. Rosenzweig, M.L. (2001) Loss of speciation rate will impoverish future diversity. Proceedings of the Edited by Joaquín Hortal National Academy of Sciences USA, 89, 5404– 5410. update Extinct or extant? Woodpeckers and rhinoceros

Biogeographical research needs accurate data on statistics to model changes in the rate of valid the distribution of species. For many species this is sightings and to assess the quality of uncertain exceedingly difficult to obtain, leading to a lack of sightings for the ivory‐billed woodpecker global information collectively known as the Wal‐ (Campephilus principalis) in North America. The lacean shortfall. Fortunately, new tools are being woodpecker was controversially rediscovered in developed that allow conservationists and bio‐ 2005, but a lack of clear documentary evidence geographers to determine the existence of extant and the failure of subsequent intensive surveys populations with much greater accuracy. have led many scientists to doubt the veracity of Foremost among these new tools is the in‐ this claim. The Bayesian model applied by Solow creasing use of genetic analysis. This was recently to 68 historical sightings (29 of which were classi‐ used to great effect to confirm the extinction of fied as uncertain) strongly suggests that the bird is the Javan rhinoceros (Rhinoceros sondaicus anna‐ indeed extinct, and the 2005 sighting was sadly a miticus) in Cat Tien National Park in Vietnam case of mistaken identity. (Brook et al. 2011). Despite their enormous size, Javan rhinoceros are remarkably shy forest‐ Richard Ladle dwelling animals that are difficult to see under Federal University of Alagoas, Institute of Biological natural conditions and were only rediscovered in Sciences and Health, Brazil and Oxford University, mainland Asia in 1988. Given the difficulty of tra‐ School of Geography and the Environment, UK. ditional surveying techniques, scientists from e‐mail: [email protected]; WWF and the Cat Tien National park had been http://www.geog.ox.ac.uk/staff/rladle.html monitoring the population by conducting genetic analysis of dung samples collected in the park be‐ tween 2009 and 2010. The analysis indicated that References all the dung belonged to a single individual, the Brook, S., de Groot, P.V.C., Mahood, S. & Long, B. (2011) Extinction of the Javan Rhinoceros body of which was found April 2010, thereby con‐ (Rhinoceros sondaicus) from Vietnam. WWF firming the extinction of the population. Report. Available at: http:// Of course, genetic analysis is costly, time www.worldwildlife.org/who/media/press/2011/ consuming and requires some form of biological WWFBinaryitem24584.pdf tissue (hair, dung, etc.). For many rare animals the Solow, A., Smith, W., Burgman, M., Rout, T., Wintle, B. and Roberts, D. (2011), Uncertain sightings and only information that exists is the occasional sight‐ the extinction of the ivory‐billed woodpecker. ing, the reliability of which is often highly ques‐ Conservation Biology. doi: 10.1111/j.1523‐ tionable. Andrew Solow and his colleagues have 1739.2011.01743.x recently come up with an ingenious method to account for this inevitable uncertainty (Solow et Edited by Joaquín Hortal al. 2011). They use Bayesian (probability‐based) frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 83 news and update ISSN 1948‐6596 update Climate wars

Links between climate and societal instability, and diseases) seem plausible, but psychological conflict and war have increasingly been suggested effects of unusual weather conditions on a large and analyzed (Diamond 2005), thereby fusing tra‐ number of individuals may also increase a soci‐ ditionally distinct academic disciplines such as ety’s conflict potential. (bio‐)geography, (agro‐)ecology and economics, Zhang et al. (2011) presented a detailed history and peace research. Studies exploring causality analysis based on a time series of cli‐ these relationships are particularly pertinent in matic fluctuations over a 300 year period in pre‐ times of anthropogenic climate change. industrial Europe. They provide strong support for Recent research has provided quantitative the idea that climatic variation caused fluctuations support for such climate–culture linkages, but in agricultural productivity, and hence food avail‐ most of these studies have either been based on ability and prices. The latter was identified as the correlative evidence (e.g., Zhang et al. 2007), ana‐ root cause for a number of societal phenomena lyzed short‐term climate fluctuations (e.g., Burke such as migrations, epidemics, population growth et al. 2009) or addressed specific hypotheses on and war. A temperature‐based model based on the causes of human conflict (Beck and Sieber these mechanisms could successfully predict peri‐ 2010). However, in order to make conflict predic‐ ods of crisis and harmony for past eras with less‐ tions under climate‐change scenarios reliable and detailed historical records. to engage in conflict prevention or mitigation, it is An important future direction of research in important to be certain about causal relationships this field will certainly be the identification of and to fully understand the mechanistic links be‐ natural factors and societal traits that explain tween past climatic changes and historical con‐ variation around such climate‐determined pat‐ flicts. Two new studies have attempted this. terns. Demography and economic performance Hsiang et al. (2011) made use of the recur‐ have sometimes been analyzed in this context ring yet irregular El Niño Southern Oscillation (Samson et al. 2011, Hsiang et al. 2011). However, (ENSO) climatic changes as a natural experiment. it will require the further integration of the above‐ This allowed them to show, on a global scale and mentioned disciplines to sort out the ultimate for a time period of more than half a century, that causes of why certain regions and/or societies (within the same localities and societies) civil con‐ navigated smoother and less violent routes flicts were more likely to arise during El Niño through times of crisis than others (my current events as compared to La Niña periods. Further‐ location, Switzerland, is a prime example within more, no such effect was observed for countries the last few centuries). outside the ENSO‐affected zone of the world. This provides strong evidence that climate is indeed causal to these events. However, the authors can Jan Beck only speculate on a variety of mechanisms for University of Basel, Dept. Environmental Science how (warmer and drier) El Niño periods could lead (Biogeography section), Basel, Switzerland. to conflict. Effects mediated by decreased agricul‐ e‐mail: [email protected]; tural productivity and/or economic disturbance http://www.biogeography.unibas.ch/beck (e.g., resulting from increases in natural disasters Your participation in frontiers of biogeography is encouraged. Please send us your articles, com‐ ments and/or reviews, as well as pictures, drawings and/or cartoons. We are also open to sug‐ gestions on content and/or structure. Please check http://www.biogeography.org/html/fb.html for more information, or contact us at [email protected] and [email protected].

84 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update References Zhang, D.D., Lee, H.F., Wang, C., Lie, B., Pei, Q., Zhang, J. & An, Y. (2011) The causality analysis of cli‐ Beck J., & Sieber, A. (2010) Is the spatial distribution of mankind’s most basic economic traits deter‐ mate change and large‐scale human crisis. Pro‐ ceedings of the National Academy of Sciences mined by climate and soil alone? PLoS ONE 5(5): e10416. USA, 108, 17296–17301. Burke, M., Miguel, E., Satyanath, S., Dykema, J. & Lo‐ Zhang, D.D., Brecke, P., Lee, H.F., He, Y.‐Q. & Zhang, J. bell, D. (2009) Warming increases risk of civil (2007) Global climate change, war and popula‐ tion decline in recent human history. Proceed‐ war in Africa. Proceedings of the National Acad‐ emy of Sciences USA, 106, 20670–20674. ings of the National Academy of Sciences USA, 104, 19214–19219. Diamond, J. (2005) Collapse: how societies choose to fail or succeed. Viking. Hsiang, S.M., Meng, K.C. & Cane, M.A. (2011) Civil con‐ Edited by Richard Ladle flicts are associated with the global climate. Na‐ ture, 476, 438–411. Samson, J., Berteaux, D., McGill, B.J., Humphries, M.M. (2011) Geographic disparities and moral hazards in the predicted impacts of climate change on human populations. Global Ecology and Bio‐ geography, 20, 532–544. update Emerging research opportunities in global urban ecology

Biogeographers have examined how human activi‐ Urban areas typically contain spatially heteroge‐ ties have affected patterns of biological diversity neous collections of native and non‐native species from a variety of perspectives, with special atten‐ (McKinney 2008); these unique assemblages can tion often given to oceanic islands. With the cur‐ be examined based on their compositional rent accelerating pace of environmental change, (Niemelä et al. 2002) and phylogenetic structures these effects are increasingly evident at global (Ricotta et al. 2009). Three nested sampling ap‐ scales. Human industry, commerce, agriculture proaches are currently used to investigate urban and transportation all have the potential now to systems at broad spatial scales: urban plots or affect natural systems globally through an assort‐ transects, the entire urban matrix and the urban ment of drivers; primary among these are land‐ matrix embedded within a regional context use change, species introductions and climate (Werner 2011). Each sampling approach provides change. a unique inferential basis, although the third al‐ Human activities and their consequences lows for more refined interpretation, controlling come to a unique focus in urban areas, an expand‐ for regional differences. ing form of land use that is attracting increasing A recent study in Global Ecology and Bio‐ research attention from ecologists (Grimm et al. geography adopts a novel perspective and exam‐ 2008). Urban areas contain similar environmental ines how avian assemblages sampled within plots conditions worldwide and act as a focal point for of intact vegetation in urban and semi‐natural ar‐ species introductions and extinctions. These hu‐ eas differ based on several common mac‐ man‐dominated environments offer unique op‐ roecological relationships. Pautasso et al. (2011) portunities to investigate the broad‐scale dynam‐ compiled data on species composition and abun‐ ics of human‐mediated biotic interchange (La dance from all around the globe, although the Sorte et al. 2007), its consequences for β diversity majority of the samples are from Europe and (La Sorte et al. 2008) and the regional factors and North America. A primary finding of the study was biological traits associated with native species ex‐ a lack of evidence for differences in the species– tinctions (Hahs et al. 2009, Duncan et al. 2011). area, species–abundance or species–biomass rela‐ frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 85 news and update tionships between urban and semi‐natural locali‐ Frank A. La Sorte ties. The number of exotic bird species in urban Cornell Lab of Ornithology, Ithaca, NY, USA. areas is low, suggesting that these relationships e‐mail: [email protected]; are defined primarily by native species in both http://www.birds.cornell.edu/ environments. These findings highlight the impor‐ tance of maintaining intact vegetation within ur‐ References ban landscapes and the role of urban diversity as a Duncan, R.P., Clemants, S.E., Corlett, R.T., Hahs, A.K., tool for promoting conservation initiatives and McCarthy, M.A., McDonnell, M.J., Schwartz, biological awareness, as emphasized in many ur‐ M.W., Thompson, K., Vesk, P.A. & Williams, ban‐ecology studies. Nevertheless, the findings N.S.G. (2011) Plant traits and extinction in urban from Pautasso et al. (2011) contrast with current areas: a meta‐analysis of 11 cities. Global Ecol‐ expectations on how urbanization affects patterns ogy and Biogeography, 20, 509–519. Grimm, N.B., Faeth, S.H., Golubiewski, N.E., Redman, of diversity, and should be a motivating factor in C.L., Wu, J., Bai, X. & Briggs, J.M. (2008) Global promoting further research. The increasing preva‐ change and the ecology of cities. Science, 319, lence and quality of global data sources provides 756–760. an exciting basis to examine the structure and de‐ Hahs, A.K., McDonnell, M.J., McCarthy, M.A.,et al. terminants of these macroecological relationships (2009) A global synthesis of plant extinction across more refined temporal, spatial and anthro‐ rates in urban areas. Ecology Letters, 12, 1165– 1173. pogenic gradients. La Sorte, F.A., McKinney, M.L. & Pyšek, P. (2007) Com‐ By taking a global perspective, novel in‐ positional similarity among urban floras within sights can be gained on the unique position urban and across continents: biogeographical conse‐ areas have, both as a source for global change and quences of human‐mediated biotic interchange. Global Change Biology, 13, 913–921. as regions capable of maintaining important as‐ La Sorte, F.A., McKinney, M.L., Pyšek, P., Klotz, S., Rap‐ pects of biological diversity. Global comparative son, G.L., Celesti‐Grapow, L. & Thompson, K. studies also have the potential to bolster and re‐ (2008) Distance decay in similarity among Euro‐ fine current recommendations about how to pean urban floras: the impacts of anthropogenic maintain biological diversity within human‐ activities on β diversity. Global Ecology and Bio‐ dominated landscapes. Specifically, the preserva‐ geography, 17, 363–371. McKinney, M.L. (2008) Effects of urbanization on spe‐ tion or restoration of patches of intact vegetation cies richness: a review of plants and animals. within urban areas is as valuable in maintaining Urban Ecosystems, 11, 161–176. basic macroecological patterns of avian diversity Niemelä, J., Kotze, D.J., Venn, S., Penev, L., Stoyanov, I., as conducting these activities outside urban areas. Spence, J., Hartley, D. & Montes de Oca, E. Importantly, this work takes the focus away from (2002) Carabid beetle assemblages (Coleoptera, Carabidae) across urban‐rural gradients: an in‐ Europe and North America, where the vast major‐ ternational comparison. Landscape Ecology, 17, ity of the research has been conducted, allowing 387–401. for a more inclusive set of inferences and recom‐ Pautasso, M., Böhning‐Gaese, K., Clergeau, P., et al. mendations. Urban data are becoming increas‐ (2011) Global macroecology of bird assemblages ingly available through remote sensing activities, in urbanized and semi‐natural ecosystems. Global Ecology and Biogeography, 20, 426–436. citizen science initiatives and broader collabora‐ Ricotta, C., La Sorte, F.A., Pyšek, P., Rapson, G.L., Celesti tive efforts. Exploring how anthropogenic activi‐ ‐Grapow, L. & Thompson, K. (2009) Phyloecol‐ ties are impacting natural systems globally is criti‐ ogy of urban alien floras. Journal of Ecology, 97, cal in supporting a truly comprehensive under‐ 1243–1251. standing of the current dynamics and long‐term Werner, P. (2011) The ecology of urban areas and their consequences of global environmental change. functions for species diversity. Landscape and Ecological Engineering, 7, 231–240.

Edited by Joaquín Hortal

86 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update update Beyond taxonomical space: large‐scale ecology meets func‐ tional and phylogenetic diversity

Community ecology traditionally focuses on hypo‐ FD, which initially was considered the holy grail of thetical‐deductive and experimental approaches the biodiversity‐ecosystem functioning agenda, and often is criticized for narrowing our under‐ also was rapidly applied as a metric for investigat‐ standing of nature to local idiosyncrasies, ignoring ing assembly rules (see Pavoine & Bonsall 2011). the importance of historical explanations. On the How could macroecology and biogeography bene‐ other hand, approaches taken by macroecologists fit from these two approaches? The answer lies in and biogeographers have been excessively ex‐ understanding what FD and PD should represent ploratory and correlative, with limited success in and how they relate to each other: while phyloge‐ elucidating the mechanisms responsible for many netic community ecology links evolutionary and of the large‐scale patterns we observe in nature biogeographic history to present‐day ecology, (see Gaston & Blackburn 1999, Ricklefs 2008 and functional diversity (as any trait‐based approach) references therein). Recognizing that both ap‐ links niche theory to large‐scale approaches, such proaches can learn from each other is pivotal in as macroecology, biogeography or phylogeogra‐ the challenge of integrating data from different phy. Therefore, combining ecological and phyloge‐ scales in order to unravel the ecological and evo‐ netic frameworks to explain large scale patterns of lutionary mechanisms that influence current pat‐ biodiversity is an important step, taken recently. terns in biodiversity and ecosystem functioning. Large‐scale studies involving PD and FD seems to Species richness has been the most com‐ be increasing at similar rates (Fig.1). Recently, it mon metric used to represent all aspects of bio‐ was shown that both measures can be decom‐ logical diversity (from genetic and taxonomic to posed into gamma (regional), alpha (local) and phenetic diversity). However, species richness beta (turnover) components. Whereas large‐scale alone cannot describe the processes involved in studies and any‐scale studies follows a similar species coexistence and ecosystem functioning trend for beta‐PD, there were few studies with and also does not describe properly the differ‐ beta‐FD (none at large‐scale). This is perhaps be‐ ences in community structure. In contrast, phy‐ cause biogeographers and macroecologists were logenetic and functional diversities allow us to more aware of evolutionary and historical hy‐ understand the relative importance of species potheses, so the conceptual framework of beta‐ composition in terms of evolutionary history and PD was likely to be absorbed first. Also, this could ecological similarities. Phylogenetic diversity (PD) reflect the assumption that closely related species is a biodiversity measure that accounts for the should be ecologically more similar than distant phylogenetic relationship (hence evolutionary his‐ related species and, thus, PD should be a good tory) among species, whereas functional diversity surrogate for FD (in fact this is what most large (FD) represents how species are distributed in a and local‐scale PD studies used to assume). This multidimensional niche space defined by ecologi‐ traditional assumption is now debated (e.g. Losos cal traits. 2008), and these two measures may be viewed as Phylogenetic and functional approaches to complementary, rather than competing, ap‐ community ecology emerged as prominent fields proaches (Gómez et al. 2010, Diniz‐Filho et al. of research in the last decade (Fig. 1), but some‐ 2011, Meynard et al. 2011, Pavoine & Bonsall how independently and without much crossover 2011, Safi et al. 2011). in the first years. Early PD measures were pro‐ While some large‐scale studies involving PD posed as a tool to select conservation areas, but and FD are exploratory (e.g. Meynard et al. 2011) later the idea was extended to understand how others have presented hypotheses and predic‐ communities are assembled from a regional pool. tions. Safi et al. (2011) investigated global pat‐ frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 87 news and update terns of mammal PD and FD and found that when neity could also result in an overdispersion in FD controlling mammal assemblages for their evolu‐ because coexisting species could adapt and spe‐ tionary history the tropics were characterized by a cialize to the different environmental conditions. FD deficit. This suggests that more species can be Some light has been shed on beta‐PD pat‐ closely packed into the ecological space in tropical terns by Gómez et al. (2010), studying Neotropical than in temperate regions (see figure 3 in their Forest antbirds at different spatial scales. If speci‐ paper), a paradoxical situation in which competi‐ ation occurred mainly among ecoregions, there is tion seems to limit trait evolution in a group, but a lower probability of sister species co‐occurring does not decrease the co‐occurrence of species in the same ecoregion, resulting in phylogenetic with similar trait values (Wiens 2011). There are evenness at this smaller scale. If so, we would ex‐ several non‐mutually exclusive mechanisms that pect high species turnover (taxonomic beta diver‐ could be responsible for this pattern (see Figure 1 sity) and low phylogenetic turnover (beta‐PD) in Safi et al. 2011). In temperate regions, for ex‐ among ecoregions, because species would tend to ample, if resources are limited, species need to be close relatives. An alternative scenario is when occupy wider ecological niches in order to secure phylogenetic structure at the regional scale is a their energy demands and therefore communities product of limited dispersal of lineages. In this would show signs of overdispersion in functional case we would expect both high species turnover traits. In addition, high environmental heteroge‐ and high beta‐PD among regions, because each

200 6 180 Any spatial scale 4 160 FD

140 2 PD

beta-FD 120 0 beta-PD 2007 2008 2009 2010 2011 100

80 published studies Large spatial scale 60

FD 40 PD 20 bet a-P D

0 1975 1980 1985 1990 1995 2000 2005 2010 year Figure 1. The number of articles published in peer‐reviewed journals indexed by ISI with functional and phylogenetic diversity in the title, abstract or key‐words from 1976 to 2010. Any spatial scale means all studies published in all sub ‐disciplines of ecology and evolutionary biology, irrespectively of scale. Large spatial scale are those studies con‐ strained by the search expression Topic=(geograph* OR macroecol* OR biogeogr*), that is, those studies most likely to be related to macroecology and biogeography. FD = any study with topic “functional diversity”; PD = any study with topic “phylogenetic diversity”; beta‐FD = any study with topic “functional beta diversity” or “functional turn‐ over”; beta‐PD = any study with topic “phylogenetic diversity” or “phylogenetic turnover”. The inset is provided to show currently starting publication trends concerning beta‐PD and beta‐FD. There was no large‐scale study involving beta‐FD up to 2010; but a few were published in 2011 or are in press.

88 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 news and update region would contain distinct clades, with inde‐ proposed, each year! This may come at the ex‐ pendent diversifications. Finally, if observed val‐ penses of the more important (and exciting) steps ues of species turnover and beta‐PD do not differ of doing science: how can we move forward the from what would be expected by chance (using theory by using novel approaches? null‐models where random assemblages are built All existing hypotheses that have been ap‐ from the species pool), phylogenetic structure at plied to taxonomic diversity can be extended to the regional scale is unlikely to be the result of phylogenetic and functional diversity (Meynard et historical processes. In that case using FD should al. 2011). However, PD and FD can be used to cre‐ be better because niche‐based processes are ate more rigorous and direct predictions for most more likely to explain the pattern. For example, of the hypotheses in macroecology and biogeog‐ along a strong environmental gradient where spe‐ raphy, such as attempts to explain latitudinal pat‐ cies are sorted from the regional pool according to terns of biodiversity (Willig et al. 2003). These their traits, we expect both species and functional metrics also present an opportunity to formulate turnover. However, if the species pool is com‐ new hypotheses about how species evolutionary posed of ecologically similar species – an indica‐ history and trait diversity are distributed across tion that species were sorted according to their communities at different scales. For example, traits at a higher spatial scale (for example, due to Wiens et al. (2011) showed situations where after a climatic filter or historical processes) – we a major evolutionary radiation within a region, the should expect low functional turnover because region can still be invaded by ecologically similar the pool already contains very similar species. species from another clade, challenging the para‐ Also, in the absence of environmental filters, spe‐ digm that communities are ‘saturated’. Large‐ cies turnover should occur independently of func‐ scale phylogenies and trait databases are cur‐ tional turnover (Mouchet et al. 2010). Neverthe‐ rently becoming available for a wide range of less, species traits should have – at least to some taxonomic groups, facilitating estimates of FD and extent – some phylogenetic signal and, therefore, PD. Including these two aspects of biological di‐ partitioning the relative contribution of evolution‐ versity will be crucial if we want to advance from ary history to trait dissimilarities among species exploratory studies which report interesting rela‐ may be important. A potential, and unexplored, tionships between biodiversity and environment solution is to decouple functional diversity into to also identifying their causal mechanisms. “phylogenetic structured” and “specific (ecological)” components. This would help us to Acknowledgements better understand historical and recent processes I thank Joaquín Hortal, Thiago Rangel, and Michael on biodiversity patterns and assembly rules (Diniz‐ Dawson for valuable comments on the manu‐ Filho et al. 2011). script. This work was supported by CAPES (project The ground is reasonably well settled to #012/09). start “rebuilding community ecology from func‐ tional traits” (McGill et al. 2006) and “merging Marcus V. Cianciaruso community ecology with evolutionary biol‐ Departamento de Ecologia, Instituto de Ciências Bioló‐ ogy” (Cavender‐Bares et al. 2009). Yes, there are gicas, Universidade Federal de Goiás, Goiânia, GO, some methodological challenges – how to prop‐ Brazil. e‐mail: [email protected]; erly define the species pool and null models, http://www.wix.com/cianciaruso/home which traits should be used, what is the most suit‐ able measure of PD and FD, and so on (see References Pavoine & Bonsall 2011), but we should avoid be‐ Cavender‐Bares, J., Kozak, K., Fine, P. & Kembel, S. coming locked into a blinkered debate about (2009) The merging of community ecology and methodological issues. For example, in the last phylogenetic biology. Ecology Letters, 12, 693– decade more than two measures of PD or FD were 715. frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 89 news and update

Diniz‐Filho, J.A.F, Cianciaruso, M.V., Rangel, T. & Bini, L. Mouchet, M.A., Villéger, S., Mason, N.W.H. & Mouillot, (2011) Eigenvector estimation of phylogenetic D. (2010) Functional diversity measures: an and functional diversity. Functional Ecology, 25, overview of their redundancy and their ability to 735–744. discriminate community assembly rules. Func‐ Gaston, K.J. & Blackburn, T.M. (1999) A critique for tional Ecology, 24, 867–876. macroecology. Oikos, 84, 353–368. Pavoine, S. & Bonsall, M. (2011) Measuring biodiversity Gómez, J.P., Bravo, G.A., Brumfield, R.T., Tello, J.G. & to explain community assembly: a unified ap‐ Cadena, C.D. (2010) A phylogenetic approach to proach. Biological Reviews, 86, 792–812. disentangling the role of competition and habi‐ Ricklefs, R.E. (2008) Disintegration of the ecological tat filtering in community assembly of Neotropi‐ community. American Naturalist, 172, 741–750. cal forest birds. Journal of Animal Ecology, 79, Safi, K., Cianciaruso, M.V., Loyola, R.D., Brito, D., Ar‐ 1181–1192. mour‐Marshall, K. & Diniz‐Filho, J.A.F. (2011) Jenkins, D.G. & Ricklefs, R.E. (2011) Biogeography and Understanding global patterns of mammalian ecology: two views of one world. Philosophical functional and phylogenetic diversity. Philoso‐ Transactions of the Royal Society of London B, phical Transactions of the Royal Society of Lon‐ 366, 2331–2335. don B, 366, 2536‐2544. Losos, J.B. (2008) Phylogenetic niche conservatism, Wiens, J.J. (2011) The niche, biogeography and species phylogenetic signal and the relationship be‐ interactions. Philosophical Transactions of the tween phylogenetic relatedness and ecological Royal Society of London B, 366, 2336–2350. similarity among species. Ecology Letters, 11, Wiens, J.J., Pyron, R.A. & Moen, D.S. (2011) Phyloge‐ 995–1003. netic origins of local‐scale diversity patterns and McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. the causes of Amazonian megadiversity. Ecology (2006) Rebuilding community ecology from Letters, 14, 643–652. functional traits. Trends in Ecology and Evolu‐ Willig, M.R., Kaufmann, D.M. & Stevens, R.D. (2003) tion, 21, 178–185. Latitudinal gradients of biodiversity: pattern, Meynard, C.N., Devictor, V., Mouillot, D., Thuiller, W., process, scale and synthesis. Annual Review of Jiguet, F. & Mouquet, N. (2011) Beyond taxo‐ Ecology, Evolution, and Systematics, 34, 273– nomic diversity patterns: how do α, β and γ 309. components of bird functional and phylogenetic diversity respond to environmental gradients Edited by Thiago F. Rangel across France? Global Ecology and Biogeogra‐ phy, 20, 893–903.

Remember that being a member of IBS means you can get free online access to four biogeo‐ graphy journals: Journal of Biogeography, Ecography, Global Ecology and Biogeography and Diversity and Distributions. You can also obtain a 20% discount on the journals Oikos and Jour‐ nal of Avian Biology. Additional information is available at http://www.biogeography.org/.

90 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update book review A mangrove compendium World atlas of mangroves, by Mark Spalding, Mami Kainuma and Lorna Collins (editors) 2010, Earthscan, 336 pp.ISBN: 9781844076574 Price: £65 (Hardback); http://www.earthscan.co.uk/

The World atlas of mangroves, an update to Spal‐ country fact sheets serve as an excellent compen‐ ding et al. (1997), is a must‐have publication for dium and make this atlas perfect as a quickstart everyone loving and working with, in, or near to guide for students as well as experienced re‐ mangroves. It celebrates the wonderful world of searchers approaching a new region. these beautiful forests with astonishing figures Considering the presentation of global and photographs. The informative maps and ta‐ trends as the main purpose of the World Atlas Of bles provide captivating facts about the ecological Mangroves, this book fulfils expectations. Unnec‐ and economic values of mangroves and the conse‐ essary uncertainties and errors in the introduction quences of their loss. to the ecology of mangroves leave, however, a The atlas scores with the presentation of drop of bitterness. The first chapters (Mangrove recent findings on carbon sequestration, showing ecosystems and Mangroves and people) notably that mangroves store more carbon than tropical omit explicit references to any publications. The forests (Donato et al. 2011); and with the suitabil‐ authors state that these chapters and the boxes ity of intact mangroves for protecting coastal re‐ therein ‘draw heavily’ on the relevant literature, gions against tsunamis (Wibisono and Suryadipu‐ but information presented is confusing or even tra 2006). This will arm (with powerful arguments) erroneous, and does not always reflect the con‐ ecologists, conservation biologists and policy‐ tent of the publications loosely mentioned at the makers, who urgently need to communicate this end of each subchapter, nor established knowl‐ knowledge in order to increase public awareness edge available in textbooks (e.g. Tomlinson 1986) and political willingness to protect and rehabili‐ or extended reviews (e.g. Feller et al. 2010). For tate one of the most vulnerable ecological sys‐ example, the classification of mangroves into tems on earth. fringing mangroves, basin mangroves, and over‐ As indicated by its title, the World atlas of wash mangroves is needlessly incomplete; it could mangroves gives a comprehensive overview of the be easily improved by following standard man‐ global distribution of mangrove species at country grove literature (e.g. Lugo & Snedaker 1974, level. A detailed description of the particular Woodroffe 1992). The heterogeneous handling of status of mangrove systems in each country, ac‐ outdated theories and debated hypotheses about companied by information about their specific the functioning of mangroves is also surprising. threats, level of degradation and extent of reha‐ For instance, the editors correctly do away with bilitation programs guides the reader through a the perspective that the land creates the capabil‐ multitude of distinct features, while keeping simi‐ ity for mangrove formation, but then present ele‐ larities and general principles in mind. vation and the subsequent gradient of inundation Mangrove experts of international repute as the only factors driving patterns of species contribute boxes on particular topics of interest, zonation. There are, however, four other major such as mangroves’ responses to climate change hypotheses to explain this striking feature: geo‐ (Gilman, Duke et al.) or their functioning in highly morphological influences, propagule dispersal, dynamic coastal regions (Fromard and Proisy). predation and species competition (see e.g. Smith They summarise up‐to‐date research as well as III 1992 for detailed discussion). Further errors in the hot topics that will be developed in the near the classification of aerating roots and also in the future. In addition, the annexes containing tree systematics and geographical distribution of some species descriptions, national species lists and mangrove species have been already listed and frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 91 news and update discussed in detail by Dahdouh‐Guebas (2010). It mangrove research is a challenge, an urgent de‐ remains a mystery why these chapters have not mand for mankind and that being involved is an been written or carefully revised by the leading accolade. On the other hand, if you are looking for mangrove experts mentioned above, or the nu‐ a general text spanning the interdisciplinary as‐ merous others who contributed to this book with pects of mangrove ecology, this is not the book for specific boxes. you. The roots of this book largely come from ge‐ This volume appears 14 years after Man‐ ography and remote sensing. If you are searching groves – The forgotten forest between land and for an up‐to‐date text about the present scientific sea (Mastaller 1997). It seems that the world has understanding and recent findings in mangrove changed and the forgotten forest has been redis‐ research, I recommend supplementing the atlas covered. Obviously neither the simple existence of with textbooks, recent reviews or more detailed this remarkable ecosystem, nor its fascinating publications on mangrove ecosystems and peo‐ functioning based on adaptation to the harsh con‐ ple’s depency on their health and functioning. ditions of tidal zones, were sufficient to convince people that it is worth protecting mangroves Uta Berger against aquaculture, agriculture, land use and the Institut für Waldwachstum und Forstliche Informatik, many types of waste water we produce. The Technische Universität Dresden monetary expression of the value of mangroves e‐mail: [email protected]‐dresden.de; –1 –1 (US$ 2000–9000 ha yr according to the statis‐ http://www.forst.tu‐dresden.de/SystemsAnalysis/uta‐berger tics in this book), and the change from the eco‐ logical perspective to the human perspective in References terms of coastal protection against hurricanes and Dahdouh‐Guebas, F. (2011) World Atlas of Mangroves: tsunamis and in carbon sequestration, is neces‐ Mark Spalding, Mami Kainuma and Lorna Collins sary to improve public awareness about the im‐ (eds). Human Ecology, 39, 107–109. portance of mangroves for our present life and a Donato, D.C., Kauffman, J.B., Murdiyarso, D., Kurnianto, critical part of our response to the challenges of S., Stidham, M. & Kanninen, M. (2011) Man‐ groves among the most carbon‐rich forests in environmental changes, including sea level rise the tropics. Nature Geoscience, 4, 293–297. and climate change. The World atlas of mangroves Feller, I.C., Lovelock, C.E., Berger, U., McKee, K.L., Joye, is a strong contribution towards this goal and, I S.B. & Ball, M.C. (2010). Biocomplexity in Man‐ hope, another step towards ushering in a new era grove Ecosystems. Annual Review of Marine where mangroves are valued for their beauty in Science, 2, 395–417. Lugo, A.E. & Snedaker, S.C. (1974). The ecology of man‐ the same way as many rain forests or coral reefs. groves. Annual Review of Ecology and Systemat‐ In summary, if you are working in the field ics, 5, 39–64. of mangrove conservation or related issues in the Mastaller, M. (1997) Mangroves – the forgotten forest context of tropical coastal zones, or if your work is between land and sea. Tropical Press Sdn. BhD. targeted towards practitioners, stakeholders or Kuala Lumpur, Malaysia. 189 pp. Smith III, Th.J. (1992). Forest Structure. In: Tropical users of at‐risk mangrove ecosystem services, the mangrove ecosystems (ed. by A.I. Robertson and World atlas of mangroves is your book; it will sup‐ D.M. Alongi), pp.101–136. American Geophysi‐ port your daily work with easy‐to‐understand in‐ cal Union, Washington. formation and strong facts about the ecological Spalding, M., Blasco, F. & Field, C. (1997). World man‐ and economic values of this forest. If you are a grove atlas. The International Society for Man‐ mangrove ecologist, this book should also be on grove Ecosystems, Okinawa, Japan. 178 pp. Tomlinson, P.B. (1986). The botany of mangroves. Cam‐ your shelf because it provides you with a quick bridge University Press, Cambridge, UK. 419 pp. overview of mangrove distribution and current Wibisono,I.T.C. & Suryadiputra, N.N. (2006). Study of status on Earth. It also acts as an enormous source lessons learned from mangrove/coastal ecosys‐ of suitable maps and material to round off your tem restoration efforts in Aceh since the tsu‐ lectures. This should convince your students that nami. Wetlands International – Indonesia Pro‐ gramme, Bogor. 86 pp.

Woodroffe, C.D. (1992). Mangrove sediments and geo‐ morphology. In: Tropical mangrove ecosystems (ed. by A.I. Robertson and D.M. Alongi), pp.7– 41. American Geophysical Union, Washington.

Edited by Markus Eichhorn book review A comprehensive foundation for the application of biogeogra‐ phy to conservation Conservation biogeography, by Richard J. Ladle and Robert J. Whittaker (editors) 2011, Blackwell Publishing, 301 pp. ISBN: 9781444335033 Price: £95 (Hardback) / £34.95 (Paperback); http://eu.wiley.com/

It is becoming increasingly clear that the diversity tion biogeography (Chapter 1). A distinction is of plant and animal species in the world is con‐ made between approaches that focus on the com‐ tinuing to decline in spite of ambitious targets set position of biological communities and those that by governments to prevent this (Butchart et al. focus on ecosystem function through an under‐ 2010). It is also becoming evident that the contin‐ standing of ecosystem processes such as nutrient ued functioning of ecosystems depends on this cycling (p. 31). An interesting and growing field in diversity (Isbell et al. 2011). In order to conserve ecology, which receives little attention in the what is left of biodiversity, it is crucial that we un‐ book, uses the functional traits of species to ex‐ derstand the diversity of life and how it is distrib‐ plain the link between the composition of biologi‐ uted across the biomes and ecosystems of the cal communities and the function of the ecosys‐ world. Since understanding the distribution of bio‐ tems that contain them. Functional traits – such as diversity is a central tenet of biogeography, it body mass, diet, habitat affinity and development seems obvious that the field of biogeography mode of animals, and height and photosynthetic should be of central importance in conservation. pathway of plants – can help explain how species In this volume, Richard Ladle and Robert contribute to the processes underlying the func‐ Whittaker bring together chapters by a number of tioning of ecosystems and can also help in predict‐ biogeographers to summarise progress to date in ing how ecosystems will respond to environ‐ applying the principles of biogeography to conser‐ mental change (McGill et al. 2006). vation and to identify areas where there is still The second section reviews our current un‐ work to be done. The book is a comprehensive but derstanding of the distribution of biodiversity, digestible summary of the field of conservation summarises the history of the global protected biogeography and should make essential reading, areas network and describes the methods avail‐ not only for the students at whom it is primarily able for more systematically representing biodi‐ aimed, but also for more experienced scientists. versity in future extensions to this network. There The editors profess at the outset that the aim was is a strong terrestrial focus here, indeed through‐ to achieve a degree of coherence among the out the entirety of the book, which the authors chapters, an aim that is achieved remarkably well acknowledge and which is owing to a less com‐ to give a very coherent text. plete understanding of the distribution of diversity The first section of the book provides a brief in the oceans and in freshwater habitats. It is but interesting history of the conservation move‐ worth noting, though, that the Census of Marine ment and the contrasting values held by different Life, an ambitious $650 million project that fin‐ sectors of this movement (Chapters 2 and 3), as ished recently, has made huge progress towards well as some background to the field of conserva‐ understanding the biogeography of the oceans frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 93 news and update

(e.g. see Tittensor et al. 2010). Even in the terres‐ and environmental factors (e.g. Ferrier et al. trial realm, knowledge about the number and 2007). identity of the world’s species and how they are With a growing need to understand changes distributed remains very far from complete: the in the natural environment and the impact of Linnaean and Wallacean shortfalls respectively these changes on human society, the emerging (Chapter 4). A recent paper (Joppa et al. 2011) field of conservation biogeography is likely to be‐ addressed both of these knowledge gaps simulta‐ come increasingly important in providing the nec‐ neously by predicting the spatial distribution of essary theoretical basis and tools for doing so. undiscovered plant species, predicting that most This book provides an excellent foundation for new plant species will be discovered in areas al‐ that field and is highly recommended reading for ready identified as hotspots of plant diversity, em‐ students, scientists and practitioners of conserva‐ phasising the importance of these areas for con‐ tion. servation. Chapter 5 provides an excellent sum‐ Tim Newbold mary of the many different types of protected United Nations Environment Programme World Con‐ areas in the global network and the different val‐ servation Monitoring Centre, Cambridge, UK ues that underpin these, while Chapter 6 provides e‐mail: Tim.Newbold@unep‐wcmc.org; a useful and succinct review of the enormous and http://www.unep‐wcmc.org/tim‐newbold_368.html ever‐growing literature on systematic conserva‐ tion planning. References The third section of the book describes how Butchart, S.H.M., Walpole, M., Collen, B. et al. (2010). the tools of biogeography can be used to plan for Global biodiversity: indicators of recent declines. environmental change in conservation. This is the Science, 328, 1164–1168. only part of the book where the chapters appear Isbell, F., Calcagno, V., Hector, A. et al. (2011). High somewhat disjointed, but this is probably owing to diversity is needed to maintain ecosystem ser‐ the attempt to summarise a vast literature in a vices. Nature, 477, 199–202. very small number of chapters. Nevertheless, the Joppa, L.N., Roberts, D.L., Myers, N. et al. (2011). Biodi‐ versity hotspots house most undiscovered plant chapters in this section provide excellent descrip‐ species. Proceedings of the National Academy of tions of some of the available methods, from phe‐ Sciences of the United States of America 108, nomenological models that infer future changes 13171–13176. from current patterns (Chapter 7) to more process McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. ‐based models that use the theory of island bio‐ (2006). Rebuilding community ecology from functional traits. Trends in Ecology & Evolution, geography to predict the consequences for biodi‐ 21, 178–185. versity of shrinking and increasingly isolated natu‐ McKnight, M.W., White, P.S., McDonald, R.I., Lam‐ ral habitat patches (Chapter 8). Chapter 9 deals oreux, J.F., Sechrest, W., Ridgely, R.S. & Stuart, with invasive species, which are an important S.N. (2007). Putting beta‐diversity on the map: driver of environmental change, and the homog‐ broad‐scale congruence and coincidence in the extremes. PLoS Biology, 5, e272. enisation of biological communities, i.e. the ero‐ Tittensor, D.P., Mora, C., Jetz, W., Lotze, H.K., Ricard, sion of beta diversity. Most of the studies investi‐ D., Vanden Berghe, E. & Worm, B.(2010). Global gating broad‐scale patterns of diversity have fo‐ patterns and predictors of marine biodiversity cused on inventory diversity, commonly measured across taxa. Nature, 466, 1098–1101. as species richness, and it is only recently that studies have attempted to map beta diversity (e.g. Edited by Markus Eichhorn McKnight et al. 2007) and to relate it to spatial

One of the benefits open to IBS members is the opportunity to have job openings posted on the IBS blog (http://biogeography.blogspot.com/). If you have a position you would like to have ad‐ vertised, please contact Karen Faller ([email protected]) or Michael Dawson ([email protected]) with details.

94 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update book review A new encyclopedia for biological invasions Encyclopedia of biological invasions, by Daniel Simberloff and Marcel Rejmánek (editors) 2011, University of California Press, 792 pp. ISBN: 9780520264212 Price US$95 (Hardback or e‐book); http://www.ucpress.edu/

Despite existing in some form for many decades ries of key topics relating (in this case) to biologi‐ (Davis 2005), invasion ecology/biology is in many cal invasions, without citations but with relevant ways a nascent and emerging field, and is still en‐ further reading at the end. The entries vary in gendering discussion regarding whether it indeed length from 1 to 8 pages, and often incorporate truly exists as a field or discipline in its own right, useful figures and occasionally tables. The book is or is rather a particularly focused aspect of com‐ impressively glossy (all figures are in full colour) munity ecology or biogeography (e.g. Marris 2009, and well presented, which is all the more remark‐ Pyšek and Hulme 2009). As with many ecological able considering the relatively modest price. The disciplines, invasion ecology has seen fundamen‐ editors, Daniel Simberloff and Marcel Rejmánek, tal disagreements over aspects ranging from core are leading invasion ecologists and are well quali‐ definitions (including ‘invasion’ itself; Falk‐ fied to compile such a text; this is reflected not Petersen et al. 2006, Ricciardi and Cohen 2007) to just in the broad range of well‐selected topics that level of scientific objectivity (e.g. Larson 2007). the volume includes (of which there are 153) but The field is at a stage in its development where (1) also the roll‐call of esteemed contributors that dedicated journals exist (e.g. Biological Invasions) have supplied the entries (of which there are 197, and there is a substantial number of academic many of them high‐profile international research‐ articles published every year (for example a ers). The book is aimed not just at an academic search of ‘invasive species’ in Web of Knowledge audience, however, and the articles are written returns 1181 articles published in 2010 alone), 2) with the interested and educated general public in there is clear and significant international interest mind. and action in relation to invasions and (3) an ex‐ The individual articles cover various aspects tended peer community is involved in researching of invasions, ranging from particular attributes of and managing the threat of invasive species, from invasive species and invaded ecosystems to im‐ world‐leading academics at research‐intensive pacts and management, interesting case studies universities to local government and conservation and historical perspectives. Clearly it is not possi‐ volunteers. The result of the burgeoning informa‐ ble to cover all of the entries in a review such as tion and uneven levels of understanding and focus this, but I did find several articles especially inter‐ across the peer community is confusion and un‐ esting, particularly because they highlight the certainty, right from the fundamentals (what is an many socioecological factors that complicate our invasive species exactly, and why is it invasive?) to relationships with potentially problematic species. the specifics (what is the best technique for reduc‐ The entry on Xenophobia for example does an ex‐ ing populations of Crassula helmsii in my pond, cellent job of summarising how society’s relation‐ and how does that differ from managing spread in ship with non‐native species is constructed in cer‐ the local lake?). The time is ripe therefore for an tain ways by the use of loaded terms or cultural encyclopaedia such as this one by Daniel Simber‐ metaphors, for example the negative personifica‐ loff and Marcel Rejmánek to form a baseline for tion of zebra mussels as ‘outlaws’ on the west future definitions and discussions. coast of the US, or the badging of ‘harmful’ or The book is one of University of California ‘distasteful’ species with appellations that note Press’ Encyclopedias of the Natural World series, their foreign status (Japanese knotweed, Chinese and as with the other volumes has a wide range of mitten crab, English sparrow and so on). As a entries that are effectively short essays or summa‐ starting point for a discussion of scientific objec‐ frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 95 news and update tivity related to invasion biology it works excep‐ other reasons), in most cases failed to be realised tionally well, and is exactly the right size for diges‐ and also (with some notable exceptions) that tion by students or interested amateurs. many societies were unsuccessful in actually natu‐ Indeed, one of the best uses I find for refer‐ ralising many species at all. But much is left un‐ ence works such as these are as opening forays said: in some cases one is left wanting to know into topics for class discussions, whether at gradu‐ more about whether species referred to as ate or undergraduate level. Good examples in‐ ‘released’ became naturalised, whether regions clude the entry on Succession, which very effec‐ such as South America maintained any such socie‐ tively and concisely summarises key concepts that ties (these countries are ignored, while others take up whole chapters in many textbooks, and such as Germany and Italy receive only one sen‐ although invasion biology is only addressed to‐ tence) and ultimately whether such societies indi‐ wards the end, it is clear how the two link to‐ rectly provided evidence to force their own dis‐ gether. Likewise, the discussion on Native invad‐ continuation. As a taster to whet the appetite, the ers, in which issues of ‘invasive’ terminologies article succeeds very well (and relevant books on (and when they are appropriate) are covered, is the subject are provided in the Further Reading excellently written and illuminating at a range of section), but it is not an authoritative, encyclopae‐ levels, particularly in relation to the many exam‐ dic summary in itself. ples of ‘invasion’ given. Certainly students and As with any vast topic, covering all aspects researchers new to the subject will have any initial in a single volume is difficult – in this case there is confusion over what is meant by invasions dis‐ differential coverage of ecosystems (e.g. entries pelled by the article, and it will also help them to for canals, lakes, rivers and wetlands, but no cov‐ think objectively about whether a species really erage of urban ecosystems, despite these being may be considered invasive or not. All of the arti‐ important points of introduction for some invasive cles I read through were of a high quality and well taxa); hypotheses (e.g. Enemy Release Hypothesis, written/edited, with very little wasted space for Novel Weapons Hypothesis, but no Tens Rule); such a large volume (although on occasion figures geographical areas (Australia, the Great Lakes, are not always relevant – I’m not sure why an im‐ Hawaiian islands, the Mediterranean, the Ponto‐ age of Frank Buckland ‘physicking a por‐ Caspian, New Zealand and South Africa receive a poise’ (page 2) is worthy of inclusion for example, particular focus) and species (good examples of despite his role in founding the main UK acclimati‐ some key species or groups such as zebra mussel, sation society). earthworms and fishes, but understandably not Of course, it is always hard to get the right comprehensive coverage). This is entirely reason‐ balance between conciseness and detail in such able, and is not a criticism of the volume – it is entries, and to retain the relevant focus. The impossible to cover the vast range of topics asso‐ opening entry, Acclimatisation societies is a case ciated with biological invasions in sufficient depth in point: the article does an excellent job of sum‐ in a single volume, and the material that is in‐ marising the development and impact of such so‐ cluded is impressive. The division of the book be‐ cieties in different countries, many of which were tween invader attributes, processes, taxa, ecosys‐ responsible for the introduction of significant tems, pathways to invasion and so on is very well numbers of non‐native species around the globe done and represents a huge effort on the part of before dying out in the face of increasing legisla‐ the editors, for which they should be roundly con‐ tion, awareness of ecological risk from introduc‐ gratulated. I would encourage consideration of a tions and lack of interest from the general public. second volume, however, at least with regard to The article elegantly conveys how originally be‐ key concepts and hypotheses. The opening guide nevolent intentions, such as the introduction of to the Encyclopedia notes that there is a website non‐natives to improve food resources, control with a list of articles, sample entries and so, and pests and to soothe homesick colonists (among notes that the site ‘will evolve with the addition of

96 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update new information’, p. xxii). The web address has References since changed and I was unable to locate the new Davis, M.A. (2005) Invasion biology 1958‐2004: the one. Though I happily agree that this could poten‐ pursuit of science and conservation. In: Concep‐ tially be a very useful resource, given the rapidly tual ecology and invasions biology: reciprocal approaches to nature (ed. by Cadotte, W.M, changing environment of the internet, the publi‐ McMahon, S.M. and Fukami, T.) , pp. 35–64. cation of a second volume would perhaps be the Kluwer Publishers, London. most reliable option. Falk‐Petersen, J., Bøhn, T. & Sandlund, O.T. (2006) On In summary, this is an excellent reference the numerous concepts in invasion biology. Bio‐ work that combines readability with academic logical Invasions, 8, 1409–1424. rigour throughout. Its broad coverage of the field, Larson, B.M.H. (2007) An alien approach to invasive species: objectivity and society in invasion biol‐ high quality of production and reasonable price ogy. Biological Invasions, 9, 947–956. makes it an essential purchase for any university Marris, E. (2009) The end of the invasion? Nature, 459, with departments teaching or researching within 327–328. the broad spectrum of ecology, as well as for indi‐ Pysek, P. & Hulme, P.E. (2009) Invasion biology is a dis‐ vidual researchers of species invasions. cipline that’s too young to die. Nature, 460, 324 –324.

Ricciardi, A. & Cohen, J. (2007) The invasiveness of an Robert A. Francis introduced species does not predict its impact. Department of Geography, King’s College London Biological Invasions, 9, 309–315. e‐mail: [email protected]; http://rg.kcl.ac.uk/ staffprofiles/staffprofile.php?pid=1961 Edited by Markus Eichhorn book review A piscine history of the Neotropics Historical biogeography of Neotropical freshwater fishes, by J.S. Albert and R.R. Reis (editors) 2011, University of California Press, 408 pp. ISBN: 9780520268685 Price £59 (Hardback); http://www.ucpress.edu/

The Neotropics leave an indelible impression on the planet and around 10% of all vertebrate spe‐ everyone who visits them. The seeds of some of cies. the most important concepts in ecology and evo‐ James Albert and Roberto Reis’ goal as edi‐ lution were sown during the South American trav‐ tors of the Historical Biogeography of Neotropical els of influential 19th century thinkers. For exam‐ Freshwater Fishes is to examine the evolutionary ple, the latitudinal gradient of diversity, now rec‐ forces responsible for this diversity. In doing so ognized as ecology’s oldest pattern (Hawkins, they make the case that multiple processes of di‐ 2001), was first identified by von Humboldt, while versification were involved and that these oper‐ Bates documented the variety and adaptations of ated over long periods of time as well as on a con‐ species in Amazonian forests, and Wallace and tinental scale. The book itself is divided into two Darwin pondered the mechanisms responsible for parts, the first of which examines current knowl‐ the myriad forms of life they encountered. Al‐ edge on the biogeography of the region, while the though the Neotropics have played a crucial role second is a regional analysis that links contempo‐ in our understanding of the diversity of life on rary geographical patterns with geological history. earth, in many ways they continue to represent an The book is ambitious in scope and brings to‐ unexplored frontier. This is particularly clear in the gether previously fragmented material to provide case of Neotropical freshwater fish, a group esti‐ an authoritative overview of this impressive group mated to consist of more than 7000 species, and of fish. And while a fish‐eye view of the Neotropi‐ that accounts for over half the freshwater fish on cal ichthyofauna is inevitably drawn to the Ama‐ frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 97 news and update zon, the book has broad coverage, embracing the cies than others. The Neotropics are no exception. Andes and extending through Central America and Ten families of fish account for 75% of the into southern Mexico. As it makes clear, it is nec‐ Neotropical icthyofauna. Characidae (including essary to have a continental perspective to under‐ piranhas and tetras) and Cichlidae (such as discus) stand the diversity and distribution of this impres‐ are particularly big hitters. One possibility is that sive group. this unevenness is simply the result of chance. I particularly liked the care and thought in‐ Alternatively, historical and biological factors, ei‐ volved in putting the book together. It is a beauti‐ ther separately or together, could contribute. E.O. fully presented volume with informative tables Wilson (2003) has argued that an ancient origin, and figures, many of them in colour. However, combined with small body size, widespread geo‐ more important than this is that the editors have graphic distribution and key innovations contrib‐ a strong sense of what the important issues are ute to the success of some groups relative to oth‐ and how these should be best dealt with. Indeed ers. On the basis of the evidence presented by the book is an essential reference for anyone Neotropical fish, Albert, Bart and Reis conclude wanting to learn more about the diversity or his‐ that these features are necessary but not suffi‐ tory of South American fishes. cient. Indeed they note that clades can be ancient One of the most challenging questions in (e.g. Arapaima, which is of Cretaceous origin), ecology is explaining why different habitats sup‐ widespread (Arapaima again) or with small body port different numbers of species. The extent of a size (e.g. Amazonsprattus) yet be represented by a habitat accounts for much of the variation but handful of species at most. On the other hand sex‐ South America has an excess of species relative to ual and trophic innovation may play a role. Eco‐ its area. The core of the continent, particularly the logical specialisation is also important. For exam‐ Amazon, is responsible for a disproportionate ple, Crampton notes that groups of closely related amount of this diversity. It is tempting to attribute Gymnotiform electric fish species tend to be this exceptional richness to the unique geological found in a narrow range of habitat types but may and environmental features of the Amazon. How‐ be spread across large geographic areas. The fac‐ ever many of the fishes that inhabit this river sys‐ tors that underpin diversification are the same as tem are older than the Amazon Basin itself. More‐ those that come into play in the explosive speci‐ over, the Amazonian ichthyofauna has been accu‐ ation that characterizes the African rift lakes. The mulated gradually through tens of millions of difference here is that the game is played out on a years. The explanation, Albert, Petry and Reis ar‐ continental scale as opposed to a local arena. gue, is rooted in the repeated subdivision and Of course, much remains to be learnt about merging of adjacent river basins and their faunas, the phylogenetic histories of Neotropical fishes with dispersal limitation and environmental filter‐ and of the geological context in which these spe‐ ing playing important roles. The exceptionally high cies evolved. Nonetheless, as this book makes diversity seems to be less to do with exceptional clear, the nature and timing of key events is be‐ speciation rates than with low rates of extinction. coming much better understood. The contribu‐ However, diversity is not just a measure of the tions to the book demonstrate how the growing numbers of species that co‐occur but also of the body of molecular data, and its integration with types of species that are found together. A univer‐ ecological theory and earth sciences, has under‐ sal feature of natural assemblages is that some pinned the recent and rapid progress in under‐ families contribute a much higher fraction of spe‐ standing this system.

Your participation in frontiers of biogeography is encouraged. Please send us your articles, com‐ ments and/or reviews, as well as pictures, drawings and/or cartoons. We are also open to sug‐ gestions on content and/or structure. Please check http://www.biogeography.org/html/fb.html for more information, or contact us at [email protected] and [email protected].

98 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update There have been many studies of tropical References diversity but until now Neotropical fishes fish have Hawkins, B. A. (2001). Ecology’s oldest pattern. Trends received relatively little attention. This contrasts in Ecology and Evolution 16, 470. with South American birds, a group that has been Rahbek, C., Gotelli, N. J., Colwell, R. K., Entsminger, G. prominent in tests of macroecological hypotheses L., Rangel, T. F. L. V. B. and Graves, G. R. (2007). Predicting continental‐scale patterns of bird (e.g. Rahbek et al., 2007). Fish are responsible for species richness with spatially explicit models. more diversity and deserve to be more fully stud‐ Proceedings of the Royal Society B: Biological ied. This book provides the knowledge that will Sciences 274, 165‐174. inform these exciting research opportunities. Wilson, E.O. (2003). The origins of hyperdiversity. pp. 13‐18 in Pheidole in the New World: A Dominant Hyperdiverse Ant Genus, Wilson, E.O. (ed). Har‐ Anne E. Magurran vard University Press.

University of St Andrews Edited by Markus Eichhorn e‐mail: aem1@st‐andrews.ac.uk; http://biology.st‐andrews.ac.uk/magurran/ books noted with interest

Principles of terrestrial ecosystem ecol‐ Guide to standard floras of the World: ogy An annotated, geographically ar‐ F. Stuart Chapin III, Pamela A. Matson & Peter ranged systematic bibliography of the M. Vitousek principal floras, enumerations, check‐ 2011, 2nd edition, Springer, 529 pp. lists and chorological atlases of differ‐ £135 (Hardback), £44.99 (Paperback) ISBN: 9781441995032 / 9781441995025 ent areas http://www.springer.com/ David F. Frodin 2001, 2nd edition, Cambridge University Press, An outstanding textbook which, after definitions, 1100 pp. sets the stage with primers on Earth’s climate sys‐ £198 (Hardback), £90 (Paperback), US$120 (e‐ tem and geological processes. What follows is a book) magisterial and comprehensive account of the ISBN: 9780521790772 / 9780521189774 movements of water, energy, carbon and nutri‐ http://www.cambridge.org/ ents though natural systems. Along with standard generalisations, the authors delve into the finer While not generally our policy to feature reprints, detail and explain how biological processes can this standard text has newly appeared in paper‐ have important modulating effects through space back, bringing it within affordable reach of a and time. A final reflective pair of chapters consid‐ greater number of researchers. It does exactly ers global changes and the implications for ecosys‐ what it says on the cover, making it the definitive tem management. The book is well written reference for anyone commencing work on the throughout and punctuated with excellent colour flora of a new region. Despite its not receiving any illustrations; no‐one from undergraduates to es‐ further updates and its coverage ending in 1999, tablished researchers can fail to learn something there remain no resources to rival it, either in from it. print or online. It also contains insightful reviews on the history of floristic description. An essential book which belongs in the library of every plant biogeographer.

frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 99 news and update Field guide Afghanistan: Flora and Community ecology vegetation Peter J. Morin Siegmar‐W. Breckle & M. Daud Rafiqpoor 2011, 2nd edition, Wiley‐Blackwell, 407 pp. 2011, Scientia Bonnensis, Bonn, 864 pp. £90 (Hardback), £34.99 (Paperback) Price: Contact publishers ISBN 9781444338218 / 9781405124119 ISBN: 9783940766304 http://www.wiley.com/ http://www.scientia‐bonnensis.com/ Community ecology straddles conventional inter‐ The flora of this vast, environmentally diverse and action‐based ecology and biogeography; recent biogeographically central country has yet to be heated debate in the pages of American Naturalist fully catalogued, but this field guide represents a has even disputed whether communities truly ex‐ landmark accomplishment on the path to doing ist as natural entities. Unsurprisingly the author so, filling an anomalous gap at the junction of sev‐ makes a strong case for communities, stressing eral floristic realms. It contains a pictorial guide to patterns and processes that can only be under‐ over 1200 species (>25% of the flora) plus general stood at this level, and pleasingly devotes equal chapters on vegetative formations and should fa‐ attention to both models and experimental data. cilitate both local and international study. Copies The textbook is intended for a graduate course have been freely distributed to universities and and represents a major update on the previous institutes throughout Afghanistan as well as her‐ edition. One might query the balance of coverage baria and museums worldwide. A feature on this of various topics but nevertheless this remains the project is planned for a future edition of Frontiers only textbook exclusively devoted to this scale of of Biogeography. study.

Markus Eichhorn Book Review Editor. e‐mail: [email protected]

Editorial policy for book reviews Frontiers of Biogeography will publish in‐depth reviews of recently published books (typically less than one year old) on biogeography or of interest to biogeographers, alongside a ‘Noted with Interest’ section provi‐ ding brief details of new publications. Authors, editors or third parties are invited to suggest books for re‐ view to the Book Review Editor, Dr Markus Eichhorn, School of Biology, University Park, Nottingham NG7 2RD, United Kingdom; telephone ++44 (0)115 951 3214; e‐mail [email protected]. We welcome offers to review books for Frontiers of Biogeography, but will not accept an offer to review a speci‐ fic book. 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, ex‐ pressing an opinion about the value of the book, its focus and breadth, setting it in the context of recent developments within the field of study. Textbook reviews should consider their utility as resources for tea‐ ching and learning. Avoid describing the book chapter by chapter or listing typographical errors. The length should normally be 1000 words (1500 words for joint reviews of related texts) including a maximum 10 refe‐ rences. Authors may suggest a short heading for the review, followed by the title of the book(s), the aut‐ hors/editors, publisher, publication date, price, hbk/pbk, pages, ISBN and website (where available). Figures or tables will not ordinarily be included. Authors of reviews must verify that they have not offered (and will not offer) a review of the same book to another journal, and must declare any potential conflict of interest that might interfere with their objectivity. This may form a basis for editorial decisions and such disclosures may be published. Book reviews will usually go through a light editorial review, though in some circumstan‐ ces also will be considered by one or more referees.

100 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 news and update thesis abstract Applying species distribution modeling for the conservation of Iberian protected invertebrates Rosa María Chefaoui PhD Thesis, Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales, c/ José Gutiérrez Abascal 2, 28006 Madrid, Spain. e‐mail: [email protected]; http://www.biogeografia.org/

Abstract. This article outlines the approaches to modeling the distribution of threatened invertebrates using data from atlases, museums and databases. Species Distribution Models (SDMs) are useful for esti‐ mating species’ ranges, identifying suitable habitats, and identifying the primary factors affecting species’ distributions. The study tackles the strategies used to obtain SDMs without reliable absence data while exploring their applications for conservation. I examine the conservation status of Copris species and Graellsia isabelae by delimiting their populations and exploring the effectiveness of protected areas. I show that the method of pseudo‐absence selection strongly determines the model obtained, generating different model predictions along the gradient between potential and realized distributions. After assess‐ ing the effects of species’ traits and data characteristics on accuracy, I found that species are modeled more accurately when sample sizes are larger, no matter the technique used. Keywords: Environmental niche modeling, Iberian Peninsula, invertebrates, predictive accuracy, species distribution models

The rapid disappearance of habitats and species Geographic Information Systems (GIS) sig‐ starkly contrasts the need to conserve biodiversity nificantly advanced the conservation of endan‐ against our inability to inventory and protect all gered species because they allow us to delimit species individually. Knowledge about biodiversity species’ potential distributions (e.g. Hortal et al. remains insufficient because many species are still 2005), to control their populations not described (the "Linnean Shortfall"; Brown (e.g. Davies et al. 2005), to analyze their niche and Lomolino 1998) and the distributions of de‐ (Peterson et al. 2002), design networks of pro‐ scribed species often are inadequately defined tected areas (e.g. Pearce and Boyce 2006), and to (the "Wallacean Shortfall"; Lomolino 2004). It is forecast the future (e.g. Hill et al. 2002). Together, therefore essential to identify threatened species the databases taken from atlases, museums and and describe their distributions using approaches herbaria have emerged as a valuable source of that overcome the time and budget constraints of species’ occurrence records (e.g. Elith and Leath‐ systematic conservation planning. wick 2007). Unfortunately, these data from het‐ Araújo et al. (2007) demonstrated the need erogeneous sources may contain errors or for additional protected areas for the effective have been obtained using a biased sampling pro‐ conservation of the diversity of plants and verte‐ cedure (Hortal et al. 2007, 2008, Newbold brates in the Iberian Peninsula. Preliminary data 2010). Besides, they do not usually provide reli‐ suggest that the existing network of reserves also able absences needed to perform consistent pre‐ would be ineffective in representing invertebrate dictive models (Anderson et al. 2003, Lobo et al. species (Verdú and Galante 2009). Unfortunately, 2007), so alternatives have been sought generat‐ the conservation of invertebrates faces serious ing models based only on presences (Hirzel et al. challenges due to their high diversity, complex life 2002, Pearce and Boyce 2006), sometimes em‐ cycles and difficult taxonomy, among other factors ploying pseudo‐absences obtained in different (see New 1998). ways (Zaniewski et al. 2002, Engler et al. 2004, frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 101 SDM applied to invertebrate conservation

Lobo et al. 2006, 2010). presence of Copris hispanus, while Copris lunaris For my doctoral thesis, I evaluated the util‐ requires siliceous soils and high rainfall. Both Co‐ ity of SDMs for the conservation of threatened pris species are distributed along a geographic and invertebrates in the Iberian Peninsula (Chefaoui environmental gradient from the Tajo basin 2010). The majority of the species studied (warmer, dryer, with strong annual weather varia‐ here have been designated by the European Un‐ tions) where only C. hispanus is found, towards ion as species of “community interest” requiring the mountain slopes of the Sistema Central protection and conservation (Habitats Directive). I (colder, higher rainfall) where C. lunaris predomi‐ used presence‐only data on Iberian threatened nates. The environmental niches of both species invertebrates obtained from museums, atlases are distributed along a Dry‐Mediterranean to Wet and databases. I applied presence‐only methods ‐Alpine axis, and overlap in areas of moderate such as ENFA (Ecological Niche Factor Analysis) temperatures and precipitations in the north of and MDE (Multi‐Dimensional Niche Envelope), in CM. addition to other methods that require presences We also studied the degree of protection of and absences (here, pseudo‐absences): GAM key populations of C. hispanus and C. lunaris, mak‐ (Generalized Additive Models), GLM (Generalized ing a proposal to improve their conservation. To Linear Models) and NNET (Neural Networks Mod‐ evaluate the conservation status of Copris species, els). I approached methodological issues concern‐ we took into account the size of protected sites as ing the difficulties associated with predicting the well as the values of habitat suitability in each distribution of species when reliable absence data protected natural site and Natura 2000 network. are not available, and explored the possibilities of We found that Copris species were poorly con‐ SDMs as a tool for conservation of endangered served in the previous protected sites network: and threatened Iberian invertebrates. In this res‐ for C. hispanus only two protected sites measured pect, I explored the applications of SDM to esti‐ around 30 km2, and for C. lunaris a single area mate species ranges, identify suitable habitats and measured 183 km2. However, protection provided the primary factors affecting species’ distribution by Sites of Community Importance (SCIs) seems to in order to assess the conservation status of improve the general conservation status of these threatened invertebrates. species in CM because the area and connectivity Dung beetle populations, which are in de‐ of protected sites have been increased substan‐ cline in the Iberian Peninsula, play a critical eco‐ tially. logical role in extensive pasture ecosystems by Chefaoui and Lobo (2008) assessed the ef‐ recycling organic matter. We delimited the poten‐ fects of pseudo‐absences on model performance tial distribution of the two species of Copris when reliable absence data are not available. We (Coleoptera, Scarabaeidae) that inhabit the Ibe‐ compared seven procedures to generate pseudo‐ rian Peninsula using ENFA (Chefaoui et al. 2005). absence data to be used in GLM‐logistic regressed ENFA is a presence‐only method that compares models. These pseudo‐absences were selected the environmental values of the localities where randomly or by means of presence‐only methods the species has been observed with respect to the (ENFA and MDE) to model the distribution of a environmental values of the territory studied threatened endemic Iberian moth species (Hirzel et al. 2002). We explored the environ‐ (Graellsia isabelae). Our purpose was to show the mental niche occupied by each species in a small possibility of achieving different forecasted distri‐ region, the Community of Madrid (CM), to restrict butions depending on the method and the thresh‐ the role of dispersal constraints discriminating old used to select these pseudo‐absences. possible areas of co‐occurrence and identifying The results showed that the pseudo‐ the specific environmental characteristics of each absence selection method greatly influenced the species. We identified that solar radiation and the percentage of explained variability, the scores of presence of calcareous soils are critical to the the accuracy measures and, most importantly, the

102 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 Rosa M. Chefaoui predicted range size. As we extracted pseudo‐ tris and P. nigra located both inside and near to absences from environmental regions further SCIs, we suggested that the reintroduction of the from the optimum established by presence data, species in these habitats could improve its conser‐ the models obtained better accuracy scores, and vation. over‐prediction increased. Conversely, the profile To understand the limitations and possibili‐ techniques that generated wider unsuitable areas, ties of SDM techniques, we evaluated the effects produced functions with lower percentages of of species’ traits and data characteristics on the explained deviance and poorer accuracy scores, accuracy of SDMs for red‐listed invertebrates but more restricted predictive distribution maps, (Chefaoui et al. 2011). We applied three SDM similar to the observed distribution. The random techniques (GAM, GLM and NNET) using pseudo‐ selection of pseudo‐absences generated the most absences to model the distribution of 20 threa‐ constrained predictive distribution map. tened Iberian invertebrates. We correlated the Based on results of the aforementioned accuracy of the obtained models with several data work, we identified the environmental variables characteristics and species’ ecological traits. We most relevant for explaining the distribution of examined two data characteristics, the amount of Graellsia isabelae and assessed this species’ con‐ data (N) and the relative occurrence area (ROA), servation status (Chefaoui and Lobo 2007). We and both significantly affected the accuracy of the modeled the potential distribution of the insect by models. Greater AUC values and higher sensitivity performing GLM with pseudo‐absence data se‐ scores were obtained from samples for which lected from an ENFA model. We found that the there were more than 200 records. In general, best predictor variables were summer precipita‐ species whose distributions were most accurately tion (ranging from 1250 mm to 3250 mm), aridity, modelled were those with a greater sample size or and mean elevation. This species prefers habitats smaller ROA. In addition, species related to habi‐ with mid‐range mountain conditions. With respect tats that are problematic to detect using GIS data, to host plants, the presence of G. isabelae was such as riparian or humid areas, seemed to be associated mainly with Pinus sylvestris and P. ni‐ more difficult to predict. gra. Moreover, we found 8 areas exclusively in Summary the eastern Iberian territory, and a larger unoccu‐ The performance of SDMs depends on the type of pied habitat in the western Iberian Peninsula, indi‐ data and the characteristics of the species. Pres‐ cating that this species is probably not in equili‐ ence‐only methods (ENFA and MDE) achieved brium with its environment because of historical worse validation results and overpredicted more factors (Chefaoui and Lobo 2007). We sug‐ than techniques using pseudo‐absences. Never‐ gested that the current distribution of the species theless, presence‐only methods can be very useful was associated with the dynamism of its host for obtaining pseudo‐absences and discovering plants during glacial periods of the Holocene, the environmental response of species. The when the forests of Pinus sylvestris decreased method of pseudo‐absence selection strongly de‐ strongly in the northwestern part of the penin‐ termined the predicted range size, generating dif‐ sula. After analyzing the possibility of connectivity ferent model predictions along the gradient be‐ and fragmentation of the eight populations delim‐ tween potential and realized distributions. There ited as well as the degree of protection of G. isa‐ is an added difficulty in obtaining predictions that belae on the SCIs, we found that the SCIs under closely approximate the realized distribution of protection did not seem sufficient to maintain cu‐ species under non‐equilibrium conditions, be‐ rrent populations. Moreover, our study rejected cause both presence and absence data may be the idea that the species was expanding its range possible under similar environmental conditions. due to reforestation. Because the conservation of Irrespective of the approach used, species’ distri‐ G. isabelae depends on the forests of Pinus sylves‐ butions are modelled more accurately when sam‐ frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 103 SDM applied to invertebrate conservation ple sizes are larger. Species in habitats that are status of the silver‐spotted skipper butterfly difficult to detect using GIS data, such as riparian (Hesperia comma) in Britain: a metapopulation success story. Biological Conservation, 124, 189– species, thus may tend to be more difficult than 198. most to predict. Elith, J. & Leathwick, J.R. (2007) Predicting species’ dis‐ tributions from museum and herbarium records Availability of thesis using multiresponse models fitted with multi‐ variate adaptive regression splines. Diversity Printed and PDF copies are available in the Sci‐ and Distributions, 13, 165–175. ence Faculty Library, Universidad Autónoma de Engler, R., Guisan, A. & Rechsteiner, L. (2004) An im‐ Madrid (http://biblioteca.uam.es/ciencias/). A proved approach for predicting the distribution PDF copy is also available at request from the au‐ of rare and endangered species from occurrence thor. and pseudo‐absence data. Journal of Applied Ecology, 41, 263–274. Hill, J.K., Thomas, C.D., Fox, R., Telfer, M.G., Willis, S.G., Acknowledgements Asher, J. & Huntley, B. (2002) Responses of but‐ I would like to thank my two supervisors, Jorge M. terflies to twentieth century climate warming: implications for future ranges. Proceedings of Lobo and Joaquín Hortal for their support and en‐ the Royal Society, 269, 2163–2171. couragement. Hirzel, A., Hausser, J., Chessel, D. & Perrin, N. (2002) Ecological‐Niche Factor Analysis: How to com‐ References pute habitat‐suitability maps without absence data? Ecology, 83, 2027–2036. Anderson, R.P., Lew, D. & Peterson, A.T. (2003) Evaluat‐ ing predictive models of species’ distributions: Hortal, J., Borges, P.A.V., Dinis, F., et al. (2005) Using criteria for selecting optimal models. Ecological ATLANTIS – Tierra 2.0 and GIS environmental Modelling, 162, 211–232. information to predict the spatial distribution and habitat suitability of endemic species. Direc‐ Araújo, M.B., Lobo, J.M. & Moreno, J.C. (2007) The ef‐ ção Regional de Ambiente and Universidade dos fectiveness of Iberian protected areas in con‐ Açores, Horta, Angra do Heroísmo and Ponta serving terrestrial biodiversity. Conservation Delgada, Horta, Faial. Biology, 21, 1423–1432. Hortal, J., Lobo, J.M., & Jiménez‐Valverde, A. (2007) Brown, J.H. & Lomolino, M.V. (1998) Biogeography, 2nd Limitations of biodiversity databases: case study edn. Sinauer Press, Sunderland, Massachusetts. on seed‐plant diversity in Tenerife (Canary Is‐ Chefaoui, R.M., Hortal, J., & Lobo, J.M. (2005) Potential lands). Conservation Biology, 21, 853–863. distribution modelling, niche characterization Hortal, J., Jiménez‐Valverde, A., Gómez, J.F., Lobo, J.M., and conservation status assessment using GIS & Baselga, A. (2008) Historical bias in biodiver‐ tools: a case study of Iberian Copris species. Bio‐ sity inventories affects the observed realized logical Conservation, 122, 327–338. niche of the species. Oikos, 117, 847–858. Chefaoui, R.M. & Lobo, J.M. (2007) Assessing the con‐ Lobo, J.M., Verdú, J.R. & Numa, C. (2006) Environ‐ servation status of an Iberian moth using mental and geographical factors affecting the pseudo‐absences. The Journal of Wildlife Man‐ Iberian distribution of flightless Jekelius species agement, 8, 2507–2516. (Coleoptera: Geotrupidae). Diversity and Distri‐ Chefaoui, R.M. & Lobo, J.M. (2008) Assessing the ef‐ butions, 12, 179–188. fects of pseudo‐absences on predictive distribu‐ Lobo, J.M., Baselga, A., Hortal, J., Jiménez‐Valverde, A., tion model performance. Ecological Modelling, & Gómez, J.F. (2007) How does the knowledge 210, 478–486. about the spatial distribution of Iberian dung Chefaoui, R.M. (2010) Modelos predictivos aplicados a beetle species accumulate over time? Diversity la conservación de invertebrados protegidos and Distributions, 13, 772–780. ibero‐baleares. Ph.D. Thesis. Universidad Autó‐ Lobo, J.M., Jiménez‐Valverde, A., & Hortal, J. (2010) noma de Madrid, Departamento de Biología, The uncertain nature of absences and their im‐ Facultad de Ciencias, 196 pp. portance in species distribution modelling. Eco‐ Chefaoui, R.M., Lobo, J.M. & Hortal J. (2011) Effects of graphy, 33, 103–114. species’ traits and data characteristics on distri‐ Lomolino, M.V. (2004) Conservation biogeography. bution models of threatened invertebrates. Ani‐ Frontiers of Biogeography: new directions in the mal Biodiversity and Conservation, 34, (in press). geography of nature (ed. by M. V. Lomolino and Davies, Z.G., Wilson, R.J., Brereton, T.M. & Thomas, L. R. Heaney), pp. 293–296. Sinauer Associates, C.D. (2005) The re‐expansion and improving Sunderland, Massachusetts.

New, T.R. (1998) Invertebrate surveys for conservation. Verdú, J.R. & Galante, E., eds. (2009) Atlas de los Inver‐ Oxford University Press, New York. tebrados Amenazados de España (Especies en Newbold, T. (2010) Applications and limitations of mu‐ peligro crítico y en peligro). Dirección General seum data for conservation and ecology, with para la Biodiversidad, Ministerio de Medio Am‐ particular attention to species distribution mod‐ biente, Madrid, 340 pp. els. Progress in Physical Geography, 34, 3–22. Zaniewski, A.E., Lehmann, A. & Overton, J.M. (2002) Pearce, J. & Boyce, M.S. (2006) Modelling distribution Predicting species spatial distributions using and abundance with presence‐only data. Journal presence‐only data: a case study of native New of Applied Ecology, 43, 405–412. Zealand ferns. Ecological Modelling, 157, 261– Peterson, A.T., Ball, L.G. & Cohoon, K.P. (2002) Predict‐ 280. ing distributions of Mexican birds using ecologi‐ cal niche modelling methods. Ibis, 144, E27–E32. Edited by Richard Pearson

frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 105 opinion and perspectives ISSN 1948‐6596 opinion Political erosion dismantles the conservation network existing in the Canary Islands José María Fernández‐Palacios and Lea de Nascimento Island Ecology and Biogeography Group, Instituto Universitario de Enfermedades Tropicales y Salud Pública de Canarias (IUETSPC), Universidad de La Laguna (ULL), Avda. Astrofísico Francisco Sánchez s/n, 38206, La Laguna, (Tenerife), Spain e‐mail: [email protected]; http://webpages.ull.es/users/jmferpal

Abstract. The outstanding nature of the Canary Islands has been recognized by European, national and regional administrations since the arrival of democracy in Spain. Forty‐five per cent of its emerged terri‐ tory has been declared as Natural Protected Areas, four Canarian National Parks were included within the Spanish network, more than 200 endemics were listed in the Spanish catalogue of endangered species, and 450 species were listed in the Canarian catalogue of protected species. However, in recent years, po‐ litical decisions have started dismantling this splendid conservation network, which impedes construction of large infrastructure, golf courses and resorts, despite the advice of the scientific community. Canarian nature is now facing two threats: delisting and downgrading of numerous endangered species, and trans‐ fer of the management of Canarian National Parks to the regional administration. Keywords: Biodiversity loss, endangered species, National Parks, natural protected areas, political corrup‐ tion, scientific community, species delisting

Recently the Canarian Parliament has approved a Changes in the environmental legislation of new version of the Canarian catalogue of pro‐ the Canary Islands entail a serious threat to the tected species (see Box 1) that reduces substan‐ nature of this region of biogeographical interest tially both the number of species included (from (Francisco‐Ortega et al., 2000; Juan et al., 2000; 466 species in the 2001 list to 361 species in the Fernández‐Palacios & Whittaker, 2008). Thus, we 2010 list) and the protection afforded (from 381 believe it is important to share our appraisal of threatened species to 160, and from 85 protected the current situation with the international scien‐ species to 18). These reductions have been widely tific community. criticized by environmental NGOs and the local Within the new revised catalogue a com‐ scientific community1, mainly due to the absence pletely new criterion for protection has emerged of a rigorous scientific process in its development. “especies de interés para los ecosistemas canar‐ Although certainly the first version of the cata‐ ios” (literally: “species of interest for Canarian eco‐ logue could be improved, the main reasons be‐ systems”), comprising 152 species (see Box 1). The hind the new revisions were not conservation is‐ phrase is poorly chosen. It is supposed to apply sues but rather strictly political. The reasons may only to endangered species, consequently the fre‐ include, for instance, the development of large quent and abundant species which usually struc‐ infrastructures, such as industrial harbours and ture and dominate the ecosystems are explicitly golf courses, which until the revisions were forbid‐ not listed, leading to a curious paradox: the Ca‐ den due to their impacts on protected species in‐ narian pine (Pinus canariensis) is not a species of cluded in the original version of the Canarian cata‐ interest for the Canarian pine forest, the logue. Macaronesian Laurel (Laurus novocanariensis) is

1. See different reactions at http://www.nodescatalogacion.com, http://www.wwf.es, http://www.greenpeace.org, http://www.atan.org, http://www.ecologistasenaccion.org, http://especiesamenazadascanarias.blogspot.com, http://ecooceanos.blogspot.com, http://www.seo.org, .

106 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 José María Fernández‐Palacios and Lea de Nascimento Box 1 Law 4/2010, June 4, of the Canarian Catalogue of Protected Species (see the original Spanish text at http://www.gobiernodecanarias.org/boc/2010/112/) Article 3. Canarian protected species 2) Species of interest for Canarian ecosystems The Canarian Catalogue of Protected Species will also include “species of interest for Canarian ecosys‐ tems" which are those that, without being listed in the threatening situations above (endangered or vul‐ nerable), are worthy of particular attention for its ecological significance in areas of the Canarian Network of Natural Protected Areas or Natura 2000 network. 2. Effects of inclusion in the Catalogue b) The legal regime for protection of “species of interest for Canarian ecosystems" will be applicable only in the territory of the Canarian Network of Natural Protected Areas or Natura 2000 Network. To this end, applicable measures shall be provided by the management plans of Natural Protected Areas and Habitats of the Natura 2000 Network in which they are located. Such plans shall include the determinations, con‐ trol and monitoring to ensure effectiveness of protection, or where applicable, the justification that there is no need for plans. (...) In the case of actions promoted by reasons of public interest and priority affect‐ ing the “species of interest for Canarian ecosystems" these actions could be possible as long as they do not affect the ecosystem substantially, under the terms in paragraphs 4 to 7 of the Article 45 of the Law 42/2007, December 13, of Natural Heritage and Biodiversity. not a species of concern for the Laurel forest, and The new law could have negative implica‐ so on. This is not to say that the most common tions for conservation biogeography, and this can structuring species of the Canarian ecosystems be illustrated with some examples of the Canarian have to be included in the catalogue, but we flora and fauna. The endemic legume Cicer ca‐ would like to draw attention to the inadequacy of nariensis, previously considered as vulnerable in the concept. the 2001 Canarian catalogue, is now included un‐ But this conceptual shortcoming pales in der the criterion species of interest. From its 12 comparison with the real repercussion of the new locations (ten in La Palma and two in Tenerife), criterion, which is that those species listed here the six populations in the North of La Palma2 are are only protected if present in an already desig‐ outside NPAs and therefore unprotected accord‐ nated Natural Protected Area (NPA). (In the Ca‐ ing to the new law. Metapopulation dynamics in naries, that means in either the Canarian Network this species could be affected by this new criterion of NPAs or the European Union Natura 2000 Net‐ if source populations within these northern loca‐ work, which overlap extensively). If a listed spe‐ tions are threatened, endangering sink popula‐ cies, for instance the woodcock (Scolopax rusti‐ tions included in NPAs. The same could apply to cola) or the coot (Fulica atra) which are both in‐ the Abalone or Canarian clam (Haliotis tuberculata cluded under the new criterion, dwells within the ssp. coccinea) or the Sea Horse (Hippocampus hip‐ limits of the protected area they are safe; but if pocampus). Both are marine species with sparse any birds cross those limits (which are not that populations in the meso‐ and infra‐littoral, which obvious to birds, unfamiliar as they are with GIS), do not always coincide with the geographical loca‐ they can be shot legally by hunters. The same in‐ tion of the marine Special Areas for Conservation, consistency affects, for instance, ca. 10 endemic which occupy mainly leeward fringes on the Archi‐ species of sea lavenders (Limonium spp.) pro‐ pelago’s coasts. Collection and capture of both tected in certain ravines, but not in others. species is prohibited by the Regulation of the Fish‐

2. According to the evaluation of this species by the Canarian Government (Servicio de Biodiversidad 2009), there are six population nuclei in the North of La Palma, distributed in three locations more than 10 km distant one from each other. frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 107 Canarian conservation network dismantled eries Law of the Canary Islands, but their inclusion In fact, many of the species of interest come from in the new criterion may lead to confusion on the the former list of susceptible species or are down‐ fishing ban in populations outside of the reserve graded threatened species. However in the former networks. criterion there were no restrictions in the protec‐ The case of the sea grass Cymodocea tion, such as the location or not in a NPA, and the nodosa is of particular interest for two reasons; main consideration to include a species was that this species structures a community (“sebadales”), its habitat was threatened, in regression, frag‐ considered as Natural Habitat of Community In‐ mented or limited. The previous criterion for pro‐ terest by the Habitats Directive, and its presence tection was much more appropriate if we think in the littoral zone is one of the main obstacles to about the design of the Canarian Network of the construction or enlargement of harbours. The NPAs. Unfortunately the Canarian Network was most recent is the Puerto de Granadilla, where not based on a thorough analysis of metapopula‐ conservation of a European priority ecosystem tion dynamics, genetic diversity or viability of comes into conflict with European funding of a populations, but simply in protecting less de‐ large infrastructure. The sebadales are a key com‐ graded remnants of communities that were still munity from an ecological point of view as they available. As in many parts of the world, reserves play an important role in the carbon cycle, stabi‐ were not designed to meet the principles of sys‐ lize sandy soils, export biomass and act as a fish tematic conservation planning needed to achieve nursery area (Barberá et al. 2005). The latter char‐ representativeness and persistence of biodiversity acteristic is also very important for the sustainabil‐ (Margules and Pressey 2000). The situation fur‐ ity of local fisheries. Also, the marine meadows of ther worsens in the Canaries when data, trends C. nodosa in the Canary Islands and Mauritania and viability of populations are almost unknown. are the most extensive examples at the species’ The Canarian Network is largely protecting southern limit and compromising them may there‐ species from marginal populations. Moreover, the fore lead to range contraction. The construction of protection of species present only in the current Puerto de Granadilla will severely damage one of Reserve Network inhibits re‐establishment of the most genetically diverse patches of sebadales original distributions. A good example is the laurel in the Archipelago (Alberto et al. 2008). In 2009, forest in Anaga Rural Park, which nowadays is the as a precautionary measure, the Superior Court of best representation of this forest type in Tenerife Justice of the Canary Islands suspended the pro‐ yet still an impoverished fraction of its past distri‐ posal submitted by the Canarian Government, the bution throughout the windward slope of the is‐ Port Authority and the Canarian Company of Gas land. From the point of view of mitigating the ef‐ Transportation, to delist C. nodosa3. Currently, the fects of global change, vulnerability of certain spe‐ European Courts have declared admissible the cies outside the Network would hinder altitudinal complaint filed by the NGO Ecologistas en Acción migration, especially when ecological corridors are asking for the public release of documents that not included in the design of NPAs. included alternatives to the construction of the The practice of protecting taxa only in NPAs harbour (including a renewal of the infrastruc‐ is already working in Catalonia (the only prece‐ tures of already existing harbours), that were hid‐ dent in Spain). The Catalonian Plan of Areas of den from the European Commission by Spain’s Natural Interest includes species of flora and National Government. fauna strictly protected in designated areas. To This controversial criterion — especies de our knowledge no cases of the failure of these interés para los ecosistemas canaries — is an ad‐ practices or public disapproval have been re‐ aptation of the criterion “species susceptible to ported there, but we suspect that the species with habitat disturbance”, from the previous catalogue. restricted protection in the Catalonian NPA Net‐

3. See news in http://www.laprovincia.es. 4. See http://www.laopinion.es, http://www. ecologistasenacción.org.

108 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 José María Fernández‐Palacios and Lea de Nascimento work were not demoted from higher protection. geographical area. After decentralization of the In theory, the main aim of the existence of re‐ Spanish State with the arrival of the democracy, gional catalogues is ensuring the protection of the NPs were simultaneously co‐managed by the particular species that are not considered by the Central Government (Madrid) and the Regional National Catalogue. On the other hand several Governments. However, the Spanish Constitu‐ authors have questioned and analysed the effec‐ tional Court now has determined that NPs man‐ tiveness of NPAs Networks in biodiversity conser‐ agement is exclusively a matter for the Regional vation (Jaffre et al. 1998, Rodrigues et al. 2004) Governments. Consequently the Central Govern‐ and concluded that reserve networks are geo‐ ment has transferred all management to the re‐ graphically and taxonomically unbalanced leaving gions. In the case of the Canarian archipelago, this a big proportion of endemic and threatened spe‐ management was intended to be subsequently cies unprotected. delegated to the respective island Councils This way of thinking may function well (“Cabildos”) in 2012, although recently the new when protecting a resource, for instance marine deputy of Environment of the Canarian Govern‐ sanctuaries are intended to increase catch in ment expressed her intention to discuss again this neighbouring areas outside, and this works com‐ transfer and to limit the management of the is‐ petently in the Canaries’ Marine Reserves with land Councils in the NPs. Fishery Interest, but is nonsensical when the aim The transfer to regions is not inherently of the declaration is to protect a threatened spe‐ bad, and for instance would work exceptionally cies. If a species is protected when within a NPA, well in Northern European countries. The problem but unprotected when beyond the area, what is is not the law but how it is developed when the really achieved in terms of protection? Might it be main political parties that govern in the Canary too cynical to suggest the greatest achievement Islands show no interests in conservation, and an would be the political goal of inflating the number alarming number of its politicians, including some of species included in the catalogue thus reducing who have significant responsibilities in conserva‐ the number of critics of delisting? Despite numer‐ tion, have been charged with environmental ous public protests and the clear opposition of the crimes5. Although some implications of decentrali‐ majority of the Canarian scientific community, the zation should be positive, for instance the creation new catalogue was presented by the leading po‐ of regional lists and plans considering the particu‐ litical force in the Regional Parliament. These lars of each NP or the proximity to local specialists kinds of conflicts are not exclusive to the Canary and technicians with a wider knowledge of the Islands and are nowadays taking place in different region, the result is exactly opposite. With the regions of the world (Possingham et al. 2010, proximity of the management centres to the NPs, Metzger et al. 2011). the likelihood of patronage and corruption seems If the delisting itself is not of sufficient con‐ likely to increase while unification of conservation cern, other news makes the outlook even bleaker. criteria across the archipelago’s four NPs seems The Canaries harbour four of the 13 National destined to decrease, especially if the different Parks (NPs) in Spain – Cañadas del Teide island Councils are governed by different political (Tenerife), Caldera de Taburiente (La Palma), Ti‐ parties, which is currently the case. In addition, manfaya (Lanzarote) and Garajonay (La Gomera) joint management of the NPs and the other NPAs – despite representing only 1.5% of the country’s in each island would dilute the rigor and resources

5. See press references in http://www.abc.es/20100322/canarias‐canarias/tres‐imputados‐coronan‐nueva‐ 20100322.html (last accessed August/2011); http://www.canarias‐semanal.com/elhierro.html (last accessed Au‐ gust/2011); http://www.eldia.es/2011‐04‐13/CANARIAS/5‐Es‐frecuente‐alcaldes‐esten‐imputados‐delitos‐ urbanisticos.html (last accessed August/2011); http://www.elpais.com/articulo/espana/corrupcion/presenta/ elecciones/elpepiesp/20110410elpepinac_1/Tes (last accessed August/2011); http://www.europapress.es/islas‐ canarias/noticia‐imputados‐canarias‐logran‐mantenerse‐instituciones‐20110524094822.html (last accessed Au‐ gust/2011) . frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 109 Canarian conservation network dismantled dedicated to NPs. Considering that budgets are Barberá, C., Tuya F., Boyra C., Sanchez‐Jerez P., Blanch not fixed this would imply that funding to manage I. & Haroun R.J. (2005) Spatial variation in the structural parameters of Cymodocea nodosa the NPs could eventually be used in other tasks, seagrass meadows in the Canary Islands: a mul‐ more consistent with the "needs of the moment". tiscaled approach. Botanica Marina, 48, 122– A recently created Commission of Canarian NPs, 126. constituted mainly of politicians and with only two Fernández‐Palacios, J.M. & Whittaker, R. (2008) Canar‐ advocates for environmental issues, left aside the ies. An important biogeographical meeting present directors and conservators of the NPs. It place. Journal of Biogeography, 35, 379–387. Francisco‐Ortega, J., Santos‐Guerra, A., Kim, S.C. & could also happen that once transferred to the Crawford, D. (2000) Plant genetic diversity in the Councils, the election of new directors will not Canary Islands: A conservation perspective. consider the balance between conservation and American Journal of Botany, 87, 909–919. management skills that such position requires. Jaffre, T., Bouchet, P., Veillon, J.M. (1998) Threatened The island Councils are already in charge of plants of New Caledonia: Is the system of pro‐ tected areas adequate? Biodiversity and Conser‐ the management of the Canarian Network of vation, 7, 109–135. NPAs. While some of these areas have been ac‐ Juan, C., Emerson, B.C., Oromí, P. & Hewitt, G.M. tively managed others lack any type of control. (2000) Colonization and diversification: towards The situation of similar NPAs varies among islands a phylogenetic synthesis for the Canary Islands. and for most the action plans have been partially Trends in Ecology and Evolution, 15, 104–109. Margules, C.R. & Pressey, R.L. (2000) Systematic con‐ or barely fulfilled, so that nowadays (more than servation planning. Nature, 405, 243–253. ten years after its declaration) it is still easy to find Metzger, J.P., Lewinsohn, T.M., Joly, C.A., Verdade, dumps, illegal constructions, invasive species, to‐ L.M., Martinelli, L.A., Rodrigues, R.R. (2011) Bra‐ gether with other potential emerging threats. De‐ zilian Law: Full Speed in Reverse? Science, 329, spite the capacity and good work of environ‐ 276–277. mental technicians, who struggle with budget cuts Possingham, H.P. et al. (2010) Open letter to the Prime Minister and Leader of the Opposition, Science every year, the Councils have demonstrated a tra‐ supporting marine protected areas, signed by jectory of inefficiency and lack of commitment to 152 Australian scientists. Available from http:// the management of NPAs. Within the new Ca‐ www.ecology.uq.edu.au/docs/Marine% narian NPs framework, the rabbits will receive the 20Reserve%20Scientist%20Ltr% responsibility of taking care of the lettuces. 2018Aug2010.pdf (last accessed October/2011) Rodrigues, A.S.L., Andelman, S.J., Bakarr, M.I., et al.

(2004) Effectiveness of the global protected area Acknowledgements network in representing species diversity. Na‐ We would like to thank Rafael Loyola and three ture, 428, 640–643. anonymous reviewers for their comments on the Servicio de Biodiversidad (2009). Evaluación de espe‐ manuscript. We are also grateful to the editorial cies catalogadas de Canarias: Cicer canariensis [Ciccan 06/2009]. Consejería de Medio Ambien‐ board of Frontiers in Biogeography for their help te y Ordenación Territorial, Gobierno de Cana‐ improving this paper. rias, Las Palmas de Gran Canaria. Available at http://www.gobcan.es/cmayot/ References medioambiente/medionatural/biodiversidad/ especies/especies_protegidas_amenazadas/ Alberto, F., Massa, S., Manent, P., Diaz‐Almela, E., Ar‐ naud‐Haond, S., Duarte, C.M. & Serrão, E.A. (2008) Genetic differentiation and secondary Edited by Joaquín Hortal & Michael N Dawson contact zone in the seagrass Cymodocea nodosa across the Mediterranean–Atlantic transition region. Journal of Biogeography, 35, 1279–1294.

You 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).

110 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 ISSN 1948‐6596 opinion and perspectives perspective The causes and biogeographical significance of species’ rediscovery Richard J. Ladle1,2,*, Paul Jepson2, Ana C. M. Malhado1, Steve Jennings3 and Maan Barua2 1. Institute of Biological and Health Sciences, Federal University of Alagoas, Maceió, AL, Brazil. 2. School of Geography and the Environment, University of Oxford, South Parks Road, Oxford, OX1 3QY, United Kingdom. 3. Oxfam GB, Oxfam House, John Smith Drive, Oxford, United Kingdom. *Author for correspondence: Dr Richard J. Ladle, Institute of Biological and Health Sciences, Federal Uni‐ versity of Alagoas, Praça Afrânio Jorge, s/n, Prado, Maceió, AL, Brazil, 57010‐020. e‐mail: [email protected]; http://www.geog.ox.ac.uk/staff/rladle.html

Abstract. The rediscovery of a species that was putatively considered to be extinct can provide valuable data to test biogeographical hypotheses about population decline and range collapse. Moreover, such rediscoveries often generate much‐needed publicity and additional funds for the conservation of rare species and habitats. However, like extinction, rediscovery is challenging to define. In this perspective we argue that the ‘loss’ of a species and its subsequent rediscovery can be understood in terms of the interplay among four socio‐ecological factors: (1) the state of knowledge of species loss and rediscovery; (2) the presence of people and/or organizations with the interest, motivation, resources, skills and tech‐ nology to find target species; (3) the accessibility of the areas, habitats or sites where the species are thought to survive; and (4) the ease with which a species can be located when it is present within a habi‐ tat. Thus, species are ‘lost’ from scientific knowledge for different reasons and, consequently, not all rediscoveries are equally significant for biogeographical research or conservation. Indeed, rediscoveries of species that underwent a well documented decline and disappearance – and are therefore of greatest potential importance for both conservation and biogeographical research – appear to be poorly repre‐ sented in the literature compared to rediscovered species that were only known from a handful of mu‐ seum specimens. Thus, carefully distinguishing between the causes of temporal gaps in zoological re‐ cords is essential for improving the utility of rediscovery data for biogeographical research and conser‐ vation practice. Keywords: extinction, range collapse, rarity, critically endangered, monitoring

Introduction tions to 18 countries seeking to locate 40 species Rediscoveries of putatively extinct species are of not seen for a decade or more (Conservation In‐ great potential interest to both conservationists ternational 2010) – at the time of writing 12 spe‐ and biogeographers (Crowley 2011). For the for‐ cies have been rediscovered. Moreover, since re‐ mer, ‘rediscovery’ can be a considerable conserva‐ discovered species are typically exceedingly rare tion policy and publicity asset (Ladle and Jepson and geographically localized, new knowledge on 2008, Ladle et al. 2009) – as testified by recent population status and distribution supports effec‐ global initiatives: in 2009 BirdLife International tive conservation interventions. Finally, rediscov‐ launched a “global bid to try to confirm the con‐ eries remove uncertainty from extinction risk as‐ tinued existence of 47 species of bird that have sessments; a confirmed new record moves the not been seen for up to 184 years” (BirdLife Inter‐ species from ‘extinct’ or ‘probably extinct’ and national 2009). The following year Conservation into an IUCN threat (or data deficient) category. International launched its “Search for lost Frogs” For biogeographers, species rediscovery has both which involves a dedicated campaign and expedi‐ a practical and conceptual significance. From the frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 111 rediscoveries in biogeography practical perspective, the rediscovery of a species covery reflects the temporary absence of such that has gone unrecorded for a long period of records. Moreover, rediscovery is the proof re‐ time improves geographical knowledge about quired to refute a hypothesis of extinction. Given some of the world’s rarest species, helping to ad‐ the close conceptual linkage between the con‐ dress the Wallacean shortfall – the inadequacy of cepts of rediscovery and extinction it is interesting our knowledge of the geographical distributions of that, until recently, there have been so few stud‐ species (Lomolino et al. 2006, Riddle et al. 2011). ies linking patterns of rediscovery to contempo‐ The shortfall can often be extreme, with a species rary theories of population decline and extinction. known from just one or a few museum specimens One impediment to such research is the lack of a collected decades or even centuries earlier. These systematic approach to species rediscoveries that species are sometimes incorrectly assumed or de‐ allow scientists to identify cases of rediscovery clared extinct, a phenomenon which Ladle and that have biogeographical or conservation signifi‐ Jepson (2008) refer to as a Wallacean extinction. cance, and which can be subject to meaningful As we discuss later, these extreme examples of analysis. Here, we propose a conceptual frame‐ the Wallacean shortfall are amongst the most fre‐ work for understanding and analyzing species re‐ quently rediscovered species. discovery, based on the social, institutional and More recently, biogeographers have started ecological factors that created the temporal gap in to use information on species rediscoveries to test occurrence data. We believe that formalizing the theories of population decline and range collapse concept of rediscovery in this way has the poten‐ under anthropogenic disturbance (Fisher 2011a,b; tial to create new measures of the state of knowl‐ Fisher and Blomberg 2011). The underlying idea is edge of the world’s rarest species, provide a quan‐ both simple and elegant: the location of a redis‐ tifiable metric to support existing endangerment covered species relative to its historical range re‐ categorizations, and would help to maintain the flects the pattern of range collapse. Thus, if an‐ culture of biogeographical exploration that con‐ thropogenic pressures (e.g. unsustainable exploi‐ tributes to the datasets that underpin global con‐ tation) are strongest at the periphery (Channel servation target‐setting, advocacy and monitoring. and Lomolino 2000) the rediscovery will most likely be made near the centre of the historic Conceptual framework range. Diana Fisher’s (2011a) study of 67 species The ‘loss’ of a species and its subsequent rediscov‐ of rediscovered mammals found a number of clear ery can be conceptualized as a result of the inter‐ trends, although these tended to be dependent play among four socio‐ecological aspects of redis‐ upon the ecology of the species. For example, one covery (schematically illustrated in Figure 1): (1) of the strongest patterns observed was that redis‐ the state of knowledge of species loss and redis‐ coveries were generally made at higher elevations covery; (2) the presence of people and/or organi‐ than the original record (excluding mountain‐top zations with the interest, motivation, resources, and coastally restricted species). This provides skills and technology to find target species; (3) the some support for the hypothesis that higher ele‐ accessibility of the areas, habitats or sites where vations can sometimes provide ecological refugia the species are thought to survive; and (4) the (Towns and Daugherty 1994) and fits with the fre‐ ease with which a species can be located when it quently observed pattern of habitat destruction is present within a habitat. It should be noted that and population extinction progressing from low to although these factors potentially apply to all high altitudes (Triantis et al. 2010). ‘lost’ taxa, owing to issues of historical data qual‐ However, like extinction, rediscovery is ity, funding and the culture of scientific explora‐ challenging to define. This should not be surpris‐ tion, rediscovery research has focused almost ex‐ ing since rediscovery and extinction are conceptu‐ clusively on herptiles, birds and mammals (cf. ally intertwined; extinction is the permanent ab‐ Scheffers et al. 2011). sence of current and future records while redis‐

Knowledge of ‘lost’ species Stresemann (1931) had the effect of making the Enormous advances have been made over the last species of “only marginal, regional interest” and 40 years in enumerating which species are appar‐ as a consequence “for many years [it] received ently ‘lost’. For example, BirdLife International has little attention” (p. 69). made significant investments in compiling new From the perspective of investigating range and authoritative assessments of threatened spe‐ changes, confounding different categories of re‐ cies using information from a variety of sources discovery could seriously influence research find‐ including amateur and university‐led research ex‐ ings. For example, we might expect that all other peditions and major reviews of existing museum things being equal, species whose habitat or range specimens. In particular, from the mid 1980s two has not been surveyed for a significant period of major regional Red List reviews were compiled for time and for which there are no strong reasons to the Americas (Collar et al. 1992) and Asia (Collar assume have become extinct (Table 1, category 4), et al. 2001), the findings of which were then fed are as likely to be rediscovered at the edge or cen‐ back to the BirdLife network of pioneering profes‐ tre of their historic range as are better‐known sional and amateur ornithologists (Tobias et al. species. Moreover, all four categories of rediscov‐ 2006, Butchart 2007). ery may contain species that were only known The knowledge of what is ‘lost’ is compli‐ from a small number of museum specimens – the cated, as rediscoveries can logically be split into rediscovery of which may tells us more about the four categories that reflect different degrees of history of biogeographical exploration than the uncertainty (and authority) about the continued ecology of decline and extinction. Indeed, Schef‐ existence of a target species (Table 1). An addi‐ fers et al. (2011) found that the majority of re‐ tional category could potentially be added to this cently claimed amphibian, bird and mammal re‐ typology to account for cases where an unre‐ discoveries represent first documentations since corded sub‐species is elevated to full species their original scientific description. It should also status. For example, the Sangihe Shrike‐thrush be noted that such rare species may have re‐ (Colluricincla sanghirensis) was rediscovered in mained unrecorded because of intrinsic biological 1985 but its status as a full species was only estab‐ characteristics (e.g. nocturnal habits, cryptic lished in 1999 (Rozendaal and Lambert 1999). colouration, etc.) rather than a lack of sampling Changes in taxonomic status may have profound effort and that these factors need to be carefully impacts on survey effort: according to Rasmussen untangled in any analysis of patterns of rediscov‐ et al. (2000), the demotion of the Sangihe White‐ ery (see McCarthy 2008; Fisher and Blomberg eye (Zosterops nehrkorni) to sub‐specific status by 2011).

Figure 1. The four major dimensions of species rediscovery (see text). frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 113 rediscoveries in biogeography

Type Rediscovery of… Example 1. a species declared extinct by an authori‐ The Pohnpei Starling (Aplonis pelzelni) was declared tative source extinct by the IUCN (1990) and rediscovered in 1995 (Buden 1996)

2. a species considered probably extinct by The Sao Tome Grosbeak (Neospiza concolor) was an authoritative source described as probably extinct by Greenway (1967) and rediscovered in 1991 (Sergeant et al. 1992) 3. a species believed to be still extant but According to the NGO BirdLife International the for which substantive searches over dec‐ Madagascar Serpent Eagle (Eutriorchis astur) was ades have drawn a blank. not definitely recorded between 1930 and 1993 de‐ spite considerable search‐effort within its habitat. 4. a species whose habitat or range had not The Chestnut‐bellied Flowerpiercer (Diglossa glorio‐ been surveyed for a significant period of sissima) was unrecorded for 38 years: since 2003 it time, but for which there is no real rea‐ has been recorded from three locations (Tobias et son to assume has become extinct al. 2006)

Table 1. A crude typology of species rediscovery based on decreasing level of certainty that the rediscovered species was extinct.

Perhaps the most important type of redis‐ Thus, the Black‐hooded Antwren (Formicivora covery for conservation is where a previously well erythronotos) was known only from a 19th Century known species undergoes a population decline, is type specimen, for which the type locality was lost from biogeographical knowledge, and is then probably incorrect, and which was also put in the rediscovered. A possible example is the Australian wrong genus. Balchon (2007) suggests that this Pygmy Blue‐tongue Lizard Tiliqua adelaidensis. led to researchers “looking in the wrong place, for This rather secretive lizard was relatively well the wrong sort of bird and listening for inappropri‐ known up to its disappearance in 1959; its redis‐ ate vocalizations”. Thus, ‘lost’ species can some‐ covery in 1992 (in the stomach of a snake) con‐ times turn up thousands of kilometres away from firmed that the species now has “a dramatically where they were last seen, or in completely differ‐ reduced geographical range” (Milne and Bull ent habitats. For example, the Large‐billed Reed 2000, p. 296). The rediscovery of the Ivory‐billed Warbler (Acrocephalus orinus) was previously Woodpecker (Campephilus principalis) (Fitzpatrick known from just a single specimen collected in et al. 2005) would be an even better example, ex‐ 1867 in the Sutlej Valley, Himachal Pradesh, India. cept that this rediscovery is increasingly looking However, a living specimen was trapped in March like a case of mistaken identity (Dalton 2005, 2006 at Laem Phak Bia, Phatchaburi Province, 2010, Stokstad 2007). The apparent scarcity of south‐west Thailand, over 3000 km from the type such rediscoveries (cf. Scheffers et al. 2011) locality (Round et al. 2007). The renewed interest strongly suggests that a species that undergoes a in this species led to the unearthing of ten new well documented decline and disappearance is museum specimens (Svensson et al. 2008) and, likely to be extinct. However, formally testing this shortly afterwards, to the discovery of a breeding hypothesis would require good information on population in north‐east Afghanistan (Timmins et population trends of rediscovered species prior to al. 2010). their original disappearance – data that rarely ex‐ ist for older cases of species loss. Institutional, scientific and technical capacity A final aspect of the knowledge needed to Even when a species is identified as possibly still find ‘lost’ species is the reliability of biogeographic extant, the institutional and technical capacity to information on where to search for the species. find it may not exist. Such capacity, at a global

114 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 Richard J. Ladle et al. level, has varied considerably over time and space ments and analysis of mitochondrial DNA and mi‐ in response to various cultural and ecological fac‐ crosatellites, to conclusively demonstrate the hy‐ tors. Most notably, the mainstreaming of biodiver‐ brid origin of the imperial pheasant (Lophura im‐ sity into international development following the perialis). This mysterious bird had first been cap‐ 1992 Earth Summit created many new sources of tured in 1924 when a single pair had been shipped funds and employment opportunities for scientists to the private aviary of Jean Delacour in France in less‐developed countries. With respect to birds, and was not seen again until one was trapped in this increase in local capacity coincided with the 1990 (Hennache et al. 2003). creation of BirdLife International in 1993. BirdLife It is not only advances in molecular biology emerged from the International Council for Bird that are facilitating rediscoveries. The ready avail‐ Preservation (founded in 1922) when its leaders ability of sophisticated audiovisual equipment has devised the compelling proposition of forming an been especially important in the evolution of bird international partnership, under a single name, surveying. Two such technological advances, the with smaller, national, bird‐orientated conserva‐ increased availability of less expensive sound‐ tion organizations (Jepson and Ladle 2010). More recording and playback equipment in the late generally, increased funding of expeditions by in‐ 1990s and the more recent internet‐based bird‐ ternational NGOs has probably been the driving sound archives, have dramatically increased the force behind the increasing frequency of rediscov‐ capacity of both amateurs and professionals to eries of various taxa (Scheffers et al. 2011). locate and identify rare and cryptic bird species. Other trends within science and conserva‐ Moreover, advances in the quality of cameras and tion also help determine the capacity and motiva‐ lenses, especially digital cameras and video re‐ tion that enables rediscoveries, especially the in‐ corders, have also been important in documenting troduction of new technology. For example, ad‐ and providing definitive proof of the existence of vances in molecular biology have made it much very rare species. For example, the New Zealand easier to genetically compare preserved type Storm Petrel (Pealeornis maoriana) was identified specimens in museums with contemporary mate‐ from the details on a digital image taken in 2003 rial collected directly or acquired from hunters or (Stephenson et al. 2008). It had previously been from rural markets. This has opened the way for known only from putative fossil material, and completely new ways of rediscovering lost spe‐ from three specimens collected in the 19th Cen‐ cies, where a fragment of hair or a faecal sample tury, 150 years before its rediscovery. may be sufficient to prove the continuing exis‐ tence of a species that has still not been physically Accessibility observed. Even if a species is extant and potential habitats An excellent example of such a technology‐ have been located, the species may not be found. aided discovery is provided by Pitra et al. (2006), Access to suitable habitat may be limited because who recently announced the continuing existence of political instability/restrictions, or simply the of the giant sable antelope (Hippotragus niger remoteness of potential sites. Although in the era variani), a sub‐species unique to Angola that was of cheap international air travel this is arguably feared extinct after almost three decades of civil less important, it may have played a critical role in war. They compared the mitochondrial DNA se‐ restricting the intensity of surveys and therefore quences derived from old museum specimens the rate of rediscoveries in many parts of the with samples extracted from dung samples re‐ globe. Examples of rediscoveries that were proba‐ cently collected in the field. Such remotely col‐ bly delayed, and possibly even caused, by political lected DNA evidence can also be used to discount instability include that of the Large‐billed Reed presumed discoveries or rediscoveries. For exam‐ Warbler in Afghanistan (see above) and the ple, Hennache et al. (2003) used a range of tech‐ Gabela Helmet‐shrike (Prionops gabela), rediscov‐ niques, including captive hybridization experi‐ ered in 2003 in Angola (Ryan et al 2004). frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 115 rediscoveries in biogeography

A closely related factor is a lack of commu‐ prisingly, between 1912 and 1990 there were no nication with remote and isolated rural communi‐ records of the Night Parrot until one was hit by ties who may already have knowledge of the con‐ traffic (Boles et al. 1994). tinued existence of a putatively extinct species, or of a species new to science. Thus, a productive Rediscoveries reconsidered route to increasing rediscoveries (and new species Given the very loose usage of the term discoveries) might be through better communica‐ ‘rediscovery’ and the varying factors, social and tion with remote tribes and communities whose ecological, that contribute to rediscoveries, both knowledge of local biodiversity may extend con‐ biogeography and conservation may benefit from siderably beyond that of conservationists. How‐ adopting a stricter policy of usage. One strategy ever, Fisher and Blomberg (2011) found that hu‐ would be to strictly confine the term ‘rediscovery’ man population overlap did not predict rediscov‐ to species categorized as extinct in the IUCN sys‐ ery rate in mammals, possibly because expedi‐ tem (Mace et al. 2008) or as ‘possibly extinct’, or tions and surveys may intentionally focus on more ‘lost’ by authoritative sources (Table 1, categories remote areas. 1, 2 and 3). It should be noted that many species that are considered possibly extinct are listed as Ecological factors “critically endangered” in the IUCN system. For The final aspect of rediscovery is the ecological example, Fisher (2011a) restricts her analysis to characteristics of the putatively extinct species rediscovered mammal species that had been pre‐ that may make verification of its continued exis‐ viously reported as globally extinct or possibly ex‐ tence problematic. For example, if the species is tinct. It should be noted, however, that this ap‐ very rare and/or dispersed, then it may be difficult proach will not completely eliminate all the cases to locate an individual/population within an area of species that are missing through low levels of of potentially suitable habitat. Even if the survey surveying. team is in the same area as the target species, it An alternative strategy could be to classify may still not be encountered because of pheno‐ rediscovery purely in terms of the length of time typic and ecological traits (e.g. cryptic coloration, without a formal record. If this were adopted, the lack of vocalizations, skulking behaviour, etc.) that only issue would be an appropriate time frame for reduce the probability of detection (Scheffers et a given taxon. For example, De Roland et al. al. 2011). However, the evidence for this effect is (2007) felt justified in claiming the ‘rediscovery’ of variable: Fisher and Blomberg (2011) found that in the Madagascar Pochard (Athya innotata) just 15 mammals many ecological characteristics such as years after the last confirmed sighting – conceiva‐ cryptic coloration and arboreal and nocturnal be‐ bly the same individual. haviour were not significantly associated with re‐ Using a simple time‐based criterion would discovery – although smaller rediscovered mam‐ provide a single, objective definition of rediscov‐ mals had been missing for longer periods of time ery – whatever the cause of the gap in zoological (Fisher 2011b). records. Conservation bodies could potentially use A possible example of ecology driving the this definition to periodically produce lists of spe‐ lack of records is the Night Parrot, a species that is cies that may still be extant and, by extension, are known from 23 specimens and many sightings of in need of rediscovery. These could be categorized varying reliability from a wide geographic area of according to the time since a species was last re‐ inland Australia (McDougall et al 2009). From corded (e.g. <25 years ago, 25–49 years ago, 50– what little information exists, the Night Parrot is 100 years ago, >100 years ago, etc.). One advan‐ crepuscular or nocturnal, cryptic, and when ap‐ tage of such a system would be to maintain and proached will only flush at close quarters, then fly extend the practice of biogeographical expedi‐ low over short distances before plunging back into tions to remote areas. It would also help guard cover (Forshaw and Cooper 2002). Perhaps unsur‐ against the overuse or misrepresentation of redis‐

116 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 Richard J. Ladle et al. coveries in the media (Ladle et al. 2009). It would Boles, W.E., Longmore, N.W., & Thompson, M.C. (1994) offer a viable alternative to the use of terms such A recent specimen of the Night Parrot Geopsit‐ tacus occidentalis. Emu, 94, 37–40. as ‘possibly extinct’ (Butchart et al. 2006) and Buden, D.W. (1996) Rediscovery of the Pohnpei Moun‐ ‘data deficient’, and would ensure better quality tain Starling (Aplonis pelzelni). Auk, 113, 229– of data for future biogeographical studies. 230. Butchart, S.H.M., Stattersfield, A.J. & Brooks, T.M. Conclusions (2006) Going or gone: defining ‘Possibly Extinct’ species to give a truer picture of recent extinc‐ The rediscovery of a species that was thought to tions. Bulletin of the British Ornithology Club, be extinct can generate global interest and repre‐ 126a, 7–24. sents a real opportunity for conservationists to Butchart, S.H.M (2007) Birds to find: a review of ‘lost’, reassert core values and raise funds that may help obscure and poorly known African bird species. Bulletin of the African Bird Club, 14, 138–157. protect poorly known habitats. Moreover, redis‐ Channell, R. & Lomolino, M.V. (2000) Trajectories to coveries provide a unique source of information extinction: spatial dynamics of the contraction about the rarest and least‐known species (for cer‐ of geographical ranges. Journal of Biogeography, tain taxa) that can be used to investigate bio‐ 27, 169–179. geographic theories about range loss and extinc‐ Collar, N.J., Gonzaga, L.P., Krabbe, N., Madrono Nieto, tion. Both of these important agendas would A., Naranjo, L.G., Parker III, T.A. & Wege, D.C. (1992) Threatened birds of the Americas. 3rd benefit from a greater systematization of the con‐ edn. Smithsonian Institution Press, Washington. cept of rediscovery, acknowledging the varying Collar, N.J., Andreev, A.V., Chan, S., Crosby, M.J., Subra‐ causes (both social and ecological) of gaps in the manya, S. & Tobias, J.A. (2001). Threatened temporal records of rare species. Birds of Asia. Birdlife International, Cambridge. In summary, the study of rediscoveries pro‐ Conservation International (2010) The search for lost frogs. Available from http:// vides a wonderful opportunity to assess both the www.conservation.org/campaigns/lost_frogs. subtle ecological and biogeogeographical charac‐ Accessed 24 January 2012. teristics of exceptionally rare species of well stud‐ Crowley, B. (2011) Extinction and rediscovery: where ied taxa such as amphibians, birds and mammals, the wild things are. Journal of Biogeography, 38, and the fascinating historical and cultural trends in 1633–1634. zoological surveying and exploration. Considerable Dalton, R. (2005) Sighting of ‘extinct’ bird may have been a case of mistaken identity. Nature, 436, efforts are being made to untangle these interact‐ 447. ing factors (Fisher 2011a,b; Fisher and Blomberg Dalton, R. (2010) Still looking for that woodpecker. Na‐ 2011, Scheffers et al. 2011), while the recent tar‐ ture 463, 718–719. geting of ‘lost species’ by international conserva‐ De Roland, L.R., Sam, T.S., Rakotondratsima, M.P.H. & tion NGOs is generating considerable amounts of Thorstrom, R. (2007) Rediscovery of the Mada‐ gascar Pochard Aythya innotata in Northern valuable new data. Nevertheless, the lack of redis‐ Madagascar. Bulletin of the African Bird Club 14, covered species that were previously well known 171–174. and which had undergone a well documented Fisher, D.O. (2011a) Trajectories from extinction: process of population decline, fragmentation and where are missing mammals rediscovered? local extinction (Scheffers et al. 2011) remains a Global Ecology & Biogeography, 20, 415–425. worrying trend for global conservation. Fisher, D.O. (2011b) Cost, effort and outcome of mam‐ mal rediscovery: neglect of small species. Bio‐ logical Conservation, 144, 1712–1718. References Fisher, D.O. & Blomberg, S.P. (2011) Correlates of redis‐ Balchon, C. (2007) Back from the dead! A potpourri of covery and the detectability of extinction in recent rediscoveries in the Neotropics. mammals. Proceedings of the Royal Society B: Neotropical Birding, 2, 4–11. Biological Sciences, 278, 1090–1097. BirdLife International (2009) Quest launched to find Fitzpatrick, J.W., Lammertink, M., Luneau Jr., et al. ‘lost’ birds. Available from http:// (2005) Ivory‐billed woodpecker (Campephilus www.birdlife.org/news/news/2009/08/ principalis) persists in continental North Amer‐ lost_and_found. Accessed 25 March 2010. ica. Science, 308, 1460–1462. frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 117 rediscoveries in biogeography

Forshaw, J.M. & Cooper, W.T. (2002) Australian par‐ matic large‐billed reed warbler Acrocephalus rots. 3rd edn. Alexander Editions, Robina. orinus rediscovered after 139 years. Journal of Greenway, J.C. (1967) Extinct and vanishing birds of the Avian Biology, 38, 133–138. world. 2nd Edition. Dover, New York. Rozendaal, F.G. & Lambert, F.R. (1999) The taxonomic Hennache, A., Rasmussen, P., Lucchini, V., Rimondi, S. and conservation status of Pinarolestes & Randi, E. (2003) Hybrid origin of the imperial sanghirensis Oustalet 1881. Forktail, 15, 1–13. pheasant Lophura imperialis (Delacour and Ryan, P.G., Sinclair, I., Cohen, C., Mills, M.S.L., Spottis‐ Jabouille, 1924) demonstrated by morphology, woode, C. & Cassidy, R. (2004) The conservation hybrid experiments, and DNA analyses. Biologi‐ status and vocalisations of threatened birds cal Journal of the Linnean Society, 80, 573–600. from the scarp forest of the Western Angola Jepson, P.J. & Ladle, R.J. (2010) Conservation: a begin‐ Endemic Bird Area. Bird Conservation Interna‐ ner’s guide. One World, Oxford. tional, 14, 247–260. Ladle, R.J. & Jepson, P.R. (2008) Toward a biocultural Scheffers, B.R., Yong, D.L., Harris, J.B.C., Giam, X. & theory of avoided extinction. Conservation Let‐ Sodhi, N.S. (2011) The World’s rediscovered ters, 1, 111–118. species: back from the brink? PLoS ONE 6, Ladle, R.J., Jepson, P., Jennings, S. & Malhado, A.C.M. e22531. (2009) Caution with claims that a species has Sergeant, D.E., Gullick, T., Turner, D.A. & Sinclair, J.C. been rediscovered. Nature, 461, 723. (1992) The rediscovery of the São Tomé Gros‐ Lomolino, M.V., Riddle, B.R. & Brown, J.H. (2006) Bio‐ beak Neospiza concolor in south‐western São geography. 3rd edn. Sinauer, Sunderland, MA. Tomé. Bird Conservation International, 2, 157– Mace, G.M., Collar, N.J., Gaston, K.J., Hilton‐Taylor, C., 159. Akçakaya, H.R., Leader‐Williams, N., Milner‐ Stephenson, B.M., Flood, R., Thomas, B. & Saville, S. Gulland, E.J. & Stuart, S.N. (2008) Quantification (2008) Rediscovery of the New Zealand storm of extinction risk: IUCN's system for classifying petrel (Pealeornis maoriana Mathews 1932): threatened species. Conservation Biology, 22, two sightings that revised our knowledge of 1424–1442. storm petrels. Notornis, 55, 77–83. McDougall, A., Porter, G., Mostert M., Cupitt R., Cupitt Stresemann, E. (1931) Die Zosteropiden der indo‐ S., Joseph, L., Murphy S., Janetzki H., Gallagher australischen Region. Mitteilungen aus dem A. & Burbidge A. (2009) Another piece in an Aus‐ Zoologischen Museum Berlin, 17, 201–238. tralian ornithological puzzle –a second Night Stokstad, E. (2007) Gambling on the ghost bird. Sci‐ Parrot is found dead in Queensland. Emu, 109, ence, 317, 888–892. 198–203. Svensson, L., Prŷs‐Jones, R., Rasmussen, P.C. & Olsson, McCarthy, M.A. (1998) Identifying declining and threat‐ U. (2008) Discovery of ten new specimens of ened species with museum data. Biological Con‐ large‐billed reed warbler Acrocephalus orinus, servation, 83, 9–17. and new insights into its distributional range. Milne, T. & Bull, C.M. (2000) Burrow choice by individu‐ Journal of Avian Biology, 39, 605–610. als of different sizes in the endangered pygmy Timmins, R.J., Mostafawi, N., Rajabi, A.M., Noori, H., blue tongue lizard Tiliqua adelaidensis. Biologi‐ Ostrowski, S., Olsson, U., Svensson, L. & Poole, cal Conservation, 95, 295–301. C.M. (2010) The discovery of Large‐billed Reed Pitra, C., VazPinto, P., O’Keeffe, B.W.J., Willows‐Munro, Warblers Acrocephalus orinus in north‐eastern S., Jansen van Vuuren, B. & Robinson, T.J. Afghanistan. BirdingASIA, 12, 42–45. (2006) DNA‐led rediscovery of the giant sable Tobias, J.A., Butchart, S.H.M. & Collar, N.J. (2006). Lost antelope in Angola. European Journal of Wildlife and found: a gap analysis for the Neotropical Research, 52, 145–152. avifauna. Neotropical Birding, 1, 4–22. Rasmussen, P.C., Wardill, J.C., Lambert, F.R., & Riley, J. Towns, D.R. & Daugherty, C.H. (1994) Patterns of range (2000) On the specific status of the Sangihe contractions and extinctions in the New Zealand White‐eye Zosterops nehrkorni, and the taxon‐ herpetofauna following human colonization. omy of the Black‐crowned White‐eye Z. atrifrons New Zealand Journal of Zoology, 21, 325–339. complex. Forktail, 16, 69–80. Triantis, K.A., Borges, P.A.V., Ladle, R.J., et al. (2010) Riddle, B., Ladle, R.J., Lourie, S. & Whittaker, R.J. (2011) Extinction debt on oceanic islands. Ecography, Basic biogeography: estimating biodiversity and 33, 1–10. mapping nature. In: Ladle, R.J. & Whittaker, R.J. (Editors) Conservation Biogeography. Oxford Edited by Jan Beck University Press, Oxford, pp. 47–92. Round, P.D., Hansson, B., Pearson, D.J., Kennerley, P.R. & Bensch, S. (2007). Lost and found: the enig‐

118 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 membership corner ISSN 1948‐6596 from the society Getting to know IBS Early Career Members

The International Biogeography Society (IBS), species’ traits, island biogeography, phylogeogra‐ founded just 10 years ago, is fast growing both in phy, global change biology, marine biogeography, terms of members and activities offered (Field and or paleobiogeography, among others. Their broad Heaney 2011). Students and early‐career bio‐ interests are also reflected in the fact that most geographers are also becoming increasingly in‐ ECM are also affiliated with societies focusing on volved within the IBS. From 2002 to 2010, the pro‐ diverse topics, including ecology, evolution, con‐ portion of new members who are students joining servation, paleontology, geography, botany, mam‐ the IBS each year has increased from 23% to 48%. malogy, entomology, etc. These are indeed very Currently, student members comprise 35% of encouraging results that show the IBS is reaching IBS’s 740 members. The IBS, aware of the rising young researchers from a wide variety of research importance of these younger members, has been topics and geographic locations. trying to increase the benefits available for them. In general terms, the IBS is meeting ECM In addition to the student travel grants, poster needs (25% responded that the IBS is doing this awards and discussion groups held at the IBS “very well”, 60% “fairly well”). However, there is meetings, the IBS is trying to foster interaction room for improvement (15% responded “not very among students and postdocs, which recently cul‐ well”), and several suggestions were made; re‐ minated in the first IBS Early Career conference sponses to open‐ended questions emphasized the that was held at Oxford University from 23 to 25 need for more off‐year meetings (regional meet‐ September 2011 (http://www.biogeography.org/ ings, workshops, etc.), more jobs/grant announce‐ html/Meetings/index.html). ments, more travel grants, online teaching re‐ With the intention of getting to know its sources, more talks at the IBS meetings by early‐career members (herein ECM) and learning younger researchers and more opportunities to their opinions on the services provided by the IBS meet other researchers. The IBS is already work‐ and on how these can be improved, the IBS in‐ ing towards improving the services it provides to vited ECM to participate in a survey that was held all its members, and new actions are being made in June 2011. Of the 48 ECM that completed this to adopt suggestions. survey, 11% were Junior Postdocs, 75% were PhD The first action was to support the IBS Early students, 8% were Masters students, and 6% were Career conference (for students and biogeogra‐ undergraduate students. Around 17% were aged phers who have finished their PhDs in the past five between 20‐25 years, 49% were 26‐30 years, 23% years). Almost ninety young researchers partici‐ were 31‐35 years, and 11% were more than 35 pated and had the chance to present their work, years young; 56% were female and 44% were and to interact with each other and with the IBS male. Although most ECM are currently affiliated board members. This conference was organized either with North American or European institu‐ into ten different sessions that covered several tions (50% and 33% respectively; total of 42 an‐ aspects of macroecology, island biogeography, swers), they represent a total of 24 nationalities; phylogeography, paleobiogeography, evolutionary 26% are from North America, 17% from Central biogeography and conservation biogeography. and South America, 15% from Northern Europe, Second, we are also working towards in‐ 28% from Southern Europe, and the other 12% creasing regular communication among IBS mem‐ from Australia/New Zealand, the Middle East, Af‐ bers. One way of doing this is through online so‐ rica and Asia. ECM work on a very broad range of cial networks, such as Facebook, and other web‐ topics, from species distribution patterns (the based platforms (e.g. the IBS blog; http:// most mentioned topic), to evolutionary biogeog‐ biogeography.blogspot.com/). Currently, the IBS raphy, dispersal and colonization, biogeography of has a Facebook group with ~590 members, where frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 119 membership corner anyone can post announcements, share ideas and and 43%, respectively; 11% would prefer to have publications of general interest, start discussions mainly senior researchers and 6% showed no pref‐ and interact with other members. Most ECM are erence). There was no overwhelming support for in fact Facebook users (80%; only 7% are Twitter student‐only sessions in future meetings (55% users), but only 8% of these members read the IBS found it important), but most respondents Facebook page on a weekly basis, and 44% actu‐ showed some willingness to extend their stay in ally never read it (31% read it once per month, order to attend this type of event (83%). In the and 17% every 3‐6 months). Regarding the IBS previous meetings, students (particularly those blog, again only a small number of people read it who have been awarded with a student travel on a weekly basis (6%), with most people reading grant) have been invited to attend discussion it once per month (38%; 31% read it every 3‐6 groups, where senior biogeographers lead the months and 25% never read it). Another platform discussion on several subjects, from career and the IBS has for communicating with its members, publishing advice to specific research topics. and to foster communication between its mem‐ Those who have attended these student discus‐ bers, is the online journal Frontiers of Biogeogra‐ sion groups in past meetings (41%) found them phy (http://www.biogeography.org/html/fb.html). helpful (63%). Suggestions for discussion topics in This journal has a section especially devoted for future meetings, other than those already covered this purpose – the membership corner – of which in these discussion groups, included advanced most ECM were not aware (66%). Thirty‐six per‐ analysis in biogeography and partnerships and cent of ECM read every issue, while 31% read 2‐3 international activities among researchers. There issues per year (27% read it rarely and only 6% was some support for future off‐year meetings never read it). Main sections of interest to the (33% found it useful; 61% said it was somewhat ECM are (i) mini‐reviews on a particular taxon, useful, and over 90% said they would at least try biogeographic topic, or question, (ii) thesis ab‐ to attend), especially if these are focused on spe‐ stracts, and (iii) symposium/congress summaries. cific research topics and methodologies (31% and In fact, 88% showed interest in submitting a 29%, respectively; there was a tie between meet‐ manuscript to any of these sections. ings on specific geographic realms and on a broad One of the most important activities organ‐ scope within biogeography – 20% each). Some ized by the IBS is the biennial meeting. The next respondents also called for workshops and semi‐ one will be held at Florida International University nars, online courses, cross‐society ventures to in Miami, Florida, in January 2013 (http://www. boost interaction between similarly oriented aca‐ biogeography.org/html/Meetings/2013). Most demics and excursions into biogeographically in‐ ECM are planning to attend this meeting (79%) teresting regions covering a broad range of taxa. and would prefer to give a talk (51%; 23% prefer a There was also a significant interest in having a poster presentation and 26% have no particular showcase at the next IBS meeting of funding agen‐ preference). One of IBS’ concerns is to maximize cies from different countries (70%), with most re‐ compatibility between high quality talks and fair spondents being willing to provide information on representation of researchers from different this matter (55%). countries, gender, and career stages. There was The long‐term success of any growing soci‐ almost an even split among ECM on favoring a ety depends on the involvement and interest of its similar number of talks by established and youngest members. We’re fortunate that many younger researchers, and having more talks by ECM have shown willingness to get involved in senior researchers plus some younger ones (40% promoting communication between IBS members, Did you know that any member of the IBS may raise an issue or appeal a decision of the gover‐ ning Board 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's Secretary (check current list of officers at http://www.biogeography.org/).

120 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 membership corner to help organizing off‐year activities, and to sub‐ Ana M. C. Santos mit manuscripts to Frontiers of Biogeography. The IBS Student‐at‐Large; Departamento de Ecologia, IBS wants to hear and share more of the early ca‐ Instituto de Ciências Biológicas, Universidade Federal reer members’ opinions and ideas; this article is de Goiás, Brazil. intended as both thanks and encouragement for e‐mail: [email protected] your active involvement, especially in the readily accessible platforms such as Frontiers of Biogeog‐ References raphy and Facebook. Finally, we would like to Field, R. & Heaney, L.R. (2011) Looking to the future of thank all the members who participated in this the IBS: the 2011 IBS membership survey. Fron‐ survey, and particularly those who have shown tiers of Biogeography, 3, 71‐73. interest in devoting some of their time to the soci‐ ety. We look forward to working with and for you Edited by Matthew Heard in the coming years. from the society Call for proposals for hosting 7th Biennial Conference of the IBS

We are seeking proposals for hosting the 7th bien‐ 3. Actual site of the meeting and the capacity of nial conference of the International Biogeography the auditorium. Society to be held in early January 2015. Proposals 4. Space for poster sessions‐‐general size and lo‐ should be submitted by individuals who are inte‐ cation relative to the auditorium. rested in chairing the local (host) committee. The 5. Approximate cost for three‐day use of the ve‐ duties of the local host include conducting contra‐ nue. A specific quote is not needed, but evi‐ ct negotiations with the venue and the hotel as dence of the price competitiveness is crucial. well as all local logistics including field trip organi‐ 6. Transportation infrastructure, including travel zation and production of the abstract bo‐ from airport. ok. Minimum requirements of the venue are 1) 7. Attractions in the vicinity of the conference one auditorium with a capacity of 450‐550 people site, including field trip potential. (2 days), 2) three or four smaller rooms with a ca‐ 8. Who would potentially serve on the local orga‐ pacity of 75‐150 people (1 day), and 3) various nizing committee? smaller meeting rooms. The IBS is interested in Proposals from prospective hosts of the holding the biennial conference in locations fairly biennial conference must be received before 20 convenient with respect to the majority of its January 2012. Please send proposals by email to membership base in North America and Euro‐ Daniel Gavin, IBS Vice‐President for Conferences pe. Locations of past (and upcoming) conferences at [email protected]. can be seen here: http://www.biogeography.org/ html/Meetings/index.html. Please include the following information in Dan Gavin the proposal: IBS Vice‐President for Conferences; 1. Location of the meeting (city) and the host ins‐ Department of Geography, University of Oregon, USA. titution or organization. e‐mail: [email protected]

2. What would be the benefit of hosting the con‐ Erratum ‐ 11 Feb 2011: The original article published on 09 Feb 2011 ference at this location? incorrectly identified Dan Gavin as the "IBS Student‐at‐Large".

If you want to announce a meeting, event or job offer that could be of interest for (some) bio‐ geographers, or you want to make a call for manuscripts or talks, please contact us at [email protected] and [email protected]. frontiers of biogeography 3.3, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society 121 membership corner Job announcements

Three Professorships and One Tenure‐Track arch areas of interest include statistical methods Lectureship for experimental design, epidemiology, medical informatics, evolutionary biology, sequence bioin‐ University of California, Merced, USA formatics, genomics, evolution of microbial sys‐ The School of Natural Sciences at the University of tems and pathogens, and systems biology. The California, Merced seeks applicants for four facul‐ Lecturer position closely parallels a tenure‐track ty positions: Ecology (Full or Associate with tenu‐ Assistant Professor but with an emphasis on un‐ re, or Assistant tenure‐track), Systems Biology dergraduate education. All applicants must be (Assistant tenure‐track), and Biostatistics able to teach effectively at both undergraduate (Assistant tenure‐track), and one tenure‐track Bio‐ and graduate levels. For more information and to logy Lecturer. For the Ecology position, we seek apply go to: http://jobs.ucmerced.edu/n/ outstanding individuals with research interests in academic/listings.jsf;jsessionid=95FADBAFFF4C13 any ecological field using experimental, field, com‐ F912A3B023DA4F1F80?seriesId=1 putational, and/or theoretical approaches and Interested applicants should submit mate‐ working at population to global scales. The Sys‐ rials online. Applications will be considered star‐ tems Biology position includes research areas that ting 05 December 2011 (Biostatistics, Systems Bio‐ use comprehensive datasets and multiple types of logy professorships), or 16 December 2011 analysis to relate overall biological function to un‐ (Ecology professorship and Biology Lecturer). UC derlying biochemical or biophysical processes for Merced is an AA/EOP employer. predictive understanding. The Biostatistics rese‐ upcoming events

VIPCA Molecular Ecology VertNet biodiversity informatics training 4–7 February 2012 – Vienna, Austria workshop http://www.vipca.at/MOLECOL/ 24–30 June 2012 – Boulder, USA http://vertnet.org/about/BITW.php Annual Conference of the Society for Tropical Ecology (gtö) 97th ESA Annual Meeting Islands in land‐ and seascape: The challenges of frag‐ Life on Earth: Preserving, Utilizing, and Sustaining our mentation Ecosystems 22–25 February 2012 – Erlangen, Germany 5–10 August 2012 – Portland, USA http://www.gtoe‐conference.de/ http://esa.org/meetings/

6th Annual Meeting of the Specialist Group on Macroecology of the Ecological Society of 3rd European Congress of Conservation Biol‐ Germany, Austria and Switzerland (GfÖ) ogy 29 February – 2 March 2012 – Frankfurt, Germany Conservation on the edge http://www.bik‐f.de/ 28 August – 1 September 2012 – Glasgow, UK http://www.eccb2012.org/ st 21 Workshop of the European Vegetation Survey (EVS) 6th International Conference of the IBS 24–27 May 2012 – Vienna, Austria January 2013 – Florida, USA http://evs2012.vinca.at/ http://www.biogeography.org/

122 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011 table of contents frontiers of biogeography the scientific magazine of the International Biogeography Society volume 3, issue 3 ‐ November 2011 ISSN 1948‐6596 news and update update: Species–area curves and the estimation of extinction rates, by J. Beck 81 update: Extinct or extant? Woodpeckers and rhinoceros, by R. Ladle 83 update: Climate wars, by J. Beck 84 update: Emerging research opportunities in global urban ecology, by F.A. La Sorte 85 update: Beyond taxonomical space: large‐scale ecology meets functional and phylogenetic diversity, by M.V. 87 Cianciaruso book review: A mangrove compendium, by U. Berger 91 book review: A comprehensive foundation for the application of biogeography to conservation, by T. Newbold 93 book review: A new encyclopedia for biological invasions, by R.A. Francis 95 book review: A piscine history of the Neotropics, by A.E. Magurran 97 books noted with interest 99 thesis abstract: Applying species distribution modeling for the conservation of Iberian protected invertebra‐ 101 tes, by R.M. Chefaoui opinion and perspectives opinion: Political erosion dismantles the conservation network existing in the Canary Islands, by J.M. Fernán‐ 106 dez‐Palacios & L. de Nascimento perspective: The causes and biogeographical significance of species’ rediscovery, by R.J. Ladle et al. 111 membership corner from the society: Getting to know IBS Early Career Members, by A.M.C. Santos 119 from the society: Call for proposals for hosting 7th Biennial Conference of the IBS, by D. Gavin 121 Job announcements 122 Upcoming meetings 122 frontiers of biogeography copyright notice Copyright © 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. It is also strictly forbidden to make copies of whole issues of this journal for any commercial purpose without the express written permission of the IBS. 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 journal during one year after its publication. Any active distribution of any part or the whole issue of the journal is explicitly permitted since the date of publication, and any passive distribution is explicitly permitted after one year of the date of publication. Any individual and/or institution can download, read and/or print a copy of any article or the whole journal for non‐commercial educational or non‐commercial research purposes at any time. This includes an express permission to 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 copyright notice without modification. All articles are copyrighted by their authors under a universal Creative Com‐ mons Attribute License (CCAL). This license permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. All authors endorse, permit and license the IBS to grant any third party the copying and use privileges specified above without additional consideration or payment to them or to the IBS. These endorsements, in writing, 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 to distribute 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 materials prepared for the International Biogeography Society Inc activities will be held solely by the International Biogeography Society Inc and in the name of the International Biogeography Society Inc.". And "Article 8. Publications. The publications of the Society shall include journals, newsletters, and such 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.