frontiers of biogeography - The International Biogeography Society

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/


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 

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editorial board 

editor‐in‐chief: 

Joaquín Hortal – Museo Nacional de Ciencias Naturales (CSIC), 

Spain and Universidade Federal de Goiás, Brazil 

associate editors: 

Antje Ahrends – Royal Botanic Garden Edinburgh, UK 

Jan Beck – University of Basel, Switzerland 

Jessica Blois – University of Wisconsin, Madison, USA 

Chris Burridge – University of Tasmania, Australia 

Marcus V. Cianciaruso – Universidade Federal de Goiás, Brazil 

Markus Eichhorn – University of Nottingham, UK 

Roy Erkens – Universiteit Utrecht, The Netherlands 

Camilla Fløjgaard – Aarhus University, Denmark 

Dan Gavin – University of Oregon, USA 

Matthew J. Heard – Brown University, USA 

David G. Jenkins – University of Central Florida, Orlando, USA 

Frank A. La Sorte – Cornell lab of Ornithology, USA 

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 

deputy editors‐in‐chief: 

Michael N Dawson – University of California, Merced, USA 

Richard Field – University of Nottingham, UK 

editorial assistant: 

Lauren Schiebelhut – University of California, Merced, USA 

advisory board: 

Miguel B. Araújo – Museo Nacional de Ciencias Naturales (CSIC), 

Spain and Universidade de Évora, Portugal 

Lawrence R. Heaney – Field Museum of Natural History, Chicago, 

USA 

David G. Jenkins – University of Central Florida, Orlando, USA 

Richard Ladle – Universidade Federal de Alagoas, Brazil and Oxford 

University, UK 

Mark V. Lomolino – State University of New York, USA 

IBS V. P. for Public Affairs & Communications 

President: Lawrence R. Heaney 

President Elect: Rosemary Gillespie 

VP for Conferences: Daniel Gavin 

VP for Public Affairs & Communications: Michael N Dawson 

VP for Development & Awards: George Stevens 

Secretary: Richard Field 

Treasurer: Lois F. Alexander 

Director‐at‐large: Catherine Graham 

Director‐at‐large: Kathy Willis 

Student‐at‐large: Ana M. C. Santos 

First Past President: James H. Brown 

Second Past President: Mark V. Lomolino 

Third Past President: Brett R. Riddle 

Fourth Past President: Vicki Funk 

Fifth Past President: Robert J. Whittaker 

Upcoming meeting host (ex officio): Kenneth Feeley 

Past Graduate student representative (ex officio): Matthew Heard 

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 

longest‐known, most intuitive and empirically best 

‐proven patterns of biodiversity (Arrhenius 1921). 

Various authors determined theoretically that the 

SAR can be approximated as a power‐law function 

(i.e., S = cA z where S is species richness, A is area 

and c and z are constants; Preston 1962, May 

1975, Harte et al. 1999), with z ≈ 0.25 in continental 

areas but higher when dispersal barriers are 

involved (e.g., ‘island species–area relationship’). 

Empirical data suggested lower z in continental 

areas (0.13‐0.18) and values up to 0.35 for island 

systems (Rosenzweig 1995). Dengler (2009) recently 

came to the conclusion that the power law 

fits empirical data best in most cases (see also 

Dengler & Odeland 2010). Various authors observed 

further systematic variations of z, such as 

when considering spatial scale or sampling design 

(Plotkin et al. 2001, Scheiner 2006, Tjørve 2006, 

Dengler 2009). Kinzig & Harte (2000) pointed out 

the difference between SAR and the endemics– 

area curve (EAR), which considers only species 

endemic to a part of the region under analysis. So 

what could He & Hubbell (2011) report that was 

so novel and generally relevant about SARs to 

merit recent publication in Nature 

Since area seems always to affect biodiversity, 

no matter what taxon, system or scale, SARs 

have frequently been used to estimate species 

richness loss resulting from anthropogenic habitat 

destruction, i.e. extinction rates in a conservation 

context. The loss of a certain amount of area leads 

to fewer species existing in a region – at least 

some regional extinctions occur – and the shape 

of the SAR has typically been used to retrieve 

quantitative estimates of how many species will 

go (regionally) extinct. 

Providing empirical evidence for the extinction 

of a species is challenging and estimating extinction 

rates across a community even more so 

(Ladle et al. 2011, this issue). Yet this is needed for 

many conservation applications, such as schemes 

for offsetting biodiversity loss (Curran et al. 2011) 

or, not least, for political argument. It is therefore 

not surprising that SAR‐based estimates of extinction 

have been welcome despite critical studies 

that often found lower extinction rates than predicted 

(e.g., Kinzig & Harte 2000). It was argued, 

reasonably, that on top of imminent extinction in 

some species, others will be doomed to future 

extinction because of reductions in their population 

size, and that this ‘extinction debt’ explains 

apparent misfits. Other sources of uncertainty of 

the SAR‐based estimates are the (often false) assumption 

of a completely inhospitable matrix between 

remaining habitat patches (Koh & Ghazoul 

2010) or the use of default slope values (z) in the 

absence of system‐specific fitted data. 

He & Hubbell (2011) pointed out that a 

backward interpolation of SARs is a flawed concept 

of measuring extinction rates (see also Kinzig 

& Harte 2000). This is because the area gain 

needed to encounter the first individual of a new 

species (which shapes the SAR) is always smaller 

than the area loss needed to remove the last individual. 

To show this, they formulated both as spatially 

explicit sampling processes (SAR for first encounters, 

EAR for last encounters). They concluded 

that SAR‐derived estimates of imminent 

extinction will always be too high, unless individuals 

are randomly distributed (i.e., no aggregated 

occurrence of individuals within a species), which 

is an unrealistic assumption. He & Hubbell (2011) 

also showed that the EAR is a good predictor of 

empirical extinction rates even if no spatial aggregation 

is modelled, which offers an alternative 

(but a more challenging one) for estimating immediate 

extinction of endemics from area loss. 

He & Hubbell (2011) clearly acknowledged 

that there is an anthropogenic extinction crisis 

and that habitat loss causes extinction. Furthermore, 

they did not claim that small population 

sizes of remaining species could not lead to further, 

lagged extinction (in He & Hubbell’s view, 

EARs model only imminent extinction – and so do 

SARs, but wrongly). Despite this, He & Hubbell 

(2011) already anticipated that pointing out this 

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, 

and the correspondence on the paper (Evans et al. 

2011, Brooks 2011; see also online comments at 

http://www.nature.com/nature/journal/v474/ 

n7351/full/474284b.html) seems to confirm that. 

The paper is viewed as irresponsibly undermining 

conservation efforts by allowing anti‐conservation 

groups to claim that things are not as bad as previously 

asserted (fossil fuel lobbying in the climate 

change discussion is cited as example of this tactic). 

Conserving nature is not only about science, 

but it is to a large degree politics – and correcting 

an error leads to better science but might weaken 

political success. I think scientists must correct 

themselves and not hold on to preconceived 

ideas, even if it creates such dilemmas. 

However, He & Hubbell (2011) studied area 

effects as a sampling problem in continental regions, 

which is probably appropriate for capturing 

immediate extinction in many conservation settings 

which occur at the regional or landscape 

scale. It remains to be understood and tested 

whether their conclusions – that (a) EAR estimates 

extinction better than SAR (cf. Kinzig & Harte 

2000, Pereira et al. 2012) and (b) z differs systematically 

between SAR and EAR (which is presented 

confusingly) – are generalities. Thus it remains to 

be seen whether SARs always overestimate extinction, 

as He and Hubbell (2011) claimed. A further 

task will be to quantitatively estimate how 

many more species may go extinct after a time 

lag: how large the extinction debt really is (see 

also Pereira et al., in press). In this context, it may 

be worthwhile to thoroughly investigate under 

which circumstances, if any, the consequences of 

area lost to habitat destruction could be understood 

solely on the basis of island biogeographic 

mechanisms (Rosenzweig 2001) – that is, species 

richness as equilibrium between immigration + 

speciation and extinction. The spatial and temporal 

scales of analysis, among other factors, may be 

relevant for this. Under such circumstances, SARs 

may estimate the new equilibrium state, accounting 

for imminent and time‐lagged extinctions. 

Jan Beck 

University of Basel, Dept. Environmental Science 

(Biogeography section), Basel, Switzerland. 

e‐mail: jan.beck@unibas.ch; 

http://www.biogeography.unibas.ch/beck 

References 

Arrhenius, O. (1921) Species and area. Journal of Ecology, 

9, 95–99. 

Brooks, T.M. (2011) Extinctions: consider all species. 

Nature, 474, 284. 

Curran, M., De Baan, L., de Schryver, A.M., van Zelm, 

R., Hellweg, S., Koellner, T., Sonnemann, G. & 

Huijbregts, M.A.J. (2011) Toward meaningful 

end points of biodiversity in life cycle assessment. 

Environmental Science and Technology, 

45, 70–79. 

Dengler, J. (2009) Which function describes the species 

–area relationship best A review and empirical 

evaluation. Journal of Biogeography, 36, 728– 

744. 

Dengler, J. & Oldeland, J. (2010) Effects of sampling 

protocol on the shapes of species richness 

curves. Journal of Biogeography, 37, 1698–1705. 

Evans, M., Possingham, H. & Wilson, K. (2011) Extinctions: 

conserve not collate. Nature, 474, 284. 

Harte, J., Kinzig, A. & Green, J. (1999) Self‐similarity in 

the distribution and abundance of species. Science, 

284, 334–336. 

He, F. & Hubbell, S.P. (2011) Species–area relationships 

always overestimate extinction rates from habitat 

loss. Nature, 473, 368–371. 

Kinzig, A. & Harte, J. (2000) Implications of endemics– 

area relationships for estimates of species extinctions. 

Ecology, 81, 3305–3311. 

Koh, L.P. & Ghazoul, J. (2010) A matrix‐calibrated species–area 

model for predicting biodiversity 

losses due to land‐use change. Conservation 

Biology, 24, 994–1001. 

Ladle, T.J., Jepson, P., Malhado, A.C.M., Jennings, S. & 

Barua, M. (2011) The causes and biogeographical 

significance of species’ rediscovery. Frontiers 

of Biogeography, 3, 111–118. 

May, R.M. (1975) Patterns of species abundance and 

distribution. In Cody M.C. & Diamond J.M. 

(eds.), Ecology and evolution of communities, 

pp. 81–120; Belknap Press, Cambridge (Mass.). 

Pereira, H.M., Borda‐de‐Agua, L. & Martins, I.S. (2012) 

Geometry and scale in species–area relationships. 

Nature, in press. 

Plotkin, J.B., Potts, M.D., Yu, D.W., et al. (2000) Predicting 

species diversity in tropical forests. Proceedings 

of the National Academy of Sciences USA, 

97, 10850–10854. 

82 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

ISSN 1948‐6596 

Preston, F.W. (1962) The canonical distribution of commonness 

and rarity: Part I. Ecology, 43, 185– 

215. 

Rosenzweig, M.L. (1995) Species diversity in space and 

time. Cambridge University Press, Cambridge. 

Rosenzweig, M.L. (2001) Loss of speciation rate will 

impoverish future diversity. Proceedings of the 

National Academy of Sciences USA, 89, 5404– 

5410. 

Scheiner, S.M. (2003) Six types of species–area curves. 

Global Ecology and Biogeography, 12, 441–447. 

Tjørve, E. (2006) Shapes and functions of species–area 

curves: a review of possible models. Journal of 

Biogeography, 30, 827–835. 

Edited by Joaquín Hortal 


update 

Extinct or extant Woodpeckers and rhinoceros 

Biogeographical research needs accurate data on 

the distribution of species. For many species this is 

exceedingly difficult to obtain, leading to a lack of 

global information collectively known as the Wallacean 

shortfall. Fortunately, new tools are being 

developed that allow conservationists and biogeographers 

to determine the existence of extant 

populations with much greater accuracy. 

Foremost among these new tools is the increasing 

use of genetic analysis. This was recently 

used to great effect to confirm the extinction of 

the Javan rhinoceros (Rhinoceros sondaicus annamiticus) 

in Cat Tien National Park in Vietnam 

(Brook et al. 2011). Despite their enormous size, 

Javan rhinoceros are remarkably shy forestdwelling 

animals that are difficult to see under 

natural conditions and were only rediscovered in 

mainland Asia in 1988. Given the difficulty of traditional 

surveying techniques, scientists from 

WWF and the Cat Tien National park had been 

monitoring the population by conducting genetic 

analysis of dung samples collected in the park between 

2009 and 2010. The analysis indicated that 

all the dung belonged to a single individual, the 

body of which was found April 2010, thereby confirming 

the extinction of the population. 

Of course, genetic analysis is costly, time 

consuming and requires some form of biological 

tissue (hair, dung, etc.). For many rare animals the 

only information that exists is the occasional sighting, 

the reliability of which is often highly questionable. 

Andrew Solow and his colleagues have 

recently come up with an ingenious method to 

account for this inevitable uncertainty (Solow et 

al. 2011). They use Bayesian (probability‐based) 

statistics to model changes in the rate of valid 

sightings and to assess the quality of uncertain 

sightings for the ivory‐billed woodpecker 

(Campephilus principalis) in North America. The 

woodpecker was controversially rediscovered in 

2005, but a lack of clear documentary evidence 

and the failure of subsequent intensive surveys 

have led many scientists to doubt the veracity of 

this claim. The Bayesian model applied by Solow 

to 68 historical sightings (29 of which were classified 

as uncertain) strongly suggests that the bird is 

indeed extinct, and the 2005 sighting was sadly a 

case of mistaken identity. 

Richard Ladle 

Federal University of Alagoas, Institute of Biological 

Sciences and Health, Brazil and Oxford University, 

School of Geography and the Environment, UK. 

e‐mail: richard.ladle@ouce.ox.ac.uk; 

http://www.geog.ox.ac.uk/staff/rladle.html 

References 

Brook, S., de Groot, P.V.C., Mahood, S. & Long, B. 

(2011) Extinction of the Javan Rhinoceros 

(Rhinoceros sondaicus) from Vietnam. WWF 

Report. Available at: http:// 

www.worldwildlife.org/who/media/press/2011/ 

WWFBinaryitem24584.pdf 

Solow, A., Smith, W., Burgman, M., Rout, T., Wintle, B. 

and Roberts, D. (2011), Uncertain sightings and 

the extinction of the ivory‐billed woodpecker. 

Conservation Biology. doi: 10.1111/j.1523‐ 

1739.2011.01743.x 



83


ISSN 1948‐6596 

update 

Climate wars 

Links between climate and societal instability, 

conflict and war have increasingly been suggested 

and analyzed (Diamond 2005), thereby fusing traditionally 

distinct academic disciplines such as 

(bio‐)geography, (agro‐)ecology and economics, 

history and peace research. Studies exploring 

these relationships are particularly pertinent in 

times of anthropogenic climate change. 

Recent research has provided quantitative 

support for such climate–culture linkages, but 

most of these studies have either been based on 

correlative evidence (e.g., Zhang et al. 2007), analyzed 

short‐term climate fluctuations (e.g., Burke 

et al. 2009) or addressed specific hypotheses on 

the causes of human conflict (Beck and Sieber 

2010). However, in order to make conflict predictions 

under climate‐change scenarios reliable and 

to engage in conflict prevention or mitigation, it is 

important to be certain about causal relationships 

and to fully understand the mechanistic links between 

past climatic changes and historical conflicts. 

Two new studies have attempted this. 

Hsiang et al. (2011) made use of the recurring 

yet irregular El Niño Southern Oscillation 

(ENSO) climatic changes as a natural experiment. 

This allowed them to show, on a global scale and 

for a time period of more than half a century, that 

(within the same localities and societies) civil conflicts 

were more likely to arise during El Niño 

events as compared to La Niña periods. Furthermore, 

no such effect was observed for countries 

outside the ENSO‐affected zone of the world. This 

provides strong evidence that climate is indeed 

causal to these events. However, the authors can 

only speculate on a variety of mechanisms for 

how (warmer and drier) El Niño periods could lead 

to conflict. Effects mediated by decreased agricultural 

productivity and/or economic disturbance 

(e.g., resulting from increases in natural disasters 

and diseases) seem plausible, but psychological 

effects of unusual weather conditions on a large 

number of individuals may also increase a society’s 

conflict potential. 

Zhang et al. (2011) presented a detailed 

causality analysis based on a time series of climatic 

fluctuations over a 300 year period in preindustrial 

Europe. They provide strong support for 

the idea that climatic variation caused fluctuations 

in agricultural productivity, and hence food availability 

and prices. The latter was identified as the 

root cause for a number of societal phenomena 

such as migrations, epidemics, population growth 

and war. A temperature‐based model based on 

these mechanisms could successfully predict periods 

of crisis and harmony for past eras with lessdetailed 

historical records. 

An important future direction of research in 

this field will certainly be the identification of 

natural factors and societal traits that explain 

variation around such climate‐determined patterns. 

Demography and economic performance 

have sometimes been analyzed in this context 

(Samson et al. 2011, Hsiang et al. 2011). However, 

it will require the further integration of the abovementioned 

disciplines to sort out the ultimate 

causes of why certain regions and/or societies 

navigated smoother and less violent routes 

through times of crisis than others (my current 

location, Switzerland, is a prime example within 

the last few centuries). 

Jan Beck 

University of Basel, Dept. Environmental Science 

(Biogeography section), Basel, Switzerland. 

e‐mail: jan.beck@unibas.ch; 

http://www.biogeography.unibas.ch/beck 

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ISSN 1948‐6596 

References 

Beck J., & Sieber, A. (2010) Is the spatial distribution of 

mankind’s most basic economic traits determined 

by climate and soil alone PLoS ONE 5(5): 

e10416. 

Burke, M., Miguel, E., Satyanath, S., Dykema, J. & Lobell, 

D. (2009) Warming increases risk of civil 

war in Africa. Proceedings of the National Academy 

of Sciences USA, 106, 20670–20674. 

Diamond, J. (2005) Collapse: how societies choose to 

fail or succeed. Viking. 

Hsiang, S.M., Meng, K.C. & Cane, M.A. (2011) Civil conflicts 

are associated with the global climate. Nature, 

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 Biogeography, 

20, 532–544. 


Zhang, D.D., Lee, H.F., Wang, C., Lie, B., Pei, Q., Zhang, 

J. & An, Y. (2011) The causality analysis of climate 

change and large‐scale human crisis. Proceedings 

of the National Academy of Sciences 

USA, 108, 17296–17301. 

Zhang, D.D., Brecke, P., Lee, H.F., He, Y.‐Q. & Zhang, J. 

(2007) Global climate change, war and population 

decline in recent human history. Proceedings 

of the National Academy of Sciences USA, 

104, 19214–19219. 

Edited by Richard Ladle 

update 

Emerging research opportunities in global urban ecology 

Biogeographers have examined how human activities 

have affected patterns of biological diversity 

from a variety of perspectives, with special attention 

often given to oceanic islands. With the current 

accelerating pace of environmental change, 

these effects are increasingly evident at global 

scales. Human industry, commerce, agriculture 

and transportation all have the potential now to 

affect natural systems globally through an assortment 

of drivers; primary among these are landuse 

change, species introductions and climate 

change. 

Human activities and their consequences 

come to a unique focus in urban areas, an expanding 

form of land use that is attracting increasing 

research attention from ecologists (Grimm et al. 

2008). Urban areas contain similar environmental 

conditions worldwide and act as a focal point for 

species introductions and extinctions. These human‐dominated 

environments offer unique opportunities 

to investigate the broad‐scale dynamics 

of human‐mediated biotic interchange (La 

Sorte et al. 2007), its consequences for β diversity 

(La Sorte et al. 2008) and the regional factors and 

biological traits associated with native species extinctions 

(Hahs et al. 2009, Duncan et al. 2011). 

Urban areas typically contain spatially heterogeneous 

collections of native and non‐native species 

(McKinney 2008); these unique assemblages can 

be examined based on their compositional 

(Niemelä et al. 2002) and phylogenetic structures 

(Ricotta et al. 2009). Three nested sampling approaches 

are currently used to investigate urban 

systems at broad spatial scales: urban plots or 

transects, the entire urban matrix and the urban 

matrix embedded within a regional context 

(Werner 2011). Each sampling approach provides 

a unique inferential basis, although the third allows 

for more refined interpretation, controlling 

for regional differences. 

A recent study in Global Ecology and Biogeography 

adopts a novel perspective and examines 

how avian assemblages sampled within plots 

of intact vegetation in urban and semi‐natural areas 

differ based on several common macroecological 

relationships. Pautasso et al. (2011) 

compiled data on species composition and abundance 

from all around the globe, although the 

majority of the samples are from Europe and 

North America. A primary finding of the study was 

a lack of evidence for differences in the species– 

area, species–abundance or species–biomass rela‐ 


85


tionships between urban and semi‐natural localities. 

The number of exotic bird species in urban 

areas is low, suggesting that these relationships 

are defined primarily by native species in both 

environments. These findings highlight the importance 

of maintaining intact vegetation within urban 

landscapes and the role of urban diversity as a 

tool for promoting conservation initiatives and 

biological awareness, as emphasized in many urban‐ecology 

studies. Nevertheless, the findings 

from Pautasso et al. (2011) contrast with current 

expectations on how urbanization affects patterns 

of diversity, and should be a motivating factor in 

promoting further research. The increasing prevalence 

and quality of global data sources provides 

an exciting basis to examine the structure and determinants 

of these macroecological relationships 

across more refined temporal, spatial and anthropogenic 

gradients. 

By taking a global perspective, novel insights 

can be gained on the unique position urban 

areas have, both as a source for global change and 

as regions capable of maintaining important aspects 

of biological diversity. Global comparative 

studies also have the potential to bolster and refine 

current recommendations about how to 

maintain biological diversity within humandominated 

landscapes. Specifically, the preservation 

or restoration of patches of intact vegetation 

within urban areas is as valuable in maintaining 

basic macroecological patterns of avian diversity 

as conducting these activities outside urban areas. 

Importantly, this work takes the focus away from 

Europe and North America, where the vast majority 

of the research has been conducted, allowing 

for a more inclusive set of inferences and recommendations. 

Urban data are becoming increasingly 

available through remote sensing activities, 

citizen science initiatives and broader collaborative 

efforts. Exploring how anthropogenic activities 

are impacting natural systems globally is critical 

in supporting a truly comprehensive understanding 

of the current dynamics and long‐term 

consequences of global environmental change. 

Frank A. La Sorte 

Cornell Lab of Ornithology, Ithaca, NY, USA. 

e‐mail: fal42@cornell.edu; 

http://www.birds.cornell.edu/ 

References 

Duncan, R.P., Clemants, S.E., Corlett, R.T., Hahs, A.K., 

McCarthy, M.A., McDonnell, M.J., Schwartz, 

M.W., Thompson, K., Vesk, P.A. & Williams, 

N.S.G. (2011) Plant traits and extinction in urban 

areas: a meta‐analysis of 11 cities. Global Ecology 

and Biogeography, 20, 509–519. 

Grimm, N.B., Faeth, S.H., Golubiewski, N.E., Redman, 

C.L., Wu, J., Bai, X. & Briggs, J.M. (2008) Global 

change and the ecology of cities. Science, 319, 

756–760. 

Hahs, A.K., McDonnell, M.J., McCarthy, M.A.,et al. 

(2009) A global synthesis of plant extinction 

rates in urban areas. Ecology Letters, 12, 1165– 

1173. 

La Sorte, F.A., McKinney, M.L. & Pyšek, P. (2007) Compositional 

similarity among urban floras within 

and across continents: biogeographical consequences 

of human‐mediated biotic interchange. 

Global Change Biology, 13, 913–921. 

La Sorte, F.A., McKinney, M.L., Pyšek, P., Klotz, S., Rapson, 

G.L., Celesti‐Grapow, L. & Thompson, K. 

(2008) Distance decay in similarity among European 

urban floras: the impacts of anthropogenic 

activities on β diversity. Global Ecology and Biogeography, 

17, 363–371. 

McKinney, M.L. (2008) Effects of urbanization on species 

richness: a review of plants and animals. 

Urban Ecosystems, 11, 161–176. 

Niemelä, J., Kotze, D.J., Venn, S., Penev, L., Stoyanov, I., 

Spence, J., Hartley, D. & Montes de Oca, E. 

(2002) Carabid beetle assemblages (Coleoptera, 

Carabidae) across urban‐rural gradients: an international 

comparison. Landscape Ecology, 17, 

387–401. 

Pautasso, M., Böhning‐Gaese, K., Clergeau, P., et al. 

(2011) Global macroecology of bird assemblages 

in urbanized and semi‐natural ecosystems. 

Global Ecology and Biogeography, 20, 426–436. 

Ricotta, C., La Sorte, F.A., Pyšek, P., Rapson, G.L., Celesti 

‐Grapow, L. & Thompson, K. (2009) Phyloecology 

of urban alien floras. Journal of Ecology, 97, 

1243–1251. 

Werner, P. (2011) The ecology of urban areas and their 

functions for species diversity. Landscape and 

Ecological Engineering, 7, 231–240. 



ISSN 1948‐6596 


update 

Beyond taxonomical space: large‐scale ecology meets functional 

and phylogenetic diversity 

Community ecology traditionally focuses on hypothetical‐deductive 

and experimental approaches 

and often is criticized for narrowing our understanding 

of nature to local idiosyncrasies, ignoring 

the importance of historical explanations. On the 

other hand, approaches taken by macroecologists 

and biogeographers have been excessively exploratory 

and correlative, with limited success in 

elucidating the mechanisms responsible for many 

of the large‐scale patterns we observe in nature 

(see Gaston & Blackburn 1999, Ricklefs 2008 and 

references therein). Recognizing that both approaches 

can learn from each other is pivotal in 

the challenge of integrating data from different 

scales in order to unravel the ecological and evolutionary 

mechanisms that influence current patterns 

in biodiversity and ecosystem functioning. 

Species richness has been the most common 

metric used to represent all aspects of biological 

diversity (from genetic and taxonomic to 

phenetic diversity). However, species richness 

alone cannot describe the processes involved in 

species coexistence and ecosystem functioning 

and also does not describe properly the differences 

in community structure. In contrast, phylogenetic 

and functional diversities allow us to 

understand the relative importance of species 

composition in terms of evolutionary history and 

ecological similarities. Phylogenetic diversity (PD) 

is a biodiversity measure that accounts for the 

phylogenetic relationship (hence evolutionary history) 

among species, whereas functional diversity 

(FD) represents how species are distributed in a 

multidimensional niche space defined by ecological 

traits. 

Phylogenetic and functional approaches to 

community ecology emerged as prominent fields 

of research in the last decade (Fig. 1), but somehow 

independently and without much crossover 

in the first years. Early PD measures were proposed 

as a tool to select conservation areas, but 

later the idea was extended to understand how 

communities are assembled from a regional pool. 

FD, which initially was considered the holy grail of 

the biodiversity‐ecosystem functioning agenda, 

also was rapidly applied as a metric for investigating 

assembly rules (see Pavoine & Bonsall 2011). 

How could macroecology and biogeography benefit 

from these two approaches The answer lies in 

understanding what FD and PD should represent 

and how they relate to each other: while phylogenetic 

community ecology links evolutionary and 

biogeographic history to present‐day ecology, 

functional diversity (as any trait‐based approach) 

links niche theory to large‐scale approaches, such 

as macroecology, biogeography or phylogeography. 

Therefore, combining ecological and phylogenetic 

frameworks to explain large scale patterns of 

biodiversity is an important step, taken recently. 

Large‐scale studies involving PD and FD seems to 

be increasing at similar rates (Fig.1). Recently, it 

was shown that both measures can be decomposed 

into gamma (regional), alpha (local) and 

beta (turnover) components. Whereas large‐scale 

studies and any‐scale studies follows a similar 

trend for beta‐PD, there were few studies with 

beta‐FD (none at large‐scale). This is perhaps because 

biogeographers and macroecologists were 

more aware of evolutionary and historical hypotheses, 

so the conceptual framework of beta‐ 

PD was likely to be absorbed first. Also, this could 

reflect the assumption that closely related species 

should be ecologically more similar than distant 

related species and, thus, PD should be a good 

surrogate for FD (in fact this is what most large 

and local‐scale PD studies used to assume). This 

traditional assumption is now debated (e.g. Losos 

2008), and these two measures may be viewed as 

complementary, rather than competing, approaches 

(Gómez et al. 2010, Diniz‐Filho et al. 

2011, Meynard et al. 2011, Pavoine & Bonsall 

2011, Safi et al. 2011). 

While some large‐scale studies involving PD 

and FD are exploratory (e.g. Meynard et al. 2011) 

others have presented hypotheses and predictions. 

Safi et al. (2011) investigated global pat‐ 


87


terns of mammal PD and FD and found that when 

controlling mammal assemblages for their evolutionary 

history the tropics were characterized by a 

FD deficit. This suggests that more species can be 

closely packed into the ecological space in tropical 

than in temperate regions (see figure 3 in their 

paper), a paradoxical situation in which competition 

seems to limit trait evolution in a group, but 

does not decrease the co‐occurrence of species 

with similar trait values (Wiens 2011). There are 

several non‐mutually exclusive mechanisms that 

could be responsible for this pattern (see Figure 1 

in Safi et al. 2011). In temperate regions, for example, 

if resources are limited, species need to 

occupy wider ecological niches in order to secure 

their energy demands and therefore communities 

would show signs of overdispersion in functional 

traits. In addition, high environmental heterogeneity 

could also result in an overdispersion in FD 

because coexisting species could adapt and specialize 

to the different environmental conditions. 

Some light has been shed on beta‐PD patterns 

by Gómez et al. (2010), studying Neotropical 

Forest antbirds at different spatial scales. If speciation 

occurred mainly among ecoregions, there is 

a lower probability of sister species co‐occurring 

in the same ecoregion, resulting in phylogenetic 

evenness at this smaller scale. If so, we would expect 

high species turnover (taxonomic beta diversity) 

and low phylogenetic turnover (beta‐PD) 

among ecoregions, because species would tend to 

be close relatives. An alternative scenario is when 

phylogenetic structure at the regional scale is a 

product of limited dispersal of lineages. In this 

case we would expect both high species turnover 

and high beta‐PD among regions, because each 

200 

6 

180 

Any spatial scale 

160 

4 

FD 

140 

2 

PD 

published studies 

120 

100 

80 

60 

0 

2007 2008 2009 2010 2011 

beta-FD 

beta-PD 

Large spatial scale 

40 

FD 

PD 

20 

beta-PD 

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 constrained 

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 turnover”; 

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. 



region would contain distinct clades, with independent 

diversifications. Finally, if observed values 

of species turnover and beta‐PD do not differ 

from what would be expected by chance (using 

null‐models where random assemblages are built 

from the species pool), phylogenetic structure at 

the regional scale is unlikely to be the result of 

historical processes. In that case using FD should 

be better because niche‐based processes are 

more likely to explain the pattern. For example, 

along a strong environmental gradient where species 

are sorted from the regional pool according to 

their traits, we expect both species and functional 

turnover. However, if the species pool is composed 

of ecologically similar species – an indication 

that species were sorted according to their 

traits at a higher spatial scale (for example, due to 

a climatic filter or historical processes) – we 

should expect low functional turnover because 

the pool already contains very similar species. 

Also, in the absence of environmental filters, species 

turnover should occur independently of functional 

turnover (Mouchet et al. 2010). Nevertheless, 

species traits should have – at least to some 

extent – some phylogenetic signal and, therefore, 

partitioning the relative contribution of evolutionary 

history to trait dissimilarities among species 

may be important. A potential, and unexplored, 

solution is to decouple functional diversity into 

“phylogenetic structured” and “specific 

(ecological)” components. This would help us to 

better understand historical and recent processes 

on biodiversity patterns and assembly rules (Diniz‐ 

Filho et al. 2011). 

The ground is reasonably well settled to 

start “rebuilding community ecology from functional 

traits” (McGill et al. 2006) and “merging 

community ecology with evolutionary biology” 

(Cavender‐Bares et al. 2009). Yes, there are 

some methodological challenges – how to properly 

define the species pool and null models, 

which traits should be used, what is the most suitable 

measure of PD and FD, and so on (see 

Pavoine & Bonsall 2011), but we should avoid becoming 

locked into a blinkered debate about 

methodological issues. For example, in the last 

decade more than two measures of PD or FD were 

proposed, each year! This may come at the expenses 

of the more important (and exciting) steps 

of doing science: how can we move forward the 

theory by using novel approaches 

All existing hypotheses that have been applied 

to taxonomic diversity can be extended to 

phylogenetic and functional diversity (Meynard et 

al. 2011). However, PD and FD can be used to create 

more rigorous and direct predictions for most 

of the hypotheses in macroecology and biogeography, 

such as attempts to explain latitudinal patterns 

of biodiversity (Willig et al. 2003). These 

metrics also present an opportunity to formulate 

new hypotheses about how species evolutionary 

history and trait diversity are distributed across 

communities at different scales. For example, 

Wiens et al. (2011) showed situations where after 

a major evolutionary radiation within a region, the 

region can still be invaded by ecologically similar 

species from another clade, challenging the paradigm 

that communities are ‘saturated’. Largescale 

phylogenies and trait databases are currently 

becoming available for a wide range of 

taxonomic groups, facilitating estimates of FD and 

PD. Including these two aspects of biological diversity 

will be crucial if we want to advance from 

exploratory studies which report interesting relationships 

between biodiversity and environment 

to also identifying their causal mechanisms. 

Acknowledgements 

I thank Joaquín Hortal, Thiago Rangel, and Michael 

Dawson for valuable comments on the manuscript. 

This work was supported by CAPES (project 

#012/09). 

Marcus V. Cianciaruso 

Departamento de Ecologia, Instituto de Ciências Biológicas, 

Universidade Federal de Goiás, Goiânia, GO, 

Brazil. e‐mail: cianciaruso@gmail.com; 

http://www.wix.com/cianciaruso/home 

References 

Cavender‐Bares, J., Kozak, K., Fine, P. & Kembel, S. 

(2009) The merging of community ecology and 

phylogenetic biology. Ecology Letters, 12, 693– 

715. 


89


Diniz‐Filho, J.A.F, Cianciaruso, M.V., Rangel, T. & Bini, L. 

(2011) Eigenvector estimation of phylogenetic 

and functional diversity. Functional Ecology, 25, 

735–744. 

Gaston, K.J. & Blackburn, T.M. (1999) A critique for 

macroecology. Oikos, 84, 353–368. 

Gómez, J.P., Bravo, G.A., Brumfield, R.T., Tello, J.G. & 

Cadena, C.D. (2010) A phylogenetic approach to 

disentangling the role of competition and habitat 

filtering in community assembly of Neotropical 

forest birds. Journal of Animal Ecology, 79, 

1181–1192. 

Jenkins, D.G. & Ricklefs, R.E. (2011) Biogeography and 

ecology: two views of one world. Philosophical 

Transactions of the Royal Society of London B, 

366, 2331–2335. 

Losos, J.B. (2008) Phylogenetic niche conservatism, 

phylogenetic signal and the relationship between 

phylogenetic relatedness and ecological 

similarity among species. Ecology Letters, 11, 

995–1003. 

McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. 

(2006) Rebuilding community ecology from 

functional traits. Trends in Ecology and Evolution, 

21, 178–185. 

Meynard, C.N., Devictor, V., Mouillot, D., Thuiller, W., 

Jiguet, F. & Mouquet, N. (2011) Beyond taxonomic 

diversity patterns: how do α, β and γ 

components of bird functional and phylogenetic 

diversity respond to environmental gradients 

across France Global Ecology and Biogeography, 

20, 893–903. 

Mouchet, M.A., Villéger, S., Mason, N.W.H. & Mouillot, 

D. (2010) Functional diversity measures: an 

overview of their redundancy and their ability to 

discriminate community assembly rules. Functional 

Ecology, 24, 867–876. 

Pavoine, S. & Bonsall, M. (2011) Measuring biodiversity 

to explain community assembly: a unified approach. 

Biological Reviews, 86, 792–812. 

Ricklefs, R.E. (2008) Disintegration of the ecological 

community. American Naturalist, 172, 741–750. 

Safi, K., Cianciaruso, M.V., Loyola, R.D., Brito, D., Armour‐Marshall, 

K. & Diniz‐Filho, J.A.F. (2011) 

Understanding global patterns of mammalian 

functional and phylogenetic diversity. Philosophical 

Transactions of the Royal Society of London 

B, 366, 2536‐2544. 

Wiens, J.J. (2011) The niche, biogeography and species 

interactions. Philosophical Transactions of the 

Royal Society of London B, 366, 2336–2350. 

Wiens, J.J., Pyron, R.A. & Moen, D.S. (2011) Phylogenetic 

origins of local‐scale diversity patterns and 

the causes of Amazonian megadiversity. Ecology 

Letters, 14, 643–652. 

Willig, M.R., Kaufmann, D.M. & Stevens, R.D. (2003) 

Latitudinal gradients of biodiversity: pattern, 

process, scale and synthesis. Annual Review of 

Ecology, Evolution, and Systematics, 34, 273– 

309. 

Edited by Thiago F. Rangel 

Remember that being a member of IBS means you can get free online access to four biogeography 

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 Journal 

of Avian Biology. 

Additional information is available at http://www.biogeography.org/. 


ISSN 1948‐6596 

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 Spalding 

et al. (1997), is a must‐have publication for 

everyone loving and working with, in, or near to 

mangroves. It celebrates the wonderful world of 

these beautiful forests with astonishing figures 

and photographs. The informative maps and tables 

provide captivating facts about the ecological 

and economic values of mangroves and the consequences 

of their loss. 

The atlas scores with the presentation of 

recent findings on carbon sequestration, showing 

that mangroves store more carbon than tropical 

forests (Donato et al. 2011); and with the suitability 

of intact mangroves for protecting coastal regions 

against tsunamis (Wibisono and Suryadiputra 

2006). This will arm (with powerful arguments) 

ecologists, conservation biologists and policymakers, 

who urgently need to communicate this 

knowledge in order to increase public awareness 

and political willingness to protect and rehabilitate 

one of the most vulnerable ecological systems 

on earth. 

As indicated by its title, the World atlas of 

mangroves gives a comprehensive overview of the 

global distribution of mangrove species at country 

level. A detailed description of the particular 

status of mangrove systems in each country, accompanied 

by information about their specific 

threats, level of degradation and extent of rehabilitation 

programs guides the reader through a 

multitude of distinct features, while keeping similarities 

and general principles in mind. 

Mangrove experts of international repute 

contribute boxes on particular topics of interest, 

such as mangroves’ responses to climate change 

(Gilman, Duke et al.) or their functioning in highly 

dynamic coastal regions (Fromard and Proisy). 

They summarise up‐to‐date research as well as 

the hot topics that will be developed in the near 

future. In addition, the annexes containing tree 

species descriptions, national species lists and 

country fact sheets serve as an excellent compendium 

and make this atlas perfect as a quickstart 

guide for students as well as experienced researchers 

approaching a new region. 

Considering the presentation of global 

trends as the main purpose of the World Atlas Of 

Mangroves, this book fulfils expectations. Unnecessary 

uncertainties and errors in the introduction 

to the ecology of mangroves leave, however, a 

drop of bitterness. The first chapters (Mangrove 

ecosystems and Mangroves and people) notably 

omit explicit references to any publications. The 

authors state that these chapters and the boxes 

therein ‘draw heavily’ on the relevant literature, 

but information presented is confusing or even 

erroneous, and does not always reflect the content 

of the publications loosely mentioned at the 

end of each subchapter, nor established knowledge 

available in textbooks (e.g. Tomlinson 1986) 

or extended reviews (e.g. Feller et al. 2010). For 

example, the classification of mangroves into 

fringing mangroves, basin mangroves, and overwash 

mangroves is needlessly incomplete; it could 

be easily improved by following standard mangrove 

literature (e.g. Lugo & Snedaker 1974, 

Woodroffe 1992). The heterogeneous handling of 

outdated theories and debated hypotheses about 

the functioning of mangroves is also surprising. 

For instance, the editors correctly do away with 

the perspective that the land creates the capability 

for mangrove formation, but then present elevation 

and the subsequent gradient of inundation 

as the only factors driving patterns of species 

zonation. There are, however, four other major 

hypotheses to explain this striking feature: geomorphological 

influences, propagule dispersal, 

predation and species competition (see e.g. Smith 

III 1992 for detailed discussion). Further errors in 

the classification of aerating roots and also in the 

systematics and geographical distribution of some 

mangrove species have been already listed and 


91


discussed in detail by Dahdouh‐Guebas (2010). It 

remains a mystery why these chapters have not 

been written or carefully revised by the leading 

mangrove experts mentioned above, or the numerous 

others who contributed to this book with 

specific boxes. 

This volume appears 14 years after Mangroves 

– The forgotten forest between land and 

sea (Mastaller 1997). It seems that the world has 

changed and the forgotten forest has been rediscovered. 

Obviously neither the simple existence of 

this remarkable ecosystem, nor its fascinating 

functioning based on adaptation to the harsh conditions 

of tidal zones, were sufficient to convince 

people that it is worth protecting mangroves 

against aquaculture, agriculture, land use and the 

many types of waste water we produce. The 

monetary expression of the value of mangroves 

(US$ 2000–9000 ha –1 yr –1 according to the statistics 

in this book), and the change from the ecological 

perspective to the human perspective in 

terms of coastal protection against hurricanes and 

tsunamis and in carbon sequestration, is necessary 

to improve public awareness about the importance 

of mangroves for our present life and a 

critical part of our response to the challenges of 

environmental changes, including sea level rise 

and climate change. The World atlas of mangroves 

is a strong contribution towards this goal and, I 

hope, another step towards ushering in a new era 

where mangroves are valued for their beauty in 

the same way as many rain forests or coral reefs. 

In summary, if you are working in the field 

of mangrove conservation or related issues in the 

context of tropical coastal zones, or if your work is 

targeted towards practitioners, stakeholders or 

users of at‐risk mangrove ecosystem services, the 

World atlas of mangroves is your book; it will support 

your daily work with easy‐to‐understand information 

and strong facts about the ecological 

and economic values of this forest. If you are a 

mangrove ecologist, this book should also be on 

your shelf because it provides you with a quick 

overview of mangrove distribution and current 

status on Earth. It also acts as an enormous source 

of suitable maps and material to round off your 

lectures. This should convince your students that 

mangrove research is a challenge, an urgent demand 

for mankind and that being involved is an 

accolade. On the other hand, if you are looking for 

a general text spanning the interdisciplinary aspects 

of mangrove ecology, this is not the book for 

you. The roots of this book largely come from geography 

and remote sensing. If you are searching 

for an up‐to‐date text about the present scientific 

understanding and recent findings in mangrove 

research, I recommend supplementing the atlas 

with textbooks, recent reviews or more detailed 

publications on mangrove ecosystems and people’s 

depency on their health and functioning. 

Uta Berger 

Institut für Waldwachstum und Forstliche Informatik, 

Technische Universität Dresden 

e‐mail: uta.berger@forst.tu‐dresden.de; 

http://www.forst.tu‐dresden.de/SystemsAnalysis/uta‐berger 

References 

Dahdouh‐Guebas, F. (2011) World Atlas of Mangroves: 

Mark Spalding, Mami Kainuma and Lorna Collins 

(eds). Human Ecology, 39, 107–109. 

Donato, D.C., Kauffman, J.B., Murdiyarso, D., Kurnianto, 

S., Stidham, M. & Kanninen, M. (2011) Mangroves 

among the most carbon‐rich forests in 

the tropics. Nature Geoscience, 4, 293–297. 

Feller, I.C., Lovelock, C.E., Berger, U., McKee, K.L., Joye, 

S.B. & Ball, M.C. (2010). Biocomplexity in Mangrove 

Ecosystems. Annual Review of Marine 

Science, 2, 395–417. 

Lugo, A.E. & Snedaker, S.C. (1974). The ecology of mangroves. 

Annual Review of Ecology and Systematics, 

5, 39–64. 

Mastaller, M. (1997) Mangroves – the forgotten forest 

between land and sea. Tropical Press Sdn. BhD. 

Kuala Lumpur, Malaysia. 189 pp. 

Smith III, Th.J. (1992). Forest Structure. In: Tropical 

mangrove ecosystems (ed. by A.I. Robertson and 

D.M. Alongi), pp.101–136. American Geophysical 

Union, Washington. 

Spalding, M., Blasco, F. & Field, C. (1997). World mangrove 

atlas. The International Society for Mangrove 

Ecosystems, Okinawa, Japan. 178 pp. 

Tomlinson, P.B. (1986). The botany of mangroves. Cambridge 

University Press, Cambridge, UK. 419 pp. 

Wibisono,I.T.C. & Suryadiputra, N.N. (2006). Study of 

lessons learned from mangrove/coastal ecosystem 

restoration efforts in Aceh since the tsunami. 

Wetlands International – Indonesia Programme, 

Bogor. 86 pp. 


ISSN 1948‐6596 


Woodroffe, C.D. (1992). Mangrove sediments and geomorphology. 

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 biogeography 

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 

of plant and animal species in the world is continuing 

to decline in spite of ambitious targets set 

by governments to prevent this (Butchart et al. 

2010). It is also becoming evident that the continued 

functioning of ecosystems depends on this 

diversity (Isbell et al. 2011). In order to conserve 

what is left of biodiversity, it is crucial that we understand 

the diversity of life and how it is distributed 

across the biomes and ecosystems of the 

world. Since understanding the distribution of biodiversity 

is a central tenet of biogeography, it 

seems obvious that the field of biogeography 

should be of central importance in conservation. 

In this volume, Richard Ladle and Robert 

Whittaker bring together chapters by a number of 

biogeographers to summarise progress to date in 

applying the principles of biogeography to conservation 

and to identify areas where there is still 

work to be done. The book is a comprehensive but 

digestible summary of the field of conservation 

biogeography and should make essential reading, 

not only for the students at whom it is primarily 

aimed, but also for more experienced scientists. 

The editors profess at the outset that the aim was 

to achieve a degree of coherence among the 

chapters, an aim that is achieved remarkably well 

to give a very coherent text. 

The first section of the book provides a brief 

but interesting history of the conservation movement 

and the contrasting values held by different 

sectors of this movement (Chapters 2 and 3), as 

well as some background to the field of conservation 

biogeography (Chapter 1). A distinction is 

made between approaches that focus on the composition 

of biological communities and those that 

focus on ecosystem function through an understanding 

of ecosystem processes such as nutrient 

cycling (p. 31). An interesting and growing field in 

ecology, which receives little attention in the 

book, uses the functional traits of species to explain 

the link between the composition of biological 

communities and the function of the ecosystems 

that contain them. Functional traits – such as 

body mass, diet, habitat affinity and development 

mode of animals, and height and photosynthetic 

pathway of plants – can help explain how species 

contribute to the processes underlying the functioning 

of ecosystems and can also help in predicting 

how ecosystems will respond to environmental 

change (McGill et al. 2006). 

The second section reviews our current understanding 

of the distribution of biodiversity, 

summarises the history of the global protected 

areas network and describes the methods available 

for more systematically representing biodiversity 

in future extensions to this network. There 

is a strong terrestrial focus here, indeed throughout 

the entirety of the book, which the authors 

acknowledge and which is owing to a less complete 

understanding of the distribution of diversity 

in the oceans and in freshwater habitats. It is 

worth noting, though, that the Census of Marine 

Life, an ambitious $650 million project that finished 

recently, has made huge progress towards 

understanding the biogeography of the oceans 


93


(e.g. see Tittensor et al. 2010). Even in the terrestrial 

realm, knowledge about the number and 

identity of the world’s species and how they are 

distributed remains very far from complete: the 

Linnaean and Wallacean shortfalls respectively 

(Chapter 4). A recent paper (Joppa et al. 2011) 

addressed both of these knowledge gaps simultaneously 

by predicting the spatial distribution of 

undiscovered plant species, predicting that most 

new plant species will be discovered in areas already 

identified as hotspots of plant diversity, emphasising 

the importance of these areas for conservation. 

Chapter 5 provides an excellent summary 

of the many different types of protected 

areas in the global network and the different values 

that underpin these, while Chapter 6 provides 

a useful and succinct review of the enormous and 

ever‐growing literature on systematic conservation 

planning. 

The third section of the book describes how 

the tools of biogeography can be used to plan for 

environmental change in conservation. This is the 

only part of the book where the chapters appear 

somewhat disjointed, but this is probably owing to 

the attempt to summarise a vast literature in a 

very small number of chapters. Nevertheless, the 

chapters in this section provide excellent descriptions 

of some of the available methods, from phenomenological 

models that infer future changes 

from current patterns (Chapter 7) to more process 

‐based models that use the theory of island biogeography 

to predict the consequences for biodiversity 

of shrinking and increasingly isolated natural 

habitat patches (Chapter 8). Chapter 9 deals 

with invasive species, which are an important 

driver of environmental change, and the homogenisation 

of biological communities, i.e. the erosion 

of beta diversity. Most of the studies investigating 

broad‐scale patterns of diversity have focused 

on inventory diversity, commonly measured 

as species richness, and it is only recently that 

studies have attempted to map beta diversity (e.g. 

McKnight et al. 2007) and to relate it to spatial 

and environmental factors (e.g. Ferrier et al. 

2007). 

With a growing need to understand changes 

in the natural environment and the impact of 

these changes on human society, the emerging 

field of conservation biogeography is likely to become 

increasingly important in providing the necessary 

theoretical basis and tools for doing so. 

This book provides an excellent foundation for 

that field and is highly recommended reading for 

students, scientists and practitioners of conservation. 

Tim Newbold 

United Nations Environment Programme World Conservation 

Monitoring Centre, Cambridge, UK 

e‐mail: Tim.Newbold@unep‐wcmc.org; 

http://www.unep‐wcmc.org/tim‐newbold_368.html 

References 

Butchart, S.H.M., Walpole, M., Collen, B. et al. (2010). 

Global biodiversity: indicators of recent declines. 

Science, 328, 1164–1168. 

Isbell, F., Calcagno, V., Hector, A. et al. (2011). High 

diversity is needed to maintain ecosystem services. 

Nature, 477, 199–202. 

Joppa, L.N., Roberts, D.L., Myers, N. et al. (2011). Biodiversity 

hotspots house most undiscovered plant 

species. Proceedings of the National Academy of 

Sciences of the United States of America 108, 

13171–13176. 

McGill, B.J., Enquist, B.J., Weiher, E. & Westoby, M. 

(2006). Rebuilding community ecology from 

functional traits. Trends in Ecology & Evolution, 

21, 178–185. 

McKnight, M.W., White, P.S., McDonald, R.I., Lamoreux, 

J.F., Sechrest, W., Ridgely, R.S. & Stuart, 

S.N. (2007). Putting beta‐diversity on the map: 

broad‐scale congruence and coincidence in the 

extremes. PLoS Biology, 5, e272. 

Tittensor, D.P., Mora, C., Jetz, W., Lotze, H.K., Ricard, 

D., Vanden Berghe, E. & Worm, B.(2010). Global 

patterns and predictors of marine biodiversity 

across taxa. Nature, 466, 1098–1101. 


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 advertised, 

please contact Karen Faller (faller@wisc.edu) or Michael Dawson 

(mdawson@ucmerced.edu) with details. 


ISSN 1948‐6596 


ries of key topics relating (in this case) to biological 

invasions, without citations but with relevant 

further reading at the end. The entries vary in 

length from 1 to 8 pages, and often incorporate 

useful figures and occasionally tables. The book is 

impressively glossy (all figures are in full colour) 

and well presented, which is all the more remarkable 

considering the relatively modest price. The 

editors, Daniel Simberloff and Marcel Rejmánek, 

are leading invasion ecologists and are well qualified 

to compile such a text; this is reflected not 

just in the broad range of well‐selected topics that 

the volume includes (of which there are 153) but 

also the roll‐call of esteemed contributors that 

have supplied the entries (of which there are 197, 

many of them high‐profile international researchers). 

The book is aimed not just at an academic 

audience, however, and the articles are written 

with the interested and educated general public in 

mind. 

The individual articles cover various aspects 

of invasions, ranging from particular attributes of 

invasive species and invaded ecosystems to impacts 

and management, interesting case studies 

and historical perspectives. Clearly it is not possible 

to cover all of the entries in a review such as 

this, but I did find several articles especially interesting, 

particularly because they highlight the 

many socioecological factors that complicate our 

relationships with potentially problematic species. 

The entry on Xenophobia for example does an excellent 

job of summarising how society’s relationship 

with non‐native species is constructed in certain 

ways by the use of loaded terms or cultural 

metaphors, for example the negative personification 

of zebra mussels as ‘outlaws’ on the west 

coast of the US, or the badging of ‘harmful’ or 

‘distasteful’ species with appellations that note 

their foreign status (Japanese knotweed, Chinese 

mitten crab, English sparrow and so on). As a 

starting point for a discussion of scientific objecbook 

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 

(Davis 2005), invasion ecology/biology is in many 

ways a nascent and emerging field, and is still engendering 

discussion regarding whether it indeed 

truly exists as a field or discipline in its own right, 

or is rather a particularly focused aspect of community 

ecology or biogeography (e.g. Marris 2009, 

Pyšek and Hulme 2009). As with many ecological 

disciplines, invasion ecology has seen fundamental 

disagreements over aspects ranging from core 

definitions (including ‘invasion’ itself; Falk‐ 

Petersen et al. 2006, Ricciardi and Cohen 2007) to 

level of scientific objectivity (e.g. Larson 2007). 

The field is at a stage in its development where (1) 

dedicated journals exist (e.g. Biological Invasions) 

and there is a substantial number of academic 

articles published every year (for example a 

search of ‘invasive species’ in Web of Knowledge 

returns 1181 articles published in 2010 alone), 2) 

there is clear and significant international interest 

and action in relation to invasions and (3) an extended 

peer community is involved in researching 

and managing the threat of invasive species, from 

world‐leading academics at research‐intensive 

universities to local government and conservation 

volunteers. The result of the burgeoning information 

and uneven levels of understanding and focus 

across the peer community is confusion and uncertainty, 

right from the fundamentals (what is an 

invasive species exactly, and why is it invasive) to 

the specifics (what is the best technique for reducing 

populations of Crassula helmsii in my pond, 

and how does that differ from managing spread in 

the local lake). The time is ripe therefore for an 

encyclopaedia such as this one by Daniel Simberloff 

and Marcel Rejmánek to form a baseline for 

future definitions and discussions. 

The book is one of University of California 

Press’ Encyclopedias of the Natural World series, 

and as with the other volumes has a wide range of 

entries that are effectively short essays or summa‐ 


95


tivity related to invasion biology it works exceptionally 

well, and is exactly the right size for digestion 

by students or interested amateurs. 

Indeed, one of the best uses I find for reference 

works such as these are as opening forays 

into topics for class discussions, whether at graduate 

or undergraduate level. Good examples include 

the entry on Succession, which very effectively 

and concisely summarises key concepts that 

take up whole chapters in many textbooks, and 

although invasion biology is only addressed towards 

the end, it is clear how the two link together. 

Likewise, the discussion on Native invaders, 

in which issues of ‘invasive’ terminologies 

(and when they are appropriate) are covered, is 

excellently written and illuminating at a range of 

levels, particularly in relation to the many examples 

of ‘invasion’ given. Certainly students and 

researchers new to the subject will have any initial 

confusion over what is meant by invasions dispelled 

by the article, and it will also help them to 

think objectively about whether a species really 

may be considered invasive or not. All of the articles 

I read through were of a high quality and well 

written/edited, with very little wasted space for 

such a large volume (although on occasion figures 

are not always relevant – I’m not sure why an image 

of Frank Buckland ‘physicking a porpoise’ 

(page 2) is worthy of inclusion for example, 

despite his role in founding the main UK acclimatisation 

society). 

Of course, it is always hard to get the right 

balance between conciseness and detail in such 

entries, and to retain the relevant focus. The 

opening entry, Acclimatisation societies is a case 

in point: the article does an excellent job of summarising 

the development and impact of such societies 

in different countries, many of which were 

responsible for the introduction of significant 

numbers of non‐native species around the globe 

before dying out in the face of increasing legislation, 

awareness of ecological risk from introductions 

and lack of interest from the general public. 

The article elegantly conveys how originally benevolent 

intentions, such as the introduction of 

non‐natives to improve food resources, control 

pests and to soothe homesick colonists (among 

other reasons), in most cases failed to be realised 

and also (with some notable exceptions) that 

many societies were unsuccessful in actually naturalising 

many species at all. But much is left unsaid: 

in some cases one is left wanting to know 

more about whether species referred to as 

‘released’ became naturalised, whether regions 

such as South America maintained any such societies 

(these countries are ignored, while others 

such as Germany and Italy receive only one sentence) 

and ultimately whether such societies indirectly 

provided evidence to force their own discontinuation. 

As a taster to whet the appetite, the 

article succeeds very well (and relevant books on 

the subject are provided in the Further Reading 

section), but it is not an authoritative, encyclopaedic 

summary in itself. 

As with any vast topic, covering all aspects 

in a single volume is difficult – in this case there is 

differential coverage of ecosystems (e.g. entries 

for canals, lakes, rivers and wetlands, but no coverage 

of urban ecosystems, despite these being 

important points of introduction for some invasive 

taxa); hypotheses (e.g. Enemy Release Hypothesis, 

Novel Weapons Hypothesis, but no Tens Rule); 

geographical areas (Australia, the Great Lakes, 

Hawaiian islands, the Mediterranean, the Ponto‐ 

Caspian, New Zealand and South Africa receive a 

particular focus) and species (good examples of 

some key species or groups such as zebra mussel, 

earthworms and fishes, but understandably not 

comprehensive coverage). This is entirely reasonable, 

and is not a criticism of the volume – it is 

impossible to cover the vast range of topics associated 

with biological invasions in sufficient depth 

in a single volume, and the material that is included 

is impressive. The division of the book between 

invader attributes, processes, taxa, ecosystems, 

pathways to invasion and so on is very well 

done and represents a huge effort on the part of 

the editors, for which they should be roundly congratulated. 

I would encourage consideration of a 

second volume, however, at least with regard to 

key concepts and hypotheses. The opening guide 

to the Encyclopedia notes that there is a website 

with a list of articles, sample entries and so, and 

notes that the site ‘will evolve with the addition of 


ISSN 1948‐6596 

new information’, p. xxii). The web address has 

since changed and I was unable to locate the new 

one. Though I happily agree that this could potentially 

be a very useful resource, given the rapidly 

changing environment of the internet, the publication 

of a second volume would perhaps be the 

most reliable option. 

In summary, this is an excellent reference 

work that combines readability with academic 

rigour throughout. Its broad coverage of the field, 

high quality of production and reasonable price 

makes it an essential purchase for any university 

with departments teaching or researching within 

the broad spectrum of ecology, as well as for individual 

researchers of species invasions. 

Robert A. Francis 

Department of Geography, King’s College London 

e‐mail: robert.francis@kcl.ac.uk; http://rg.kcl.ac.uk/ 

staffprofiles/staffprofile.phppid=1961 

References 


Davis, M.A. (2005) Invasion biology 1958‐2004: the 

pursuit of science and conservation. In: Conceptual 

ecology and invasions biology: reciprocal 

approaches to nature (ed. by Cadotte, W.M, 

McMahon, S.M. and Fukami, T.) , pp. 35–64. 

Kluwer Publishers, London. 

Falk‐Petersen, J., Bøhn, T. & Sandlund, O.T. (2006) On 

the numerous concepts in invasion biology. Biological 

Invasions, 8, 1409–1424. 

Larson, B.M.H. (2007) An alien approach to invasive 

species: objectivity and society in invasion biology. 

Biological Invasions, 9, 947–956. 

Marris, E. (2009) The end of the invasion Nature, 459, 

327–328. 

Pysek, P. & Hulme, P.E. (2009) Invasion biology is a discipline 

that’s too young to die. Nature, 460, 324 

–324. 

Ricciardi, A. & Cohen, J. (2007) The invasiveness of an 

introduced species does not predict its impact. 

Biological Invasions, 9, 309–315. 


the planet and around 10% of all vertebrate species. 

James Albert and Roberto Reis’ goal as editors 

of the Historical Biogeography of Neotropical 

Freshwater Fishes is to examine the evolutionary 

forces responsible for this diversity. In doing so 

they make the case that multiple processes of diversification 

were involved and that these operated 

over long periods of time as well as on a continental 

scale. The book itself is divided into two 

parts, the first of which examines current knowledge 

on the biogeography of the region, while the 

second is a regional analysis that links contemporary 

geographical patterns with geological history. 

The book is ambitious in scope and brings together 

previously fragmented material to provide 

an authoritative overview of this impressive group 

of fish. And while a fish‐eye view of the Neotropical 

ichthyofauna is inevitably drawn to the Amabook 

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 

everyone who visits them. The seeds of some of 

the most important concepts in ecology and evolution 

were sown during the South American travels 

of influential 19 th century thinkers. For example, 

the latitudinal gradient of diversity, now recognized 

as ecology’s oldest pattern (Hawkins, 

2001), was first identified by von Humboldt, while 

Bates documented the variety and adaptations of 

species in Amazonian forests, and Wallace and 

Darwin pondered the mechanisms responsible for 

the myriad forms of life they encountered. Although 

the Neotropics have played a crucial role 

in our understanding of the diversity of life on 

earth, in many ways they continue to represent an 

unexplored frontier. This is particularly clear in the 

case of Neotropical freshwater fish, a group estimated 

to consist of more than 7000 species, and 

that accounts for over half the freshwater fish on 


97


zon, the book has broad coverage, embracing the 

Andes and extending through Central America and 

into southern Mexico. As it makes clear, it is necessary 

to have a continental perspective to understand 

the diversity and distribution of this impressive 

group. 

I particularly liked the care and thought involved 

in putting the book together. It is a beautifully 

presented volume with informative tables 

and figures, many of them in colour. However, 

more important than this is that the editors have 

a strong sense of what the important issues are 

and how these should be best dealt with. Indeed 

the book is an essential reference for anyone 

wanting to learn more about the diversity or history 

of South American fishes. 

One of the most challenging questions in 

ecology is explaining why different habitats support 

different numbers of species. The extent of a 

habitat accounts for much of the variation but 

South America has an excess of species relative to 

its area. The core of the continent, particularly the 

Amazon, is responsible for a disproportionate 

amount of this diversity. It is tempting to attribute 

this exceptional richness to the unique geological 

and environmental features of the Amazon. However 

many of the fishes that inhabit this river system 

are older than the Amazon Basin itself. Moreover, 

the Amazonian ichthyofauna has been accumulated 

gradually through tens of millions of 

years. The explanation, Albert, Petry and Reis argue, 

is rooted in the repeated subdivision and 

merging of adjacent river basins and their faunas, 

with dispersal limitation and environmental filtering 

playing important roles. The exceptionally high 

diversity seems to be less to do with exceptional 

speciation rates than with low rates of extinction. 

However, diversity is not just a measure of the 

numbers of species that co‐occur but also of the 

types of species that are found together. A universal 

feature of natural assemblages is that some 

families contribute a much higher fraction of species 

than others. The Neotropics are no exception. 

Ten families of fish account for 75% of the 

Neotropical icthyofauna. Characidae (including 

piranhas and tetras) and Cichlidae (such as discus) 

are particularly big hitters. One possibility is that 

this unevenness is simply the result of chance. 

Alternatively, historical and biological factors, either 

separately or together, could contribute. E.O. 

Wilson (2003) has argued that an ancient origin, 

combined with small body size, widespread geographic 

distribution and key innovations contribute 

to the success of some groups relative to others. 

On the basis of the evidence presented by 

Neotropical fish, Albert, Bart and Reis conclude 

that these features are necessary but not sufficient. 

Indeed they note that clades can be ancient 

(e.g. Arapaima, which is of Cretaceous origin), 

widespread (Arapaima again) or with small body 

size (e.g. Amazonsprattus) yet be represented by a 

handful of species at most. On the other hand sexual 

and trophic innovation may play a role. Ecological 

specialisation is also important. For example, 

Crampton notes that groups of closely related 

Gymnotiform electric fish species tend to be 

found in a narrow range of habitat types but may 

be spread across large geographic areas. The factors 

that underpin diversification are the same as 

those that come into play in the explosive speciation 

that characterizes the African rift lakes. The 

difference here is that the game is played out on a 

continental scale as opposed to a local arena. 

Of course, much remains to be learnt about 

the phylogenetic histories of Neotropical fishes 

and of the geological context in which these species 

evolved. Nonetheless, as this book makes 

clear, the nature and timing of key events is becoming 

much better understood. The contributions 

to the book demonstrate how the growing 

body of molecular data, and its integration with 

ecological theory and earth sciences, has underpinned 

the recent and rapid progress in understanding 

this system. 

Your participation in frontiers of biogeography is encouraged. Please send us your articles, comments 

and/or reviews, as well as pictures, drawings and/or cartoons. We are also open to suggestions 

on content and/or structure. 

Please check http://www.biogeography.org/html/fb.html for more information, or contact us at 



ISSN 1948‐6596 

There have been many studies of tropical 

diversity but until now Neotropical fishes fish have 

received relatively little attention. This contrasts 

with South American birds, a group that has been 

prominent in tests of macroecological hypotheses 

(e.g. Rahbek et al., 2007). Fish are responsible for 

more diversity and deserve to be more fully studied. 

This book provides the knowledge that will 

inform these exciting research opportunities. 

Anne E. Magurran 

University of St Andrews 

e‐mail: aem1@st‐andrews.ac.uk; 

http://biology.st‐andrews.ac.uk/magurran/ 

References 


Hawkins, B. A. (2001). Ecology’s oldest pattern. Trends 

in Ecology and Evolution 16, 470. 

Rahbek, C., Gotelli, N. J., Colwell, R. K., Entsminger, G. 

L., Rangel, T. F. L. V. B. and Graves, G. R. (2007). 

Predicting continental‐scale patterns of bird 

species richness with spatially explicit models. 

Proceedings of the Royal Society B: Biological 

Sciences 274, 165‐174. 

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). Harvard 

University Press. 


books noted with interest 

Principles of terrestrial ecosystem ecology 

F. Stuart Chapin III, Pamela A. Matson & Peter 

M. Vitousek 

2011, 2nd edition, Springer, 529 pp. 

£135 (Hardback), £44.99 (Paperback) 

ISBN: 9781441995032 / 9781441995025 

http://www.springer.com/ 

An outstanding textbook which, after definitions, 

sets the stage with primers on Earth’s climate system 

and geological processes. What follows is a 

magisterial and comprehensive account of the 

movements of water, energy, carbon and nutrients 

though natural systems. Along with standard 

generalisations, the authors delve into the finer 

detail and explain how biological processes can 

have important modulating effects through space 

and time. A final reflective pair of chapters considers 

global changes and the implications for ecosystem 

management. The book is well written 

throughout and punctuated with excellent colour 

illustrations; no‐one from undergraduates to established 

researchers can fail to learn something 

from it. 

Guide to standard floras of the World: 

An annotated, geographically arranged 

systematic bibliography of the 

principal floras, enumerations, checklists 

and chorological atlases of different 

areas 

David F. Frodin 

2001, 2 nd edition, Cambridge University Press, 

1100 pp. 

£198 (Hardback), £90 (Paperback), US$120 (ebook) 

ISBN: 9780521790772 / 9780521189774 

http://www.cambridge.org/ 

While not generally our policy to feature reprints, 

this standard text has newly appeared in paperback, 

bringing it within affordable reach of a 

greater number of researchers. It does exactly 

what it says on the cover, making it the definitive 

reference for anyone commencing work on the 

flora of a new region. Despite its not receiving any 

further updates and its coverage ending in 1999, 

there remain no resources to rival it, either in 

print or online. It also contains insightful reviews 

on the history of floristic description. An essential 

book which belongs in the library of every plant 

biogeographer. 


99


Field guide Afghanistan: Flora and 

vegetation 

Siegmar‐W. Breckle & M. Daud Rafiqpoor 

2011, Scientia Bonnensis, Bonn, 864 pp. 

Price: Contact publishers 

ISBN: 9783940766304 

http://www.scientia‐bonnensis.com/ 

The flora of this vast, environmentally diverse and 

biogeographically central country has yet to be 

fully catalogued, but this field guide represents a 

landmark accomplishment on the path to doing 

so, filling an anomalous gap at the junction of several 

floristic realms. It contains a pictorial guide to 

over 1200 species (>25% of the flora) plus general 

chapters on vegetative formations and should facilitate 

both local and international study. Copies 

have been freely distributed to universities and 

institutes throughout Afghanistan as well as herbaria 

and museums worldwide. A feature on this 

project is planned for a future edition of Frontiers 

of Biogeography. 

Community ecology 

Peter J. Morin 

2011, 2nd edition, Wiley‐Blackwell, 407 pp. 

£90 (Hardback), £34.99 (Paperback) 

ISBN 9781444338218 / 9781405124119 

http://www.wiley.com/ 

Community ecology straddles conventional interaction‐based 

ecology and biogeography; recent 

heated debate in the pages of American Naturalist 

has even disputed whether communities truly exist 

as natural entities. Unsurprisingly the author 

makes a strong case for communities, stressing 

patterns and processes that can only be understood 

at this level, and pleasingly devotes equal 

attention to both models and experimental data. 

The textbook is intended for a graduate course 

and represents a major update on the previous 

edition. One might query the balance of coverage 

of various topics but nevertheless this remains the 

only textbook exclusively devoted to this scale of 

study. 

Markus Eichhorn 

Book Review Editor. 

e‐mail: Markus.Eichhorn@nottingham.ac.uk 

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 providing 

brief details of new publications. Authors, editors or third parties are invited to suggest books for review 

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 markus.eichhorn@nottingham.ac.uk. We 

welcome offers to review books for Frontiers of Biogeography, but will not accept an offer to review a specific 

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, expressing 

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 teaching 

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 references. 

Authors may suggest a short heading for the review, followed by the title of the book(s), the authors/editors, 

publisher, publication date, price, hbk/pbk, pages, ISBN and website (where available). 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 

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may be published. Book reviews will usually go through a light editorial review, though in some circumstances 

also will be considered by one or more referees. 


ISSN 1948‐6596 


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: rosa.chef@gmail.com; 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 estimating 

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 assessing 

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 

starkly contrasts the need to conserve biodiversity 

against our inability to inventory and protect all 

species individually. Knowledge about biodiversity 

remains insufficient because many species are still 

not described (the "Linnean Shortfall"; Brown 

and Lomolino 1998) and the distributions of described 

species often are inadequately defined 

(the "Wallacean Shortfall"; Lomolino 2004). It is 

therefore essential to identify threatened species 

and describe their distributions using approaches 

that overcome the time and budget constraints of 

systematic conservation planning. 

Araújo et al. (2007) demonstrated the need 

for additional protected areas for the effective 

conservation of the diversity of plants and vertebrates 

in the Iberian Peninsula. Preliminary data 

suggest that the existing network of reserves also 

would be ineffective in representing invertebrate 

species (Verdú and Galante 2009). Unfortunately, 

the conservation of invertebrates faces serious 

challenges due to their high diversity, complex life 

cycles and difficult taxonomy, among other factors 

(see New 1998). 

Geographic Information Systems (GIS) significantly 

advanced the conservation of endangered 

species because they allow us to delimit 

species’ potential distributions (e.g. Hortal et al. 

2005), to control their populations 

(e.g. Davies et al. 2005), to analyze their niche 

(Peterson et al. 2002), design networks of protected 

areas (e.g. Pearce and Boyce 2006), and to 

forecast the future (e.g. Hill et al. 2002). Together, 

the databases taken from atlases, museums and 

herbaria have emerged as a valuable source of 

species’ occurrence records (e.g. Elith and Leathwick 

2007). Unfortunately, these data from heterogeneous 

sources may contain errors or 

have been obtained using a biased sampling procedure 

(Hortal et al. 2007, 2008, Newbold 

2010). Besides, they do not usually provide reliable 

absences needed to perform consistent predictive 

models (Anderson et al. 2003, Lobo et al. 

2007), so alternatives have been sought generating 

models based only on presences (Hirzel et al. 

2002, Pearce and Boyce 2006), sometimes employing 

pseudo‐absences obtained in different 

ways (Zaniewski et al. 2002, Engler et al. 2004, 


101

SDM applied to invertebrate conservation 

Lobo et al. 2006, 2010). 

For my doctoral thesis, I evaluated the utility 

of SDMs for the conservation of threatened 

invertebrates in the Iberian Peninsula (Chefaoui 

2010). The majority of the species studied 

here have been designated by the European Union 

as species of “community interest” requiring 

protection and conservation (Habitats Directive). I 

used presence‐only data on Iberian threatened 

invertebrates obtained from museums, atlases 

and databases. I applied presence‐only methods 

such as ENFA (Ecological Niche Factor Analysis) 

and MDE (Multi‐Dimensional Niche Envelope), in 

addition to other methods that require presences 

and absences (here, pseudo‐absences): GAM 

(Generalized Additive Models), GLM (Generalized 

Linear Models) and NNET (Neural Networks Models). 

I approached methodological issues concerning 

the difficulties associated with predicting the 

distribution of species when reliable absence data 

are not available, and explored the possibilities of 

SDMs as a tool for conservation of endangered 

and threatened Iberian invertebrates. In this respect, 

I explored the applications of SDM to estimate 

species ranges, identify suitable habitats and 

the primary factors affecting species’ distribution 

in order to assess the conservation status of 

threatened invertebrates. 

Dung beetle populations, which are in decline 

in the Iberian Peninsula, play a critical ecological 

role in extensive pasture ecosystems by 

recycling organic matter. We delimited the potential 

distribution of the two species of Copris 

(Coleoptera, Scarabaeidae) that inhabit the Iberian 

Peninsula using ENFA (Chefaoui et al. 2005). 

ENFA is a presence‐only method that compares 

the environmental values of the localities where 

the species has been observed with respect to the 

environmental values of the territory studied 

(Hirzel et al. 2002). We explored the environmental 

niche occupied by each species in a small 

region, the Community of Madrid (CM), to restrict 

the role of dispersal constraints discriminating 

possible areas of co‐occurrence and identifying 

the specific environmental characteristics of each 

species. We identified that solar radiation and the 

presence of calcareous soils are critical to the 

presence of Copris hispanus, while Copris lunaris 

requires siliceous soils and high rainfall. Both Copris 

species are distributed along a geographic and 

environmental gradient from the Tajo basin 

(warmer, dryer, with strong annual weather variations) 

where only C. hispanus is found, towards 

the mountain slopes of the Sistema Central 

(colder, higher rainfall) where C. lunaris predominates. 

The environmental niches of both species 

are distributed along a Dry‐Mediterranean to Wet 

‐Alpine axis, and overlap in areas of moderate 

temperatures and precipitations in the north of 

CM. 

We also studied the degree of protection of 

key populations of C. hispanus and C. lunaris, making 

a proposal to improve their conservation. To 

evaluate the conservation status of Copris species, 

we took into account the size of protected sites as 

well as the values of habitat suitability in each 

protected natural site and Natura 2000 network. 

We found that Copris species were poorly conserved 

in the previous protected sites network: 

for C. hispanus only two protected sites measured 

around 30 km 2 , and for C. lunaris a single area 

measured 183 km 2 . However, protection provided 

by Sites of Community Importance (SCIs) seems to 

improve the general conservation status of these 

species in CM because the area and connectivity 

of protected sites have been increased substantially. 

Chefaoui and Lobo (2008) assessed the effects 

of pseudo‐absences on model performance 

when reliable absence data are not available. We 

compared seven procedures to generate pseudoabsence 

data to be used in GLM‐logistic regressed 

models. These pseudo‐absences were selected 

randomly or by means of presence‐only methods 

(ENFA and MDE) to model the distribution of a 

threatened endemic Iberian moth species 

(Graellsia isabelae). Our purpose was to show the 

possibility of achieving different forecasted distributions 

depending on the method and the threshold 

used to select these pseudo‐absences. 

The results showed that the pseudoabsence 

selection method greatly influenced the 

percentage of explained variability, the scores of 

the accuracy measures and, most importantly, the 


Rosa M. Chefaoui 

predicted range size. As we extracted pseudoabsences 

from environmental regions further 

from the optimum established by presence data, 

the models obtained better accuracy scores, and 

over‐prediction increased. Conversely, the profile 

techniques that generated wider unsuitable areas, 

produced functions with lower percentages of 

explained deviance and poorer accuracy scores, 

but more restricted predictive distribution maps, 

similar to the observed distribution. The random 

selection of pseudo‐absences generated the most 

constrained predictive distribution map. 

Based on results of the aforementioned 

work, we identified the environmental variables 

most relevant for explaining the distribution of 

Graellsia isabelae and assessed this species’ conservation 

status (Chefaoui and Lobo 2007). We 

modeled the potential distribution of the insect by 

performing GLM with pseudo‐absence data selected 

from an ENFA model. We found that the 

best predictor variables were summer precipitation 

(ranging from 1250 mm to 3250 mm), aridity, 

and mean elevation. This species prefers habitats 

with mid‐range mountain conditions. With respect 

to host plants, the presence of G. isabelae was 

associated mainly with Pinus sylvestris and P. nigra. 

Moreover, we found 8 areas exclusively in 

the eastern Iberian territory, and a larger unoccupied 

habitat in the western Iberian Peninsula, indicating 

that this species is probably not in equilibrium 

with its environment because of historical 

factors (Chefaoui and Lobo 2007). We suggested 

that the current distribution of the species 

was associated with the dynamism of its host 

plants during glacial periods of the Holocene, 

when the forests of Pinus sylvestris decreased 

strongly in the northwestern part of the peninsula. 

After analyzing the possibility of connectivity 

and fragmentation of the eight populations delimited 

as well as the degree of protection of G. isabelae 

on the SCIs, we found that the SCIs under 

protection did not seem sufficient to maintain current 

populations. Moreover, our study rejected 

the idea that the species was expanding its range 

due to reforestation. Because the conservation of 

G. isabelae depends on the forests of Pinus sylvestris 

and P. nigra located both inside and near to 

SCIs, we suggested that the reintroduction of the 

species in these habitats could improve its conservation. 

To understand the limitations and possibilities 

of SDM techniques, we evaluated the effects 

of species’ traits and data characteristics on the 

accuracy of SDMs for red‐listed invertebrates 

(Chefaoui et al. 2011). We applied three SDM 

techniques (GAM, GLM and NNET) using pseudoabsences 

to model the distribution of 20 threatened 

Iberian invertebrates. We correlated the 

accuracy of the obtained models with several data 

characteristics and species’ ecological traits. We 

examined two data characteristics, the amount of 

data (N) and the relative occurrence area (ROA), 

and both significantly affected the accuracy of the 

models. Greater AUC values and higher sensitivity 

scores were obtained from samples for which 

there were more than 200 records. In general, 

species whose distributions were most accurately 

modelled were those with a greater sample size or 

smaller ROA. In addition, species related to habitats 

that are problematic to detect using GIS data, 

such as riparian or humid areas, seemed to be 

more difficult to predict. 

Summary 

The performance of SDMs depends on the type of 

data and the characteristics of the species. Presence‐only 

methods (ENFA and MDE) achieved 

worse validation results and overpredicted more 

than techniques using pseudo‐absences. Nevertheless, 

presence‐only methods can be very useful 

for obtaining pseudo‐absences and discovering 

the environmental response of species. The 

method of pseudo‐absence selection strongly determined 

the predicted range size, generating different 

model predictions along the gradient between 

potential and realized distributions. There 

is an added difficulty in obtaining predictions that 

closely approximate the realized distribution of 

species under non‐equilibrium conditions, because 

both presence and absence data may be 

possible under similar environmental conditions. 

Irrespective of the approach used, species’ distributions 

are modelled more accurately when sam‐ 


103

SDM applied to invertebrate conservation 

ple sizes are larger. Species in habitats that are 

difficult to detect using GIS data, such as riparian 

species, thus may tend to be more difficult than 

most to predict. 

Availability of thesis 

Printed and PDF copies are available in the Science 

Faculty Library, Universidad Autónoma de 

Madrid (http://biblioteca.uam.es/ciencias/). A 

PDF copy is also available at request from the author. 


I would like to thank my two supervisors, Jorge M. 

Lobo and Joaquín Hortal for their support and encouragement. 

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Chefaoui, R.M. (2010) Modelos predictivos aplicados a 

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ibero‐baleares. Ph.D. Thesis. Universidad Autónoma 

de Madrid, Departamento de Biología, 

Facultad de Ciencias, 196 pp. 

Chefaoui, R.M., Lobo, J.M. & Hortal J. (2011) Effects of 

species’ traits and data characteristics on distribution 

models of threatened invertebrates. Animal 

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C.D. (2005) The re‐expansion and improving 

status of the silver‐spotted skipper butterfly 

(Hesperia comma) in Britain: a metapopulation 

success story. Biological Conservation, 124, 189– 

198. 

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from museum and herbarium records 

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adaptive regression splines. Diversity 

and Distributions, 13, 165–175. 

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approach for predicting the distribution 

of rare and endangered species from occurrence 

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Edited by Richard Pearson 


105

opinion and perspectives 

opinion 

Political erosion dismantles the conservation network existing 

in the Canary Islands 

José María Fernández‐Palacios and Lea de Nascimento 

ISSN 1948‐6596 

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: jmferpal@ull.es; 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 territory 

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, political 

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 transfer 

of the management of Canarian National Parks to the regional administration. 

Keywords: Biodiversity loss, endangered species, National Parks, natural protected areas, political corruption, 

scientific community, species delisting 

Recently the Canarian Parliament has approved a 

new version of the Canarian catalogue of protected 

species (see Box 1) that reduces substantially 

both the number of species included (from 

466 species in the 2001 list to 361 species in the 

2010 list) and the protection afforded (from 381 

threatened species to 160, and from 85 protected 

species to 18). These reductions have been widely 

criticized by environmental NGOs and the local 

scientific community 1 , mainly due to the absence 

of a rigorous scientific process in its development. 

Although certainly the first version of the catalogue 

could be improved, the main reasons behind 

the new revisions were not conservation issues 

but rather strictly political. The reasons may 

include, for instance, the development of large 

infrastructures, such as industrial harbours and 

golf courses, which until the revisions were forbidden 

due to their impacts on protected species included 

in the original version of the Canarian catalogue. 

Changes in the environmental legislation of 

the Canary Islands entail a serious threat to the 

nature of this region of biogeographical interest 

(Francisco‐Ortega et al., 2000; Juan et al., 2000; 

Fernández‐Palacios & Whittaker, 2008). Thus, we 

believe it is important to share our appraisal of 

the current situation with the international scientific 

community. 

Within the new revised catalogue a completely 

new criterion for protection has emerged 

“especies de interés para los ecosistemas canarios” 

(literally: “species of interest for Canarian ecosystems”), 

comprising 152 species (see Box 1). The 

phrase is poorly chosen. It is supposed to apply 

only to endangered species, consequently the frequent 

and abundant species which usually structure 

and dominate the ecosystems are explicitly 

not listed, leading to a curious paradox: the Canarian 

pine (Pinus canariensis) is not a species of 

interest for the Canarian pine forest, the 

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, . 


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 ecosystems" 

which are those that, without being listed in the threatening situations above (endangered or vulnerable), 

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, control 

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 affecting 

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 

so on. This is not to say that the most common 

structuring species of the Canarian ecosystems 

have to be included in the catalogue, but we 

would like to draw attention to the inadequacy of 

the concept. 

But this conceptual shortcoming pales in 

comparison with the real repercussion of the new 

criterion, which is that those species listed here 

are only protected if present in an already designated 

Natural Protected Area (NPA). (In the Canaries, 

that means in either the Canarian Network 

of NPAs or the European Union Natura 2000 Network, 

which overlap extensively). If a listed species, 

for instance the woodcock (Scolopax rusticola) 

or the coot (Fulica atra) which are both included 

under the new criterion, dwells within the 

limits of the protected area they are safe; but if 

any birds cross those limits (which are not that 

obvious to birds, unfamiliar as they are with GIS), 

they can be shot legally by hunters. The same inconsistency 

affects, for instance, ca. 10 endemic 

species of sea lavenders (Limonium spp.) protected 

in certain ravines, but not in others. 

The new law could have negative implications 

for conservation biogeography, and this can 

be illustrated with some examples of the Canarian 

flora and fauna. The endemic legume Cicer canariensis, 

previously considered as vulnerable in 

the 2001 Canarian catalogue, is now included under 

the criterion species of interest. From its 12 

locations (ten in La Palma and two in Tenerife), 

the six populations in the North of La Palma 2 are 

outside NPAs and therefore unprotected according 

to the new law. Metapopulation dynamics in 

this species could be affected by this new criterion 

if source populations within these northern locations 

are threatened, endangering sink populations 

included in NPAs. The same could apply to 

the Abalone or Canarian clam (Haliotis tuberculata 

ssp. coccinea) or the Sea Horse (Hippocampus hippocampus). 

Both are marine species with sparse 

populations in the meso‐ and infra‐littoral, which 

do not always coincide with the geographical location 

of the marine Special Areas for Conservation, 

which occupy mainly leeward fringes on the Archipelago’s 

coasts. Collection and capture of both 

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. 


107

Canarian conservation network dismantled 

eries Law of the Canary Islands, but their inclusion 

in the new criterion may lead to confusion on the 

fishing ban in populations outside of the reserve 

networks. 

The case of the sea grass Cymodocea 

nodosa is of particular interest for two reasons; 

this species structures a community (“sebadales”), 

considered as Natural Habitat of Community Interest 

by the Habitats Directive, and its presence 

in the littoral zone is one of the main obstacles to 

the construction or enlargement of harbours. The 

most recent is the Puerto de Granadilla, where 

conservation of a European priority ecosystem 

comes into conflict with European funding of a 

large infrastructure. The sebadales are a key community 

from an ecological point of view as they 

play an important role in the carbon cycle, stabilize 

sandy soils, export biomass and act as a fish 

nursery area (Barberá et al. 2005). The latter characteristic 

is also very important for the sustainability 

of local fisheries. Also, the marine meadows of 

C. nodosa in the Canary Islands and Mauritania 

are the most extensive examples at the species’ 

southern limit and compromising them may therefore 

lead to range contraction. The construction of 

Puerto de Granadilla will severely damage one of 

the most genetically diverse patches of sebadales 

in the Archipelago (Alberto et al. 2008). In 2009, 

as a precautionary measure, the Superior Court of 

Justice of the Canary Islands suspended the proposal 

submitted by the Canarian Government, the 

Port Authority and the Canarian Company of Gas 

Transportation, to delist C. nodosa 3 . Currently, the 

European Courts have declared admissible the 

complaint filed by the NGO Ecologistas en Acción 

asking for the public release of documents that 

included alternatives to the construction of the 

harbour (including a renewal of the infrastructures 

of already existing harbours), that were hidden 

from the European Commission by Spain’s 

National Government. 

This controversial criterion — especies de 

interés para los ecosistemas canaries — is an adaptation 

of the criterion “species susceptible to 

habitat disturbance”, from the previous catalogue. 

In fact, many of the species of interest come from 

the former list of susceptible species or are downgraded 

threatened species. However in the former 

criterion there were no restrictions in the protection, 

such as the location or not in a NPA, and the 

main consideration to include a species was that 

its habitat was threatened, in regression, fragmented 

or limited. The previous criterion for protection 

was much more appropriate if we think 

about the design of the Canarian Network of 

NPAs. Unfortunately the Canarian Network was 

not based on a thorough analysis of metapopulation 

dynamics, genetic diversity or viability of 

populations, but simply in protecting less degraded 

remnants of communities that were still 

available. As in many parts of the world, reserves 

were not designed to meet the principles of systematic 

conservation planning needed to achieve 

representativeness and persistence of biodiversity 

(Margules and Pressey 2000). The situation further 

worsens in the Canaries when data, trends 

and viability of populations are almost unknown. 

The Canarian Network is largely protecting 

species from marginal populations. Moreover, the 

protection of species present only in the current 

Reserve Network inhibits re‐establishment of 

original distributions. A good example is the laurel 

forest in Anaga Rural Park, which nowadays is the 

best representation of this forest type in Tenerife 

yet still an impoverished fraction of its past distribution 

throughout the windward slope of the island. 

From the point of view of mitigating the effects 

of global change, vulnerability of certain species 

outside the Network would hinder altitudinal 

migration, especially when ecological corridors are 

not included in the design of NPAs. 

The practice of protecting taxa only in NPAs 

is already working in Catalonia (the only precedent 

in Spain). The Catalonian Plan of Areas of 

Natural Interest includes species of flora and 

fauna strictly protected in designated areas. To 

our knowledge no cases of the failure of these 

practices or public disapproval have been reported 

there, but we suspect that the species with 

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. 



work were not demoted from higher protection. 

In theory, the main aim of the existence of regional 

catalogues is ensuring the protection of 

particular species that are not considered by the 

National Catalogue. On the other hand several 

authors have questioned and analysed the effectiveness 

of NPAs Networks in biodiversity conservation 

(Jaffre et al. 1998, Rodrigues et al. 2004) 

and concluded that reserve networks are geographically 

and taxonomically unbalanced leaving 

a big proportion of endemic and threatened species 

unprotected. 

This way of thinking may function well 

when protecting a resource, for instance marine 

sanctuaries are intended to increase catch in 

neighbouring areas outside, and this works competently 

in the Canaries’ Marine Reserves with 

Fishery Interest, but is nonsensical when the aim 

of the declaration is to protect a threatened species. 

If a species is protected when within a NPA, 

but unprotected when beyond the area, what is 

really achieved in terms of protection Might it be 

too cynical to suggest the greatest achievement 

would be the political goal of inflating the number 

of species included in the catalogue thus reducing 

the number of critics of delisting Despite numerous 

public protests and the clear opposition of the 

majority of the Canarian scientific community, the 

new catalogue was presented by the leading political 

force in the Regional Parliament. These 

kinds of conflicts are not exclusive to the Canary 

Islands and are nowadays taking place in different 

regions of the world (Possingham et al. 2010, 

Metzger et al. 2011). 

If the delisting itself is not of sufficient concern, 

other news makes the outlook even bleaker. 

The Canaries harbour four of the 13 National 

Parks (NPs) in Spain – Cañadas del Teide 

(Tenerife), Caldera de Taburiente (La Palma), Timanfaya 

(Lanzarote) and Garajonay (La Gomera) 

– despite representing only 1.5% of the country’s 

geographical area. After decentralization of the 

Spanish State with the arrival of the democracy, 

the NPs were simultaneously co‐managed by the 

Central Government (Madrid) and the Regional 

Governments. However, the Spanish Constitutional 

Court now has determined that NPs management 

is exclusively a matter for the Regional 

Governments. Consequently the Central Government 

has transferred all management to the regions. 

In the case of the Canarian archipelago, this 

management was intended to be subsequently 

delegated to the respective island Councils 

(“Cabildos”) in 2012, although recently the new 

deputy of Environment of the Canarian Government 

expressed her intention to discuss again this 

transfer and to limit the management of the island 

Councils in the NPs. 

The transfer to regions is not inherently 

bad, and for instance would work exceptionally 

well in Northern European countries. The problem 

is not the law but how it is developed when the 

main political parties that govern in the Canary 

Islands show no interests in conservation, and an 

alarming number of its politicians, including some 

who have significant responsibilities in conservation, 

have been charged with environmental 

crimes 5 . Although some implications of decentralization 

should be positive, for instance the creation 

of regional lists and plans considering the particulars 

of each NP or the proximity to local specialists 

and technicians with a wider knowledge of the 

region, the result is exactly opposite. With the 

proximity of the management centres to the NPs, 

the likelihood of patronage and corruption seems 

likely to increase while unification of conservation 

criteria across the archipelago’s four NPs seems 

destined to decrease, especially if the different 

island Councils are governed by different political 

parties, which is currently the case. In addition, 

joint management of the NPs and the other NPAs 

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 August/2011); 

http://www.eldia.es/2011‐04‐13/CANARIAS/5‐Es‐frecuente‐alcaldes‐esten‐imputados‐delitosurbanisticos.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/islascanarias/noticia‐imputados‐canarias‐logran‐mantenerse‐instituciones‐20110524094822.html 

(last accessed August/2011) 

. 


109

Canarian conservation network dismantled 

dedicated to NPs. Considering that budgets are 

not fixed this would imply that funding to manage 

the NPs could eventually be used in other tasks, 

more consistent with the "needs of the moment". 

A recently created Commission of Canarian NPs, 

constituted mainly of politicians and with only two 

advocates for environmental issues, left aside the 

present directors and conservators of the NPs. It 

could also happen that once transferred to the 

Councils, the election of new directors will not 

consider the balance between conservation and 

management skills that such position requires. 

The island Councils are already in charge of 

the management of the Canarian Network of 

NPAs. While some of these areas have been actively 

managed others lack any type of control. 

The situation of similar NPAs varies among islands 

and for most the action plans have been partially 

or barely fulfilled, so that nowadays (more than 

ten years after its declaration) it is still easy to find 

dumps, illegal constructions, invasive species, together 

with other potential emerging threats. Despite 

the capacity and good work of environmental 

technicians, who struggle with budget cuts 

every year, the Councils have demonstrated a trajectory 

of inefficiency and lack of commitment to 

the management of NPAs. Within the new Canarian 

NPs framework, the rabbits will receive the 

responsibility of taking care of the lettuces. 


We would like to thank Rafael Loyola and three 

anonymous reviewers for their comments on the 

manuscript. We are also grateful to the editorial 

board of Frontiers in Biogeography for their help 

improving this paper. 

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catalogadas de Canarias: Cicer canariensis 

[Ciccan 06/2009]. Consejería de Medio Ambiente 

y Ordenación Territorial, Gobierno de Canarias, 

Las Palmas de Gran Canaria. Available at 

http://www.gobcan.es/cmayot/ 

medioambiente/medionatural/biodiversidad/ 

especies/especies_protegidas_amenazadas/ 

Edited by Joaquín Hortal & Michael N Dawson 

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ISSN 1948‐6596 

perspective 


The causes and biogeographical significance 

of species’ rediscovery 

Richard J. Ladle 1,2,* , Paul Jepson 2 , Ana C. M. Malhado 1 , 

Steve Jennings 3 and Maan Barua 2 

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 University 

of Alagoas, Praça Afrânio Jorge, s/n, Prado, Maceió, AL, Brazil, 57010‐020. 

e‐mail: richard.ladle@ouce.ox.ac.uk; 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 technology 

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 habitat. 

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 represented 

in the literature compared to rediscovered species that were only known from a handful of museum 

specimens. Thus, carefully distinguishing between the causes of temporal gaps in zoological records 

is essential for improving the utility of rediscovery data for biogeographical research and conservation 

practice. 

Keywords: extinction, range collapse, rarity, critically endangered, monitoring 

Introduction 

Rediscoveries of putatively extinct species are of 

great potential interest to both conservationists 

and biogeographers (Crowley 2011). For the former, 

‘rediscovery’ can be a considerable conservation 

policy and publicity asset (Ladle and Jepson 

2008, Ladle et al. 2009) – as testified by recent 

global initiatives: in 2009 BirdLife International 

launched a “global bid to try to confirm the continued 

existence of 47 species of bird that have 

not been seen for up to 184 years” (BirdLife International 

2009). The following year Conservation 

International launched its “Search for lost Frogs” 

which involves a dedicated campaign and expeditions 

to 18 countries seeking to locate 40 species 

not seen for a decade or more (Conservation International 

2010) – at the time of writing 12 species 

have been rediscovered. Moreover, since rediscovered 

species are typically exceedingly rare 

and geographically localized, new knowledge on 

population status and distribution supports effective 

conservation interventions. Finally, rediscoveries 

remove uncertainty from extinction risk assessments; 

a confirmed new record moves the 

species from ‘extinct’ or ‘probably extinct’ and 

into an IUCN threat (or data deficient) category. 

For biogeographers, species rediscovery has both 

a practical and conceptual significance. From the 


111

ediscoveries in biogeography 

practical perspective, the rediscovery of a species 

that has gone unrecorded for a long period of 

time improves geographical knowledge about 

some of the world’s rarest species, helping to address 

the Wallacean shortfall – the inadequacy of 

our knowledge of the geographical distributions of 

species (Lomolino et al. 2006, Riddle et al. 2011). 

The shortfall can often be extreme, with a species 

known from just one or a few museum specimens 

collected decades or even centuries earlier. These 

species are sometimes incorrectly assumed or declared 

extinct, a phenomenon which Ladle and 

Jepson (2008) refer to as a Wallacean extinction. 

As we discuss later, these extreme examples of 

the Wallacean shortfall are amongst the most frequently 

rediscovered species. 

More recently, biogeographers have started 

to use information on species rediscoveries to test 

theories of population decline and range collapse 

under anthropogenic disturbance (Fisher 2011a,b; 

Fisher and Blomberg 2011). The underlying idea is 

both simple and elegant: the location of a rediscovered 

species relative to its historical range reflects 

the pattern of range collapse. Thus, if anthropogenic 

pressures (e.g. unsustainable exploitation) 

are strongest at the periphery (Channel 

and Lomolino 2000) the rediscovery will most 

likely be made near the centre of the historic 

range. Diana Fisher’s (2011a) study of 67 species 

of rediscovered mammals found a number of clear 

trends, although these tended to be dependent 

upon the ecology of the species. For example, one 

of the strongest patterns observed was that rediscoveries 

were generally made at higher elevations 

than the original record (excluding mountain‐top 

and coastally restricted species). This provides 

some support for the hypothesis that higher elevations 

can sometimes provide ecological refugia 

(Towns and Daugherty 1994) and fits with the frequently 

observed pattern of habitat destruction 

and population extinction progressing from low to 

high altitudes (Triantis et al. 2010). 

However, like extinction, rediscovery is 

challenging to define. This should not be surprising 

since rediscovery and extinction are conceptually 

intertwined; extinction is the permanent absence 

of current and future records while rediscovery 

reflects the temporary absence of such 

records. Moreover, rediscovery is the proof required 

to refute a hypothesis of extinction. Given 

the close conceptual linkage between the concepts 

of rediscovery and extinction it is interesting 

that, until recently, there have been so few studies 

linking patterns of rediscovery to contemporary 

theories of population decline and extinction. 

One impediment to such research is the lack of a 

systematic approach to species rediscoveries that 

allow scientists to identify cases of rediscovery 

that have biogeographical or conservation significance, 

and which can be subject to meaningful 

analysis. Here, we propose a conceptual framework 

for understanding and analyzing species rediscovery, 

based on the social, institutional and 

ecological factors that created the temporal gap in 

occurrence data. We believe that formalizing the 

concept of rediscovery in this way has the potential 

to create new measures of the state of knowledge 

of the world’s rarest species, provide a quantifiable 

metric to support existing endangerment 

categorizations, and would help to maintain the 

culture of biogeographical exploration that contributes 

to the datasets that underpin global conservation 

target‐setting, advocacy and monitoring. 

Conceptual framework 

The ‘loss’ of a species and its subsequent rediscovery 

can be conceptualized as a result of the interplay 

among four socio‐ecological aspects of rediscovery 

(schematically illustrated in Figure 1): (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 technology 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 habitat. It should be noted that 

although these factors potentially apply to all 

‘lost’ taxa, owing to issues of historical data quality, 

funding and the culture of scientific exploration, 

rediscovery research has focused almost exclusively 

on herptiles, birds and mammals (cf. 

Scheffers et al. 2011). 


Richard J. Ladle et al. 

Knowledge of ‘lost’ species 

Enormous advances have been made over the last 

40 years in enumerating which species are apparently 

‘lost’. For example, BirdLife International has 

made significant investments in compiling new 

and authoritative assessments of threatened species 

using information from a variety of sources 

including amateur and university‐led research expeditions 

and major reviews of existing museum 

specimens. In particular, from the mid 1980s two 

major regional Red List reviews were compiled for 

the Americas (Collar et al. 1992) and Asia (Collar 

et al. 2001), the findings of which were then fed 

back to the BirdLife network of pioneering professional 

and amateur ornithologists (Tobias et al. 

2006, Butchart 2007). 

The knowledge of what is ‘lost’ is complicated, 

as rediscoveries can logically be split into 

four categories that reflect different degrees of 

uncertainty (and authority) about the continued 

existence of a target species (Table 1). An additional 

category could potentially be added to this 

typology to account for cases where an unrecorded 

sub‐species is elevated to full species 

status. For example, the Sangihe Shrike‐thrush 

(Colluricincla sanghirensis) was rediscovered in 

1985 but its status as a full species was only established 

in 1999 (Rozendaal and Lambert 1999). 

Changes in taxonomic status may have profound 

impacts on survey effort: according to Rasmussen 

et al. (2000), the demotion of the Sangihe Whiteeye 

(Zosterops nehrkorni) to sub‐specific status by 

Stresemann (1931) had the effect of making the 

species of “only marginal, regional interest” and 

as a consequence “for many years [it] received 

little attention” (p. 69). 

From the perspective of investigating range 

changes, confounding different categories of rediscovery 

could seriously influence research findings. 

For example, we might expect that all other 

things being equal, species whose habitat or range 

has not been surveyed for a significant period of 

time and for which there are no strong reasons to 

assume have become extinct (Table 1, category 4), 

are as likely to be rediscovered at the edge or centre 

of their historic range as are better‐known 

species. Moreover, all four categories of rediscovery 

may contain species that were only known 

from a small number of museum specimens – the 

rediscovery of which may tells us more about the 

history of biogeographical exploration than the 

ecology of decline and extinction. Indeed, Scheffers 

et al. (2011) found that the majority of recently 

claimed amphibian, bird and mammal rediscoveries 

represent first documentations since 

their original scientific description. It should also 

be noted that such rare species may have remained 

unrecorded because of intrinsic biological 

characteristics (e.g. nocturnal habits, cryptic 

colouration, etc.) rather than a lack of sampling 

effort and that these factors need to be carefully 

untangled in any analysis of patterns of rediscovery 

(see McCarthy 2008; Fisher and Blomberg 

2011). 

Figure 1. The four major dimensions 

of species rediscovery (see text). 


113


Type Rediscovery of… Example 

1. a species declared extinct by an authoritative 

source 

The Pohnpei Starling (Aplonis pelzelni) was declared 

extinct by the IUCN (1990) and rediscovered in 1995 

(Buden 1996) 

2. a species considered probably extinct by 

an authoritative source 

3. a species believed to be still extant but 

for which substantive searches over decades 

have drawn a blank. 

4. a species whose habitat or range had not 

been surveyed for a significant period of 

time, but for which there is no real reason 

to assume has become extinct 

The Sao Tome Grosbeak (Neospiza concolor) was 

described as probably extinct by Greenway (1967) 

and rediscovered in 1991 (Sergeant et al. 1992) 

According to the NGO BirdLife International the 

Madagascar Serpent Eagle (Eutriorchis astur) was 

not definitely recorded between 1930 and 1993 despite 

considerable search‐effort within its habitat. 

The Chestnut‐bellied Flowerpiercer (Diglossa gloriosissima) 

was unrecorded for 38 years: since 2003 it 

has been recorded from three locations (Tobias et 

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 rediscovery 

for conservation is where a previously well 

known species undergoes a population decline, is 

lost from biogeographical knowledge, and is then 

rediscovered. A possible example is the Australian 

Pygmy Blue‐tongue Lizard Tiliqua adelaidensis. 

This rather secretive lizard was relatively well 

known up to its disappearance in 1959; its rediscovery 

in 1992 (in the stomach of a snake) confirmed 

that the species now has “a dramatically 

reduced geographical range” (Milne and Bull 

2000, p. 296). The rediscovery of the Ivory‐billed 

Woodpecker (Campephilus principalis) (Fitzpatrick 

et al. 2005) would be an even better example, except 

that this rediscovery is increasingly looking 

like a case of mistaken identity (Dalton 2005, 

2010, Stokstad 2007). The apparent scarcity of 

such rediscoveries (cf. Scheffers et al. 2011) 

strongly suggests that a species that undergoes a 

well documented decline and disappearance is 

likely to be extinct. However, formally testing this 

hypothesis would require good information on 

population trends of rediscovered species prior to 

their original disappearance – data that rarely exist 

for older cases of species loss. 

A final aspect of the knowledge needed to 

find ‘lost’ species is the reliability of biogeographic 

information on where to search for the species. 

Thus, the Black‐hooded Antwren (Formicivora 

erythronotos) was known only from a 19 th Century 

type specimen, for which the type locality was 

probably incorrect, and which was also put in the 

wrong genus. Balchon (2007) suggests that this 

led to researchers “looking in the wrong place, for 

the wrong sort of bird and listening for inappropriate 

vocalizations”. Thus, ‘lost’ species can sometimes 

turn up thousands of kilometres away from 

where they were last seen, or in completely different 

habitats. For example, the Large‐billed Reed 

Warbler (Acrocephalus orinus) was previously 

known from just a single specimen collected in 

1867 in the Sutlej Valley, Himachal Pradesh, India. 

However, a living specimen was trapped in March 

2006 at Laem Phak Bia, Phatchaburi Province, 

south‐west Thailand, over 3000 km from the type 

locality (Round et al. 2007). The renewed interest 

in this species led to the unearthing of ten new 

museum specimens (Svensson et al. 2008) and, 

shortly afterwards, to the discovery of a breeding 

population in north‐east Afghanistan (Timmins et 

al. 2010). 

Institutional, scientific and technical capacity 

Even when a species is identified as possibly still 

extant, the institutional and technical capacity to 

find it may not exist. Such capacity, at a global 



level, has varied considerably over time and space 

in response to various cultural and ecological factors. 

Most notably, the mainstreaming of biodiversity 

into international development following the 

1992 Earth Summit created many new sources of 

funds and employment opportunities for scientists 

in less‐developed countries. With respect to birds, 

this increase in local capacity coincided with the 

creation of BirdLife International in 1993. BirdLife 

emerged from the International Council for Bird 

Preservation (founded in 1922) when its leaders 

devised the compelling proposition of forming an 

international partnership, under a single name, 

with smaller, national, bird‐orientated conservation 

organizations (Jepson and Ladle 2010). More 

generally, increased funding of expeditions by international 

NGOs has probably been the driving 

force behind the increasing frequency of rediscoveries 

of various taxa (Scheffers et al. 2011). 

Other trends within science and conservation 

also help determine the capacity and motivation 

that enables rediscoveries, especially the introduction 

of new technology. For example, advances 

in molecular biology have made it much 

easier to genetically compare preserved type 

specimens in museums with contemporary material 

collected directly or acquired from hunters or 

from rural markets. This has opened the way for 

completely new ways of rediscovering lost species, 

where a fragment of hair or a faecal sample 

may be sufficient to prove the continuing existence 

of a species that has still not been physically 

observed. 

An excellent example of such a technologyaided 

discovery is provided by Pitra et al. (2006), 

who recently announced the continuing existence 

of the giant sable antelope (Hippotragus niger 

variani), a sub‐species unique to Angola that was 

feared extinct after almost three decades of civil 

war. They compared the mitochondrial DNA sequences 

derived from old museum specimens 

with samples extracted from dung samples recently 

collected in the field. Such remotely collected 

DNA evidence can also be used to discount 

presumed discoveries or rediscoveries. For example, 

Hennache et al. (2003) used a range of techniques, 

including captive hybridization experiments 

and analysis of mitochondrial DNA and microsatellites, 

to conclusively demonstrate the hybrid 

origin of the imperial pheasant (Lophura imperialis). 

This mysterious bird had first been captured 

in 1924 when a single pair had been shipped 

to the private aviary of Jean Delacour in France 

and was not seen again until one was trapped in 

1990 (Hennache et al. 2003). 

It is not only advances in molecular biology 

that are facilitating rediscoveries. The ready availability 

of sophisticated audiovisual equipment has 

been especially important in the evolution of bird 

surveying. Two such technological advances, the 

increased availability of less expensive soundrecording 

and playback equipment in the late 

1990s and the more recent internet‐based birdsound 

archives, have dramatically increased the 

capacity of both amateurs and professionals to 

locate and identify rare and cryptic bird species. 

Moreover, advances in the quality of cameras and 

lenses, especially digital cameras and video recorders, 

have also been important in documenting 

and providing definitive proof of the existence of 

very rare species. For example, the New Zealand 

Storm Petrel (Pealeornis maoriana) was identified 

from the details on a digital image taken in 2003 

(Stephenson et al. 2008). It had previously been 

known only from putative fossil material, and 

from three specimens collected in the 19 th Century, 

150 years before its rediscovery. 

Accessibility 

Even if a species is extant and potential habitats 

have been located, the species may not be found. 

Access to suitable habitat may be limited because 

of political instability/restrictions, or simply the 

remoteness of potential sites. Although in the era 

of cheap international air travel this is arguably 

less important, it may have played a critical role in 

restricting the intensity of surveys and therefore 

the rate of rediscoveries in many parts of the 

globe. Examples of rediscoveries that were probably 

delayed, and possibly even caused, by political 

instability include that of the Large‐billed Reed 

Warbler in Afghanistan (see above) and the 

Gabela Helmet‐shrike (Prionops gabela), rediscovered 

in 2003 in Angola (Ryan et al 2004). 


115


A closely related factor is a lack of communication 

with remote and isolated rural communities 

who may already have knowledge of the continued 

existence of a putatively extinct species, or 

of a species new to science. Thus, a productive 

route to increasing rediscoveries (and new species 

discoveries) might be through better communication 

with remote tribes and communities whose 

knowledge of local biodiversity may extend considerably 

beyond that of conservationists. However, 

Fisher and Blomberg (2011) found that human 

population overlap did not predict rediscovery 

rate in mammals, possibly because expeditions 

and surveys may intentionally focus on more 

remote areas. 

Ecological factors 

The final aspect of rediscovery is the ecological 

characteristics of the putatively extinct species 

that may make verification of its continued existence 

problematic. For example, if the species is 

very rare and/or dispersed, then it may be difficult 

to locate an individual/population within an area 

of potentially suitable habitat. Even if the survey 

team is in the same area as the target species, it 

may still not be encountered because of phenotypic 

and ecological traits (e.g. cryptic coloration, 

lack of vocalizations, skulking behaviour, etc.) that 

reduce the probability of detection (Scheffers et 

al. 2011). However, the evidence for this effect is 

variable: Fisher and Blomberg (2011) found that in 

mammals many ecological characteristics such as 

cryptic coloration and arboreal and nocturnal behaviour 

were not significantly associated with rediscovery 

– although smaller rediscovered mammals 

had been missing for longer periods of time 

(Fisher 2011b). 

A possible example of ecology driving the 

lack of records is the Night Parrot, a species that is 

known from 23 specimens and many sightings of 

varying reliability from a wide geographic area of 

inland Australia (McDougall et al 2009). From 

what little information exists, the Night Parrot is 

crepuscular or nocturnal, cryptic, and when approached 

will only flush at close quarters, then fly 

low over short distances before plunging back into 

cover (Forshaw and Cooper 2002). Perhaps unsurprisingly, 

between 1912 and 1990 there were no 

records of the Night Parrot until one was hit by 

traffic (Boles et al. 1994). 

Rediscoveries reconsidered 

Given the very loose usage of the term 

‘rediscovery’ and the varying factors, social and 

ecological, that contribute to rediscoveries, both 

biogeography and conservation may benefit from 

adopting a stricter policy of usage. One strategy 

would be to strictly confine the term ‘rediscovery’ 

to species categorized as extinct in the IUCN system 

(Mace et al. 2008) or as ‘possibly extinct’, or 

‘lost’ by authoritative sources (Table 1, categories 

1, 2 and 3). It should be noted that many species 

that are considered possibly extinct are listed as 

“critically endangered” in the IUCN system. For 

example, Fisher (2011a) restricts her analysis to 

rediscovered mammal species that had been previously 

reported as globally extinct or possibly extinct. 

It should be noted, however, that this approach 

will not completely eliminate all the cases 

of species that are missing through low levels of 

surveying. 

An alternative strategy could be to classify 

rediscovery purely in terms of the length of time 

without a formal record. If this were adopted, the 

only issue would be an appropriate time frame for 

a given taxon. For example, De Roland et al. 

(2007) felt justified in claiming the ‘rediscovery’ of 

the Madagascar Pochard (Athya innotata) just 15 

years after the last confirmed sighting – conceivably 

the same individual. 

Using a simple time‐based criterion would 

provide a single, objective definition of rediscovery 

– whatever the cause of the gap in zoological 

records. Conservation bodies could potentially use 

this definition to periodically produce lists of species 

that may still be extant and, by extension, are 

in need of rediscovery. These could be categorized 

according to the time since a species was last recorded 

(e.g. 100 years ago, etc.). One advantage 

of such a system would be to maintain and 

extend the practice of biogeographical expeditions 

to remote areas. It would also help guard 

against the overuse or misrepresentation of redis‐ 



coveries in the media (Ladle et al. 2009). It would 

offer a viable alternative to the use of terms such 

as ‘possibly extinct’ (Butchart et al. 2006) and 

‘data deficient’, and would ensure better quality 

of data for future biogeographical studies. 

Conclusions 

The rediscovery of a species that was thought to 

be extinct can generate global interest and represents 

a real opportunity for conservationists to 

reassert core values and raise funds that may help 

protect poorly known habitats. Moreover, rediscoveries 

provide a unique source of information 

about the rarest and least‐known species (for certain 

taxa) that can be used to investigate biogeographic 

theories about range loss and extinction. 

Both of these important agendas would 

benefit from a greater systematization of the concept 

of rediscovery, acknowledging the varying 

causes (both social and ecological) of gaps in the 

temporal records of rare species. 

In summary, the study of rediscoveries provides 

a wonderful opportunity to assess both the 

subtle ecological and biogeogeographical characteristics 

of exceptionally rare species of well studied 

taxa such as amphibians, birds and mammals, 

and the fascinating historical and cultural trends in 

zoological surveying and exploration. Considerable 

efforts are being made to untangle these interacting 

factors (Fisher 2011a,b; Fisher and Blomberg 

2011, Scheffers et al. 2011), while the recent targeting 

of ‘lost species’ by international conservation 

NGOs is generating considerable amounts of 

valuable new data. Nevertheless, the lack of rediscovered 

species that were previously well known 

and which had undergone a well documented 

process of population decline, fragmentation and 

local extinction (Scheffers et al. 2011) remains a 

worrying trend for global conservation. 

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Edited by Jan Beck 


membership corner 

ISSN 1948‐6596 

from the society 

Getting to know IBS Early Career Members 

The International Biogeography Society (IBS), 

founded just 10 years ago, is fast growing both in 

terms of members and activities offered (Field and 

Heaney 2011). Students and early‐career biogeographers 

are also becoming increasingly involved 

within the IBS. From 2002 to 2010, the proportion 

of new members who are students joining 

the IBS each year has increased from 23% to 48%. 

Currently, student members comprise 35% of 

IBS’s 740 members. The IBS, aware of the rising 

importance of these younger members, has been 

trying to increase the benefits available for them. 

In addition to the student travel grants, poster 

awards and discussion groups held at the IBS 

meetings, the IBS is trying to foster interaction 

among students and postdocs, which recently culminated 

in the first IBS Early Career conference 

that was held at Oxford University from 23 to 25 

September 2011 (http://www.biogeography.org/ 

html/Meetings/index.html). 

With the intention of getting to know its 

early‐career members (herein ECM) and learning 

their opinions on the services provided by the IBS 

and on how these can be improved, the IBS invited 

ECM to participate in a survey that was held 

in June 2011. Of the 48 ECM that completed this 

survey, 11% were Junior Postdocs, 75% were PhD 

students, 8% were Masters students, and 6% were 

undergraduate students. Around 17% were aged 

between 20‐25 years, 49% were 26‐30 years, 23% 

were 31‐35 years, and 11% were more than 35 

years young; 56% were female and 44% were 

male. Although most ECM are currently affiliated 

either with North American or European institutions 

(50% and 33% respectively; total of 42 answers), 

they represent a total of 24 nationalities; 

26% are from North America, 17% from Central 

and South America, 15% from Northern Europe, 

28% from Southern Europe, and the other 12% 

from Australia/New Zealand, the Middle East, Africa 

and Asia. ECM work on a very broad range of 

topics, from species distribution patterns (the 

most mentioned topic), to evolutionary biogeography, 

dispersal and colonization, biogeography of 

species’ traits, island biogeography, phylogeography, 

global change biology, marine biogeography, 

or paleobiogeography, among others. Their broad 

interests are also reflected in the fact that most 

ECM are also affiliated with societies focusing on 

diverse topics, including ecology, evolution, conservation, 

paleontology, geography, botany, mammalogy, 

entomology, etc. These are indeed very 

encouraging results that show the IBS is reaching 

young researchers from a wide variety of research 

topics and geographic locations. 

In general terms, the IBS is meeting ECM 

needs (25% responded that the IBS is doing this 

“very well”, 60% “fairly well”). However, there is 

room for improvement (15% responded “not very 

well”), and several suggestions were made; responses 

to open‐ended questions emphasized the 

need for more off‐year meetings (regional meetings, 

workshops, etc.), more jobs/grant announcements, 

more travel grants, online teaching resources, 

more talks at the IBS meetings by 

younger researchers and more opportunities to 

meet other researchers. The IBS is already working 

towards improving the services it provides to 

all its members, and new actions are being made 

to adopt suggestions. 

The first action was to support the IBS Early 

Career conference (for students and biogeographers 

who have finished their PhDs in the past five 

years). Almost ninety young researchers participated 

and had the chance to present their work, 

and to interact with each other and with the IBS 

board members. This conference was organized 

into ten different sessions that covered several 

aspects of macroecology, island biogeography, 

phylogeography, paleobiogeography, evolutionary 

biogeography and conservation biogeography. 

Second, we are also working towards increasing 

regular communication among IBS members. 

One way of doing this is through online social 

networks, such as Facebook, and other webbased 

platforms (e.g. the IBS blog; http:// 

biogeography.blogspot.com/). Currently, the IBS 

has a Facebook group with ~590 members, where 


119


anyone can post announcements, share ideas and 

publications of general interest, start discussions 

and interact with other members. Most ECM are 

in fact Facebook users (80%; only 7% are Twitter 

users), but only 8% of these members read the IBS 

Facebook page on a weekly basis, and 44% actually 

never read it (31% read it once per month, 

and 17% every 3‐6 months). Regarding the IBS 

blog, again only a small number of people read it 

on a weekly basis (6%), with most people reading 

it once per month (38%; 31% read it every 3‐6 

months and 25% never read it). Another platform 

the IBS has for communicating with its members, 

and to foster communication between its members, 

is the online journal Frontiers of Biogeography 

(http://www.biogeography.org/html/fb.html). 

This journal has a section especially devoted for 

this purpose – the membership corner – of which 

most ECM were not aware (66%). Thirty‐six percent 

of ECM read every issue, while 31% read 2‐3 

issues per year (27% read it rarely and only 6% 

never read it). Main sections of interest to the 

ECM are (i) mini‐reviews on a particular taxon, 

biogeographic topic, or question, (ii) thesis abstracts, 

and (iii) symposium/congress summaries. 

In fact, 88% showed interest in submitting a 

manuscript to any of these sections. 

One of the most important activities organized 

by the IBS is the biennial meeting. The next 

one will be held at Florida International University 

in Miami, Florida, in January 2013 (http://www. 

biogeography.org/html/Meetings/2013). Most 

ECM are planning to attend this meeting (79%) 

and would prefer to give a talk (51%; 23% prefer a 

poster presentation and 26% have no particular 

preference). One of IBS’ concerns is to maximize 

compatibility between high quality talks and fair 

representation of researchers from different 

countries, gender, and career stages. There was 

almost an even split among ECM on favoring a 

similar number of talks by established and 

younger researchers, and having more talks by 

senior researchers plus some younger ones (40% 

and 43%, respectively; 11% would prefer to have 

mainly senior researchers and 6% showed no preference). 

There was no overwhelming support for 

student‐only sessions in future meetings (55% 

found it important), but most respondents 

showed some willingness to extend their stay in 

order to attend this type of event (83%). In the 

previous meetings, students (particularly those 

who have been awarded with a student travel 

grant) have been invited to attend discussion 

groups, where senior biogeographers lead the 

discussion on several subjects, from career and 

publishing advice to specific research topics. 

Those who have attended these student discussion 

groups in past meetings (41%) found them 

helpful (63%). Suggestions for discussion topics in 

future meetings, other than those already covered 

in these discussion groups, included advanced 

analysis in biogeography and partnerships and 

international activities among researchers. There 

was some support for future off‐year meetings 

(33% found it useful; 61% said it was somewhat 

useful, and over 90% said they would at least try 

to attend), especially if these are focused on specific 

research topics and methodologies (31% and 

29%, respectively; there was a tie between meetings 

on specific geographic realms and on a broad 

scope within biogeography – 20% each). Some 

respondents also called for workshops and seminars, 

online courses, cross‐society ventures to 

boost interaction between similarly oriented academics 

and excursions into biogeographically interesting 

regions covering a broad range of taxa. 

There was also a significant interest in having a 

showcase at the next IBS meeting of funding agencies 

from different countries (70%), with most respondents 

being willing to provide information on 

this matter (55%). 

The long‐term success of any growing society 

depends on the involvement and interest of its 

youngest members. We’re fortunate that many 

ECM have shown willingness to get involved in 

promoting communication between IBS members, 

Did you know that any member of the IBS may raise an issue or appeal a decision of the governing 

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/). 



to help organizing off‐year activities, and to submit 

manuscripts to Frontiers of Biogeography. The 

IBS wants to hear and share more of the early career 

members’ opinions and ideas; this article is 

intended as both thanks and encouragement for 

your active involvement, especially in the readily 

accessible platforms such as Frontiers of Biogeography 

and Facebook. Finally, we would like to 

thank all the members who participated in this 

survey, and particularly those who have shown 

interest in devoting some of their time to the society. 

We look forward to working with and for you 

in the coming years. 

Ana M. C. Santos 

IBS Student‐at‐Large; Departamento de Ecologia, 

Instituto de Ciências Biológicas, Universidade Federal 

de Goiás, Brazil. 

e‐mail: ana.margarida.c.santos@googlemail.com 

References 

Field, R. & Heaney, L.R. (2011) Looking to the future of 

the IBS: the 2011 IBS membership survey. Frontiers 

of Biogeography, 3, 71‐73. 

Edited by Matthew Heard 

from the society 

Call for proposals for hosting 7th Biennial Conference of the IBS 

We are seeking proposals for hosting the 7th biennial 

conference of the International Biogeography 

Society to be held in early January 2015. Proposals 

should be submitted by individuals who are interested 

in chairing the local (host) committee. The 

duties of the local host include conducting contract 

negotiations with the venue and the hotel as 

well as all local logistics including field trip organization 

and production of the abstract book. 

Minimum requirements of the venue are 1) 

one auditorium with a capacity of 450‐550 people 

(2 days), 2) three or four smaller rooms with a capacity 

of 75‐150 people (1 day), and 3) various 

smaller meeting rooms. The IBS is interested in 

holding the biennial conference in locations fairly 

convenient with respect to the majority of its 

membership base in North America and Europe. 

Locations of past (and upcoming) conferences 

can be seen here: http://www.biogeography.org/ 

html/Meetings/index.html. 

Please include the following information in 

the proposal: 

1. Location of the meeting (city) and the host institution 

or organization. 

2. What would be the benefit of hosting the conference 

at this location 

3. Actual site of the meeting and the capacity of 

the auditorium. 

4. Space for poster sessions‐‐general size and location 

relative to the auditorium. 

5. Approximate cost for three‐day use of the venue. 

A specific quote is not needed, but evidence 

of the price competitiveness is crucial. 

6. Transportation infrastructure, including travel 

from airport. 

7. Attractions in the vicinity of the conference 

site, including field trip potential. 

8. Who would potentially serve on the local organizing 

committee 

Proposals from prospective hosts of the 

biennial conference must be received before 20 

January 2012. Please send proposals by email to 

Daniel Gavin, IBS Vice‐President for Conferences 

at dgavin@uoregon.edu. 

Dan Gavin 

IBS Vice‐President for Conferences; 

IBS Student‐at‐Large; Department of Geography, 

University of Oregon, USA. 

e‐mail: dgavin@uoregon.edu 

If you want to announce a meeting, event or job offer that could be of interest for (some) biogeographers, 

or you want to make a call for manuscripts or talks, please contact us at 



121


Job announcements 

Three Professorships and One Tenure‐Track 

Lectureship 

University of California, Merced, USA 

The School of Natural Sciences at the University of 

California, Merced seeks applicants for four faculty 

positions: Ecology (Full or Associate with tenure, 

or Assistant tenure‐track), Systems Biology 

(Assistant tenure‐track), and Biostatistics 

(Assistant tenure‐track), and one tenure‐track Biology 

Lecturer. For the Ecology position, we seek 

outstanding individuals with research interests in 

any ecological field using experimental, field, computational, 

and/or theoretical approaches and 

working at population to global scales. The Systems 

Biology position includes research areas that 

use comprehensive datasets and multiple types of 

analysis to relate overall biological function to underlying 

biochemical or biophysical processes for 

predictive understanding. The Biostatistics research 

areas of interest include statistical methods 

for experimental design, epidemiology, medical 

informatics, evolutionary biology, sequence bioinformatics, 

genomics, evolution of microbial systems 

and pathogens, and systems biology. The 

Lecturer position closely parallels a tenure‐track 

Assistant Professor but with an emphasis on undergraduate 

education. All applicants must be 

able to teach effectively at both undergraduate 

and graduate levels. For more information and to 

apply go to: http://jobs.ucmerced.edu/n/ 

academic/listings.jsf;jsessionid=95FADBAFFF4C13 

F912A3B023DA4F1F80seriesId=1 

Interested applicants should submit materials 

online. Applications will be considered starting 

05 December 2011 (Biostatistics, Systems Biology 

professorships), or 16 December 2011 

(Ecology professorship and Biology Lecturer). UC 

Merced is an AA/EOP employer. 

upcoming events 

VIPCA Molecular Ecology 

4–7 February 2012 – Vienna, Austria 

http://www.vipca.at/MOLECOL/ 

Annual Conference of the Society for Tropical 

Ecology (gtö) 

Islands in land‐ and seascape: The challenges of fragmentation 

22–25 February 2012 – Erlangen, Germany 

http://www.gtoe‐conference.de/ 

6th Annual Meeting of the Specialist Group 

on Macroecology of the Ecological Society of 

Germany, Austria and Switzerland (GfÖ) 

29 February – 2 March 2012 – Frankfurt, Germany 

http://www.bik‐f.de/ 

21 st Workshop of the European Vegetation 

Survey (EVS) 

24–27 May 2012 – Vienna, Austria 

http://evs2012.vinca.at/ 

VertNet biodiversity informatics training 

workshop 

24–30 June 2012 – Boulder, USA 

http://vertnet.org/about/BITW.php 

97th ESA Annual Meeting 

Life on Earth: Preserving, Utilizing, and Sustaining our 

Ecosystems 

5–10 August 2012 – Portland, USA 

http://esa.org/meetings/ 

3 rd European Congress of Conservation Biology 

Conservation on the edge 

28 August – 1 September 2012 – Glasgow, UK 

http://www.eccb2012.org/ 

6th International Conference of the IBS 

January 2013 – Florida, USA 

http://www.biogeography.org/ 


table of contents 


the scientific magazine of the International Biogeography Society 

volume 3, issue 3 ‐ November 2011 


ISSN 1948‐6596 

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. 

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 invertebrates, 

by R.M. Chefaoui 


opinion: Political erosion dismantles the conservation network existing in the Canary Islands, by J.M. Fernández‐Palacios 

& L. de Nascimento 

perspective: The causes and biogeographical significance of species’ rediscovery, by R.J. Ladle et al. 111 


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 

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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 

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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 

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We gratefully acknowledge Evolutionary Ecology, Ltd. and Mike Rosenzweig in particular for the advice on copyright matters. 

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