frontiers of biogeography

vol. 3, nº 3 ‐ november 2011

the scientific magazine of the International Biogeography Society

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

frontiers of biogeography the scientific magazine of the International Biogeography Society volume 3, issue 3 ‐ November 2011

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.

editorial board

editor‐in‐chief: Joaquín Hortal – Museo Nacional de Ciencias Naturales (CSIC), Spain and Universidade Federal de Goiás, Brazil

deputy editors‐in‐chief: Michael N Dawson – University of California, Merced, USA Richard Field – University of Nottingham, UK

frontiers of biogeography is published by the International Biogeography Society (IBS), an international and interdisciplinary society contributing to the advancement of all studies of the geography of nature

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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

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ISSN 1948‐6596

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

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

International Biogeography Society officers 2011‐2012

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 = cAz 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 continen‐

tal 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) re‐

cently came to the conclusion that the power law

fits empirical data best in most cases (see also

Dengler & Odeland 2010). Various authors ob‐

served 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 biodiver‐

sity, 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 extinc‐

tion of a species is challenging and estimating ex‐

tinction 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 extinc‐

tion have been welcome despite critical studies

that often found lower extinction rates than pre‐

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

reasonably, that on top of imminent extinction in

some species, others will be doomed to future

extinction because of reductions in their popula‐

tion size, and that this ‘extinction debt’ explains

apparent misfits. Other sources of uncertainty of

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

sumption of a completely inhospitable matrix be‐

tween 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 con‐

cept 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 indi‐

vidual. To show this, they formulated both as spa‐

tially explicit sampling processes (SAR for first en‐

counters, EAR for last encounters). They con‐

cluded that SAR‐derived estimates of imminent

extinction will always be too high, unless individu‐

als 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 aggre‐

gation is modelled, which offers an alternative

(but a more challenging one) for estimating imme‐

diate extinction of endemics from area loss.

He & Hubbell (2011) clearly acknowledged

that there is an anthropogenic extinction crisis

and that habitat loss causes extinction. Further‐

more, they did not claim that small population

sizes of remaining species could not lead to fur‐

ther, lagged extinction (in He & 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

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

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 previ‐

ously asserted (fossil fuel lobbying in the climate

change discussion is cited as example of this tac‐

tic). Conserving nature is not only about science,

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 re‐

gions, which is probably appropriate for capturing

immediate extinction in many conservation set‐

tings 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 system‐

atically between SAR and EAR (which is presented

confusingly) – are generalities. Thus it remains to

be seen whether SARs always overestimate ex‐

tinction, as He and Hubbell (2011) claimed. A fur‐

ther task will be to quantitatively estimate how

many more species may go extinct after a time

lag: how large the extinction debt really is (see

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 under‐

stood solely on the basis of island biogeographic

mechanisms (Rosenzweig 2001) – that is, species

richness as equilibrium between immigration +

speciation and extinction. The spatial and tempo‐

ral scales of analysis, among other factors, may be

relevant for this. Under such circumstances, SARs

may estimate the new equilibrium state, account‐

ing 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 Ecol‐ogy, 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 assess‐ment. 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) Extinc‐tions: conserve not collate. Nature, 474, 284.

Harte, J., Kinzig, A. & Green, J. (1999) Self‐similarity in the distribution and abundance of species. Sci‐ence, 284, 334–336.

He, F. & Hubbell, S.P. (2011) Species–area relationships always overestimate extinction rates from habi‐tat loss. Nature, 473, 368–371.

Kinzig, A. & Harte, J. (2000) Implications of endemics–area relationships for estimates of species ex‐tinctions. Ecology, 81, 3305–3311.

Koh, L.P. & Ghazoul, J. (2010) A matrix‐calibrated spe‐cies–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 biogeographi‐cal 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 relation‐ships. Nature, in press.

Plotkin, J.B., Potts, M.D., Yu, D.W., et al. (2000) Predict‐ing species diversity in tropical forests. Proceed‐ings of the National Academy of Sciences USA, 97, 10850–10854.

news and update

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

Preston, F.W. (1962) The canonical distribution of com‐monness 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

news and update

update

Extinct or extant? Woodpeckers and rhinoceros

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

ISSN 1948‐6596

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 Wal‐

lacean shortfall. Fortunately, new tools are being

developed that allow conservationists and bio‐

geographers to determine the existence of extant

populations with much greater accuracy.

Foremost among these new tools is the in‐

creasing use of genetic analysis. This was recently

used to great effect to confirm the extinction of

the Javan rhinoceros (Rhinoceros sondaicus anna‐

miticus) in Cat Tien National Park in Vietnam

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

Javan rhinoceros are remarkably shy forest‐

dwelling animals that are difficult to see under

natural conditions and were only rediscovered in

mainland Asia in 1988. Given the difficulty of tra‐

ditional 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 be‐

tween 2009 and 2010. The analysis indicated that

all the dung belonged to a single individual, the

body of which was found April 2010, thereby con‐

firming 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 sight‐

ing, the reliability of which is often highly ques‐

tionable. Andrew Solow and his colleagues have

recently come up with an ingenious method to

account for this inevitable uncertainty (Solow et

al. 2011). They use Bayesian (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 classi‐

fied 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 R e p o r t . A v a i l a b l e a t : h t t p : / /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

Edited by Joaquín Hortal

news and update

Links between climate and societal instability,

conflict and war have increasingly been suggested

and analyzed (Diamond 2005), thereby fusing tra‐

ditionally 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), ana‐

lyzed 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 predic‐

tions 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 be‐

tween past climatic changes and historical con‐

flicts. Two new studies have attempted this.

Hsiang et al. (2011) made use of the recur‐

ring 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 con‐

flicts were more likely to arise during El Niño

events as compared to La Niña periods. Further‐

more, 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 agricul‐

tural productivity and/or economic disturbance

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

and diseases) seem plausible, but psychological

effects of unusual weather conditions on a large

number of individuals may also increase a soci‐

ety’s conflict potential.

Zhang et al. (2011) presented a detailed

causality analysis based on a time series of cli‐

matic fluctuations over a 300 year period in pre‐

industrial Europe. They provide strong support for

the idea that climatic variation caused fluctuations

in agricultural productivity, and hence food avail‐

ability 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 peri‐

ods of crisis and harmony for past eras with less‐

detailed 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 pat‐

terns. 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 above‐

mentioned disciplines to sort out the ultimate

causes of why certain regions and/or societies

navigated smoother and less violent routes

through times of crisis than others (my current

location, Switzerland, is a prime example within

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

ISSN 1948‐6596

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update

Climate wars

84 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

news and update

References

Beck J., & Sieber, A. (2010) Is the spatial distribution of mankind’s most basic economic traits deter‐mined by climate and soil alone? PLoS ONE 5(5): e10416.

Burke, M., Miguel, E., Satyanath, S., Dykema, J. & Lo‐bell, D. (2009) Warming increases risk of civil war in Africa. Proceedings of the National Acad‐emy 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 con‐flicts are associated with the global climate. Na‐ture, 476, 438–411.

Samson, J., Berteaux, D., McGill, B.J., Humphries, M.M. (2011) Geographic disparities and moral hazards in the predicted impacts of climate change on human populations. Global Ecology and Bio‐geography, 20, 532–544.

Zhang, D.D., Lee, H.F., Wang, C., Lie, B., Pei, Q., Zhang, J. & An, Y. (2011) The causality analysis of cli‐mate change and large‐scale human crisis. Pro‐ceedings 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 popula‐tion decline in recent human history. Proceed‐ings 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 activi‐

ties have affected patterns of biological diversity

from a variety of perspectives, with special atten‐

tion often given to oceanic islands. With the cur‐

rent accelerating pace of environmental change,

these effects are increasingly evident at global

scales. Human industry, commerce, agriculture

and transportation all have the potential now to

affect natural systems globally through an assort‐

ment of drivers; primary among these are land‐

use change, species introductions and climate

change.

Human activities and their consequences

come to a unique focus in urban areas, an expand‐

ing 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 hu‐

man‐dominated environments offer unique op‐

portunities to investigate the broad‐scale dynam‐

ics 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 ex‐

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

Urban areas typically contain spatially heteroge‐

neous 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 ap‐

proaches are currently used to investigate urban

systems at broad spatial scales: urban plots or

transects, the entire urban matrix and the urban

matrix embedded within a regional context

(Werner 2011). Each sampling approach provides

a unique inferential basis, although the third al‐

lows for more refined interpretation, controlling

for regional differences.

A recent study in Global Ecology and Bio‐

geography adopts a novel perspective and exam‐

ines how avian assemblages sampled within plots

of intact vegetation in urban and semi‐natural ar‐

eas differ based on several common mac‐

roecological relationships. Pautasso et al. (2011)

compiled data on species composition and abun‐

dance from all around the globe, although the

majority of the samples are from Europe and

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

ISSN 1948‐6596

news and update

tionships between urban and semi‐natural locali‐

ties. The number of exotic bird species in urban

areas is low, suggesting that these relationships

are defined primarily by native species in both

environments. These findings highlight the impor‐

tance of maintaining intact vegetation within ur‐

ban landscapes and the role of urban diversity as a

tool for promoting conservation initiatives and

biological awareness, as emphasized in many ur‐

ban‐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 preva‐

lence and quality of global data sources provides

an exciting basis to examine the structure and de‐

terminants of these macroecological relationships

across more refined temporal, spatial and anthro‐

pogenic gradients.

By taking a global perspective, novel in‐

sights can be gained on the unique position urban

areas have, both as a source for global change and

as regions capable of maintaining important as‐

pects of biological diversity. Global comparative

studies also have the potential to bolster and re‐

fine current recommendations about how to

maintain biological diversity within human‐

dominated landscapes. Specifically, the preserva‐

tion 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 major‐

ity of the research has been conducted, allowing

for a more inclusive set of inferences and recom‐

mendations. Urban data are becoming increas‐

ingly available through remote sensing activities,

citizen science initiatives and broader collabora‐

tive efforts. Exploring how anthropogenic activi‐

ties are impacting natural systems globally is criti‐

cal in supporting a truly comprehensive under‐

standing 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 Ecol‐ogy 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) Com‐positional similarity among urban floras within and across continents: biogeographical conse‐quences of human‐mediated biotic interchange. Global Change Biology, 13, 913–921.

La Sorte, F.A., McKinney, M.L., Pyšek, P., Klotz, S., Rap‐son, G.L., Celesti‐Grapow, L. & Thompson, K. (2008) Distance decay in similarity among Euro‐pean urban floras: the impacts of anthropogenic activities on β diversity. Global Ecology and Bio‐geography, 17, 363–371.

McKinney, M.L. (2008) Effects of urbanization on spe‐cies 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 in‐ternational 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) Phyloecol‐ogy 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.

Edited by Joaquín Hortal

86 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

ISSN 1948‐6596 news and update

Community ecology traditionally focuses on hypo‐

thetical‐deductive and experimental approaches

and often is criticized for narrowing our under‐

standing of nature to local idiosyncrasies, ignoring

the importance of historical explanations. On the

other hand, approaches taken by macroecologists

and biogeographers have been excessively ex‐

ploratory 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 ap‐

proaches can learn from each other is pivotal in

the challenge of integrating data from different

scales in order to unravel the ecological and evo‐

lutionary mechanisms that influence current pat‐

terns in biodiversity and ecosystem functioning.

Species richness has been the most com‐

mon metric used to represent all aspects of bio‐

logical diversity (from genetic and taxonomic to

phenetic diversity). However, species richness

alone cannot describe the processes involved in

species coexistence and ecosystem functioning

and also does not describe properly the differ‐

ences in community structure. In contrast, phy‐

logenetic and functional diversities allow us to

understand the relative importance of species

composition in terms of evolutionary history and

ecological similarities. Phylogenetic diversity (PD)

is a biodiversity measure that accounts for the

phylogenetic relationship (hence evolutionary his‐

tory) among species, whereas functional diversity

(FD) represents how species are distributed in a

multidimensional niche space defined by ecologi‐

cal traits.

Phylogenetic and functional approaches to

community ecology emerged as prominent fields

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

how independently and without much crossover

in the first years. Early PD measures were pro‐

posed as a tool to select conservation areas, but

later the idea was extended to understand how

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 investigat‐

ing assembly rules (see Pavoine & Bonsall 2011).

How could macroecology and biogeography bene‐

fit from these two approaches? The answer lies in

understanding what FD and PD should represent

and how they relate to each other: while phyloge‐

netic 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 phylogeogra‐

phy. Therefore, combining ecological and phyloge‐

netic 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 decom‐

posed 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 be‐

cause biogeographers and macroecologists were

more aware of evolutionary and historical hy‐

potheses, 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, ap‐

proaches (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 predic‐

tions. Safi et al. (2011) investigated global pat‐

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

update

Beyond taxonomical space: large‐scale ecology meets func‐tional and phylogenetic diversity

news and update

terns of mammal PD and FD and found that when

controlling mammal assemblages for their evolu‐

tionary 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 competi‐

tion 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 ex‐

ample, 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 heteroge‐

neity could also result in an overdispersion in FD

because coexisting species could adapt and spe‐

cialize to the different environmental conditions.

Some light has been shed on beta‐PD pat‐

terns by Gómez et al. (2010), studying Neotropical

Forest antbirds at different spatial scales. If speci‐

ation 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 ex‐

pect high species turnover (taxonomic beta diver‐

sity) 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

88 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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

1975 1980 1985 1990 1995 2000 2005 20100

20

40

60

80

100

120

140

160

180

200

2007 2008 2009 2010 20110

2

4

6

FD

PD

PD

FD

Any spatial scale

Large spatial scale

beta-PD

beta-FD

beta-PD

year

publ

ishe

d st

udie

s

news and update

region would contain distinct clades, with inde‐

pendent diversifications. Finally, if observed val‐

ues 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 spe‐

cies are sorted from the regional pool according to

their traits, we expect both species and functional

turnover. However, if the species pool is com‐

posed of ecologically similar species – an indica‐

tion 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, spe‐

cies turnover should occur independently of func‐

tional turnover (Mouchet et al. 2010). Neverthe‐

less, species traits should have – at least to some

extent – some phylogenetic signal and, therefore,

partitioning the relative contribution of evolution‐

ary history to trait dissimilarities among species

may be important. A potential, and unexplored,

solution is to decouple functional diversity into

“phylogenetic structured” and “specific

(ecological)” components. This would help us to

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 func‐

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

community ecology with evolutionary biol‐

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

some methodological challenges – how to prop‐

erly define the species pool and null models,

which traits should be used, what is the most suit‐

able measure of PD and FD, and so on (see

Pavoine & Bonsall 2011), but we should avoid be‐

coming 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 ex‐

penses 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 ap‐

plied to taxonomic diversity can be extended to

phylogenetic and functional diversity (Meynard et

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

ate more rigorous and direct predictions for most

of the hypotheses in macroecology and biogeog‐

raphy, such as attempts to explain latitudinal pat‐

terns of biodiversity (Willig et al. 2003). These

metrics also present an opportunity to formulate

new hypotheses about how species evolutionary

history and trait diversity are distributed across

communities at different scales. For example,

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 para‐

digm that communities are ‘saturated’. Large‐

scale phylogenies and trait databases are cur‐

rently becoming available for a wide range of

taxonomic groups, facilitating estimates of FD and

PD. Including these two aspects of biological di‐

versity will be crucial if we want to advance from

exploratory studies which report interesting rela‐

tionships 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 manu‐

script. This work was supported by CAPES (project

#012/09).

Marcus V. Cianciaruso Departamento de Ecologia, Instituto de Ciências Bioló‐

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

news and update

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 habi‐tat filtering in community assembly of Neotropi‐cal 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 be‐tween 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 Evolu‐tion, 21, 178–185.

Meynard, C.N., Devictor, V., Mouillot, D., Thuiller, W., Jiguet, F. & Mouquet, N. (2011) Beyond taxo‐nomic diversity patterns: how do α, β and γ components of bird functional and phylogenetic diversity respond to environmental gradients across France? Global Ecology and Biogeogra‐phy, 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. Func‐tional Ecology, 24, 867–876.

Pavoine, S. & Bonsall, M. (2011) Measuring biodiversity to explain community assembly: a unified ap‐proach. 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., Ar‐mour‐Marshall, K. & Diniz‐Filho, J.A.F. (2011) Understanding global patterns of mammalian functional and phylogenetic diversity. Philoso‐phical Transactions of the Royal Society of Lon‐don 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) Phyloge‐netic 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

90 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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

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

The World atlas of mangroves, an update to Spal‐

ding 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 ta‐

bles provide captivating facts about the ecological

and economic values of mangroves and the conse‐

quences 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 suitabil‐

ity of intact mangroves for protecting coastal re‐

gions against tsunamis (Wibisono and Suryadipu‐

tra 2006). This will arm (with powerful arguments)

ecologists, conservation biologists and policy‐

makers, who urgently need to communicate this

knowledge in order to increase public awareness

and political willingness to protect and rehabili‐

tate one of the most vulnerable ecological sys‐

tems 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, ac‐

companied by information about their specific

threats, level of degradation and extent of reha‐

bilitation programs guides the reader through a

multitude of distinct features, while keeping simi‐

larities 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 compen‐

dium and make this atlas perfect as a quickstart

guide for students as well as experienced re‐

searchers approaching a new region.

Considering the presentation of global

trends as the main purpose of the World Atlas Of

Mangroves, this book fulfils expectations. Unnec‐

essary 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 con‐

tent of the publications loosely mentioned at the

end of each subchapter, nor established knowl‐

edge 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 over‐

wash mangroves is needlessly incomplete; it could

be easily improved by following standard man‐

grove 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 capabil‐

ity for mangrove formation, but then present ele‐

vation 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: geo‐

morphological 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

news and update

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

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/

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 nu‐

merous others who contributed to this book with

specific boxes.

This volume appears 14 years after Man‐

groves – The forgotten forest between land and

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

changed and the forgotten forest has been redis‐

covered. Obviously neither the simple existence of

this remarkable ecosystem, nor its fascinating

functioning based on adaptation to the harsh con‐

ditions 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 statis‐

tics in this book), and the change from the eco‐

logical perspective to the human perspective in

terms of coastal protection against hurricanes and

tsunamis and in carbon sequestration, is neces‐

sary to improve public awareness about the im‐

portance of mangroves for our present life and a

critical part of our response to the challenges of

environmental changes, including sea level rise

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 sup‐

port your daily work with easy‐to‐understand in‐

formation and strong facts about the ecological

and economic values of this forest. If you are a

mangrove ecologist, this book should also be on

your shelf because it provides you with a quick

overview of mangrove distribution and current

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 de‐

mand for mankind and that being involved is an

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

a general text spanning the interdisciplinary as‐

pects of mangrove ecology, this is not the book for

you. The roots of this book largely come from ge‐

ography 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 peo‐

ple’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) Man‐groves 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 Man‐grove Ecosystems. Annual Review of Marine Science, 2, 395–417.

Lugo, A.E. & Snedaker, S.C. (1974). The ecology of man‐groves. Annual Review of Ecology and Systemat‐ics, 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 Geophysi‐cal Union, Washington.

Spalding, M., Blasco, F. & Field, C. (1997). World man‐grove atlas. The International Society for Man‐grove Ecosystems, Okinawa, Japan. 178 pp.

Tomlinson, P.B. (1986). The botany of mangroves. Cam‐bridge University Press, Cambridge, UK. 419 pp.

Wibisono,I.T.C. & Suryadiputra, N.N. (2006). Study of lessons learned from mangrove/coastal ecosys‐tem restoration efforts in Aceh since the tsu‐nami. Wetlands International – Indonesia Pro‐gramme, Bogor. 86 pp.

news and update

92 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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

Edited by Markus Eichhorn

news and update

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

book review

A comprehensive foundation for the application of biogeogra‐phy to conservation Conservation biogeography, by Richard J. Ladle and Robert J. Whittaker (editors)

2011, Blackwell Publishing, 301 pp. ISBN: 9781444335033

Price: £95 (Hardback) / £34.95 (Paperback); http://eu.wiley.com/

It is becoming increasingly clear that the diversity

of plant and animal species in the world is con‐

tinuing to decline in spite of ambitious targets set

by governments to prevent this (Butchart et al.

2010). It is also becoming evident that the contin‐

ued 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 un‐

derstand the diversity of life and how it is distrib‐

uted across the biomes and ecosystems of the

world. Since understanding the distribution of bio‐

diversity 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 conser‐

vation 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 move‐

ment and the contrasting values held by different

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

well as some background to the field of conserva‐

tion biogeography (Chapter 1). A distinction is

made between approaches that focus on the com‐

position of biological communities and those that

focus on ecosystem function through an under‐

standing 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 ex‐

plain the link between the composition of biologi‐

cal communities and the function of the ecosys‐

tems 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 func‐

tioning of ecosystems and can also help in predict‐

ing how ecosystems will respond to environ‐

mental change (McGill et al. 2006).

The second section reviews our current un‐

derstanding of the distribution of biodiversity,

summarises the history of the global protected

areas network and describes the methods avail‐

able for more systematically representing biodi‐

versity in future extensions to this network. There

is a strong terrestrial focus here, indeed through‐

out the entirety of the book, which the authors

acknowledge and which is owing to a less com‐

plete 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 fin‐

ished recently, has made huge progress towards

understanding the biogeography of the oceans

ISSN 1948‐6596

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

trial 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 simulta‐

neously by predicting the spatial distribution of

undiscovered plant species, predicting that most

new plant species will be discovered in areas al‐

ready identified as hotspots of plant diversity, em‐

phasising the importance of these areas for con‐

servation. Chapter 5 provides an excellent sum‐

mary of the many different types of protected

areas in the global network and the different val‐

ues that underpin these, while Chapter 6 provides

a useful and succinct review of the enormous and

ever‐growing literature on systematic conserva‐

tion 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 descrip‐

tions of some of the available methods, from phe‐

nomenological models that infer future changes

from current patterns (Chapter 7) to more process

‐based models that use the theory of island bio‐

geography to predict the consequences for biodi‐

versity of shrinking and increasingly isolated natu‐

ral habitat patches (Chapter 8). Chapter 9 deals

with invasive species, which are an important

driver of environmental change, and the homog‐

enisation of biological communities, i.e. the ero‐

sion of beta diversity. Most of the studies investi‐

gating broad‐scale patterns of diversity have fo‐

cused 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 be‐

come increasingly important in providing the nec‐

essary 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 conserva‐

tion.

Tim Newbold United Nations Environment Programme World Con‐

servation 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 ser‐vices. Nature, 477, 199–202.

Joppa, L.N., Roberts, D.L., Myers, N. et al. (2011). Biodi‐versity 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., Lam‐oreux, 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.

Edited by Markus Eichhorn

news and update

94 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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

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

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 en‐

gendering discussion regarding whether it indeed

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

or is rather a particularly focused aspect of com‐

munity ecology or biogeography (e.g. Marris 2009,

Pyšek and Hulme 2009). As with many ecological

disciplines, invasion ecology has seen fundamen‐

tal 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 ex‐

tended 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 informa‐

tion and uneven levels of understanding and focus

across the peer community is confusion and un‐

certainty, right from the fundamentals (what is an

invasive species exactly, and why is it invasive?) to

the specifics (what is the best technique for reduc‐

ing 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 Simber‐

loff 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‐

ries of key topics relating (in this case) to biologi‐

cal invasions, without citations but with relevant

further reading at the end. The entries vary in

length from 1 to 8 pages, and often incorporate

useful figures and occasionally tables. The book is

impressively glossy (all figures are in full colour)

and well presented, which is all the more remark‐

able considering the relatively modest price. The

editors, Daniel Simberloff and Marcel Rejmánek,

are leading invasion ecologists and are well quali‐

fied 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 research‐

ers). 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 im‐

pacts and management, interesting case studies

and historical perspectives. Clearly it is not possi‐

ble to cover all of the entries in a review such as

this, but I did find several articles especially inter‐

esting, particularly because they highlight the

many socioecological factors that complicate our

relationships with potentially problematic species.

The entry on Xenophobia for example does an ex‐

cellent job of summarising how society’s relation‐

ship with non‐native species is constructed in cer‐

tain ways by the use of loaded terms or cultural

metaphors, for example the negative personifica‐

tion of zebra mussels as ‘outlaws’ on the west

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

‘distasteful’ species with appellations that note

their foreign status (Japanese knotweed, Chinese

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

starting point for a discussion of scientific objec‐

book review

A new encyclopedia for biological invasions Encyclopedia of biological invasions, by Daniel Simberloff and Marcel Rejmánek (editors)

2011, University of California Press, 792 pp. ISBN: 9780520264212

Price US$95 (Hardback or e‐book); http://www.ucpress.edu/

ISSN 1948‐6596

tivity related to invasion biology it works excep‐

tionally well, and is exactly the right size for diges‐

tion by students or interested amateurs.

Indeed, one of the best uses I find for refer‐

ence works such as these are as opening forays

into topics for class discussions, whether at gradu‐

ate or undergraduate level. Good examples in‐

clude the entry on Succession, which very effec‐

tively and concisely summarises key concepts that

take up whole chapters in many textbooks, and

although invasion biology is only addressed to‐

wards the end, it is clear how the two link to‐

gether. Likewise, the discussion on Native invad‐

ers, in which issues of ‘invasive’ terminologies

(and when they are appropriate) are covered, is

excellently written and illuminating at a range of

levels, particularly in relation to the many exam‐

ples of ‘invasion’ given. Certainly students and

researchers new to the subject will have any initial

confusion over what is meant by invasions dis‐

pelled 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 arti‐

cles 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 im‐

age of Frank Buckland ‘physicking a por‐

poise’ (page 2) is worthy of inclusion for example,

despite his role in founding the main UK acclimati‐

sation 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 sum‐

marising the development and impact of such so‐

cieties 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 legisla‐

tion, awareness of ecological risk from introduc‐

tions and lack of interest from the general public.

The article elegantly conveys how originally be‐

nevolent 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 natu‐

ralising many species at all. But much is left un‐

said: in some cases one is left wanting to know

more about whether species referred to as

‘released’ became naturalised, whether regions

such as South America maintained any such socie‐

ties (these countries are ignored, while others

such as Germany and Italy receive only one sen‐

tence) and ultimately whether such societies indi‐

rectly provided evidence to force their own dis‐

continuation. 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, encyclopae‐

dic 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 cov‐

erage 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 reason‐

able, and is not a criticism of the volume – it is

impossible to cover the vast range of topics asso‐

ciated with biological invasions in sufficient depth

in a single volume, and the material that is in‐

cluded is impressive. The division of the book be‐

tween invader attributes, processes, taxa, ecosys‐

tems, 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 con‐

gratulated. I would encourage consideration of a

second volume, however, at least with regard to

key concepts and hypotheses. The opening guide

to the Encyclopedia notes that there is a website

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

notes that the site ‘will evolve with the addition of

news and update

96 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

news and update

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

The Neotropics leave an indelible impression on

everyone who visits them. The seeds of some of

the most important concepts in ecology and evo‐

lution were sown during the South American trav‐

els of influential 19th century thinkers. For exam‐

ple, the latitudinal gradient of diversity, now rec‐

ognized 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. Al‐

though 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 esti‐

mated to consist of more than 7000 species, and

that accounts for over half the freshwater fish on

the planet and around 10% of all vertebrate spe‐

cies.

James Albert and Roberto Reis’ goal as edi‐

tors 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 di‐

versification were involved and that these oper‐

ated over long periods of time as well as on a con‐

tinental scale. The book itself is divided into two

parts, the first of which examines current knowl‐

edge on the biogeography of the region, while the

second is a regional analysis that links contempo‐

rary geographical patterns with geological history.

The book is ambitious in scope and brings to‐

gether previously fragmented material to provide

an authoritative overview of this impressive group

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

cal ichthyofauna is inevitably drawn to the Ama‐

book review

A piscine history of the Neotropics Historical biogeography of Neotropical freshwater fishes, by J.S. Albert and R.R. Reis (editors)

2011, University of California Press, 408 pp. ISBN: 9780520268685

Price £59 (Hardback); http://www.ucpress.edu/

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 poten‐

tially be a very useful resource, given the rapidly

changing environment of the internet, the publi‐

cation of a second volume would perhaps be the

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 indi‐

vidual 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.php?pid=1961

References

Davis, M.A. (2005) Invasion biology 1958‐2004: the pursuit of science and conservation. In: Concep‐tual 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. Bio‐logical Invasions, 8, 1409–1424.

Larson, B.M.H. (2007) An alien approach to invasive species: objectivity and society in invasion biol‐ogy. 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 dis‐cipline 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.

Edited by Markus Eichhorn

ISSN 1948‐6596

zon, the book has broad coverage, embracing the

Andes and extending through Central America and

into southern Mexico. As it makes clear, it is nec‐

essary to have a continental perspective to under‐

stand the diversity and distribution of this impres‐

sive group.

I particularly liked the care and thought in‐

volved in putting the book together. It is a beauti‐

fully 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 his‐

tory of South American fishes.

One of the most challenging questions in

ecology is explaining why different habitats sup‐

port 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. How‐

ever many of the fishes that inhabit this river sys‐

tem are older than the Amazon Basin itself. More‐

over, the Amazonian ichthyofauna has been accu‐

mulated gradually through tens of millions of

years. The explanation, Albert, Petry and Reis ar‐

gue, is rooted in the repeated subdivision and

merging of adjacent river basins and their faunas,

with dispersal limitation and environmental filter‐

ing 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 univer‐

sal feature of natural assemblages is that some

families contribute a much higher fraction of spe‐

cies 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, ei‐

ther separately or together, could contribute. E.O.

Wilson (2003) has argued that an ancient origin,

combined with small body size, widespread geo‐

graphic distribution and key innovations contrib‐

ute to the success of some groups relative to oth‐

ers. On the basis of the evidence presented by

Neotropical fish, Albert, Bart and Reis conclude

that these features are necessary but not suffi‐

cient. 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 sex‐

ual and trophic innovation may play a role. Eco‐

logical specialisation is also important. For exam‐

ple, 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 fac‐

tors that underpin diversification are the same as

those that come into play in the explosive speci‐

ation 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 spe‐

cies evolved. Nonetheless, as this book makes

clear, the nature and timing of key events is be‐

coming much better understood. The contribu‐

tions to the book demonstrate how the growing

body of molecular data, and its integration with

ecological theory and earth sciences, has under‐

pinned the recent and rapid progress in under‐

standing this system.

news and update

98 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

Your participation in frontiers of biogeography is encouraged. Please send us your articles, com‐ments and/or reviews, as well as pictures, drawings and/or cartoons. We are also open to sug‐gestions on content and/or structure.

Please check http://www.biogeography.org/html/fb.html for more information, or contact us at ibs@mncn.csic.es and frontiersofbiogeography@gmail.com.

news and update

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

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 stud‐

ied. 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). Har‐vard University Press.

Edited by Markus Eichhorn

books noted with interest

Principles of terrestrial ecosystem ecol‐

ogy

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 sys‐

tem and geological processes. What follows is a

magisterial and comprehensive account of the

movements of water, energy, carbon and nutri‐

ents 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 consid‐

ers global changes and the implications for ecosys‐

tem management. The book is well written

throughout and punctuated with excellent colour

illustrations; no‐one from undergraduates to es‐

tablished researchers can fail to learn something

from it.

Guide to standard floras of the World:

An annotated, geographically ar‐

ranged systematic bibliography of the

principal floras, enumerations, check‐

lists and chorological atlases of differ‐

ent areas

David F. Frodin

2001, 2nd edition, Cambridge University Press, 1100 pp.

£198 (Hardback), £90 (Paperback), US$120 (e‐book)

ISBN: 9780521790772 / 9780521189774

http://www.cambridge.org/

While not generally our policy to feature reprints,

this standard text has newly appeared in paper‐

back, 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.

ISSN 1948‐6596

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 sev‐

eral floristic realms. It contains a pictorial guide to

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

chapters on vegetative formations and should fa‐

cilitate both local and international study. Copies

have been freely distributed to universities and

institutes throughout Afghanistan as well as her‐

baria and museums worldwide. A feature on this

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 inter‐

action‐based ecology and biogeography; recent

heated debate in the pages of American Naturalist

has even disputed whether communities truly ex‐

ist as natural entities. Unsurprisingly the author

makes a strong case for communities, stressing

patterns and processes that can only be under‐

stood at this level, and pleasingly devotes equal

attention to both models and experimental data.

The textbook is intended for a graduate course

and represents a major update on the previous

edition. 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

news and update

Editorial policy for book reviews

Frontiers of Biogeography will publish in‐depth reviews of recently published books (typically less than one

year old) on biogeography or of interest to biogeographers, alongside a ‘Noted with Interest’ section provi‐

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

view to the Book Review Editor, Dr Markus Eichhorn, School of Biology, University Park, Nottingham NG7

2RD, United Kingdom; telephone ++44 (0)115 951 3214; e‐mail markus.eichhorn@nottingham.ac.uk. We

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

fic book. Anyone wishing to review books should send a brief curriculum vitae, description of competencies,

and a statement of reviewing interests to the Book Review Editor. Reviews should be in an essay style, ex‐

pressing an opinion about the value of the book, its focus and breadth, setting it in the context of recent

developments within the field of study. Textbook reviews should consider their utility as resources for tea‐

ching and learning. Avoid describing the book chapter by chapter or listing typographical errors. The length

should normally be 1000 words (1500 words for joint reviews of related texts) including a maximum 10 refe‐

rences. Authors may suggest a short heading for the review, followed by the title of the book(s), the aut‐

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

or tables will not ordinarily be included. Authors of reviews must verify that they have not offered (and will

not offer) a review of the same book to another journal, and must declare any potential conflict of interest

that might interfere with their objectivity. This may form a basis for editorial decisions and such disclosures

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ces also will be considered by one or more referees.

100 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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 de‐

scribed 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 verte‐

brates 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) sig‐

nificantly advanced the conservation of endan‐

gered 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 pro‐

tected 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 Leath‐

wick 2007). Unfortunately, these data from het‐

erogeneous sources may contain errors or

have been obtained using a biased sampling pro‐

cedure (Hortal et al. 2007, 2008, Newbold

2010). Besides, they do not usually provide reli‐

able absences needed to perform consistent pre‐

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

2007), so alternatives have been sought generat‐

ing models based only on presences (Hirzel et al.

2002, Pearce and Boyce 2006), sometimes em‐

ploying pseudo‐absences obtained in different

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

news and update

thesis abstract

Applying species distribution modeling for the conservation of Iberian protected invertebrates Rosa María Chefaoui

PhD Thesis, Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales,

c/ José Gutiérrez Abascal 2, 28006 Madrid, Spain.

e‐mail: 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 esti‐

mating species’ ranges, identifying suitable habitats, and identifying the primary factors affecting species’

distributions. The study tackles the strategies used to obtain SDMs without reliable absence data while

exploring their applications for conservation. I examine the conservation status of Copris species and

Graellsia isabelae by delimiting their populations and exploring the effectiveness of protected areas. I show that the method of pseudo‐absence selection strongly determines the model obtained, generating

different model predictions along the gradient between potential and realized distributions. After assess‐

ing the effects of species’ traits and data characteristics on accuracy, I found that species are modeled

more accurately when sample sizes are larger, no matter the technique used.

Keywords: Environmental niche modeling, Iberian Peninsula, invertebrates, predictive accuracy, species

distribution models

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

ISSN 1948‐6596

Lobo et al. 2006, 2010).

For my doctoral thesis, I evaluated the util‐

ity 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 Un‐

ion 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 Mod‐

els). I approached methodological issues concern‐

ing 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 res‐

pect, I explored the applications of SDM to esti‐

mate 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 de‐

cline in the Iberian Peninsula, play a critical eco‐

logical role in extensive pasture ecosystems by

recycling organic matter. We delimited the poten‐

tial distribution of the two species of Copris

(Coleoptera, Scarabaeidae) that inhabit the Ibe‐

rian Peninsula using ENFA (Chefaoui et al. 2005).

ENFA is a 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 environ‐

mental 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 Co‐

pris species are distributed along a geographic and

environmental gradient from the Tajo basin

(warmer, dryer, with strong annual weather varia‐

tions) where only C. hispanus is found, towards

the mountain slopes of the Sistema Central

(colder, higher rainfall) where C. lunaris predomi‐

nates. 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, mak‐

ing 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 con‐

served in the previous protected sites network:

for C. hispanus only two protected sites measured

around 30 km2, and for C. lunaris a single area

measured 183 km2. 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 substan‐

tially.

Chefaoui and Lobo (2008) assessed the ef‐

fects of pseudo‐absences on model performance

when reliable absence data are not available. We

compared seven procedures to generate pseudo‐

absence 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 distri‐

butions depending on the method and the thresh‐

old used to select these pseudo‐absences.

The results showed that the pseudo‐

absence selection method greatly influenced the

percentage of explained variability, the scores of

the accuracy measures and, most importantly, the

SDM applied to invertebrate conservation

102 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

predicted range size. As we extracted pseudo‐

absences from environmental regions further

from the optimum established by presence data,

the models obtained better accuracy scores, and

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’ con‐

servation status (Chefaoui and Lobo 2007). We

modeled the potential distribution of the insect by

performing GLM with pseudo‐absence data se‐

lected from an ENFA model. We found that the

best predictor variables were summer precipita‐

tion (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. ni‐

gra.

Moreover, we found 8 areas exclusively in

the eastern Iberian territory, and a larger unoccu‐

pied habitat in the western Iberian Peninsula, indi‐

cating that this species is probably not in equili‐

brium with its environment because of historical

factors (Chefaoui and Lobo 2007). We sug‐

gested 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 penin‐

sula. After analyzing the possibility of connectivity

and fragmentation of the eight populations delim‐

ited as well as the degree of protection of G. isa‐

belae on the SCIs, we found that the SCIs under

protection did not seem sufficient to maintain cu‐

rrent 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 sylves‐

tris and P. nigra located both inside and near to

SCIs, we suggested that the reintroduction of the

species in these habitats could improve its conser‐

vation.

To understand the limitations and possibili‐

ties 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 pseudo‐

absences to model the distribution of 20 threa‐

tened 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 habi‐

tats 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. Pres‐

ence‐only methods (ENFA and MDE) achieved

worse validation results and overpredicted more

than techniques using pseudo‐absences. Never‐

theless, 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 de‐

termined the predicted range size, generating dif‐

ferent model predictions along the gradient be‐

tween potential and realized distributions. There

is an added difficulty in obtaining predictions that

closely approximate the realized distribution of

species under non‐equilibrium conditions, be‐

cause both presence and absence data may be

possible under similar environmental conditions.

Irrespective of the approach used, species’ distri‐

butions are modelled more accurately when sam‐

Rosa M. Chefaoui

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

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 Sci‐

ence Faculty Library, Universidad Autónoma de

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

PDF copy is also available at request from the au‐

thor.

Acknowledgements

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

Lobo and Joaquín Hortal for their support and en‐

couragement.

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Edited by Richard Pearson

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

Rosa M. Chefaoui

opinion

Political erosion dismantles the conservation network existing in the Canary Islands José María Fernández‐Palacios and Lea de Nascimento

Island Ecology and Biogeography Group, Instituto Universitario de Enfermedades Tropicales y Salud

Pública de Canarias (IUETSPC), Universidad de La Laguna (ULL), Avda. Astrofísico Francisco Sánchez s/n,

38206, La Laguna, (Tenerife), Spain

e‐mail: jmferpal@ull.es; http://webpages.ull.es/users/jmferpal

ISSN 1948‐6596 opinion and perspectives

Recently the Canarian Parliament has approved a

new version of the Canarian catalogue of pro‐

tected species (see Box 1) that reduces substan‐

tially both the number of species included (from

466 species in the 2001 list to 361 species in the

2010 list) and the protection afforded (from 381

threatened species to 160, and from 85 protected

species to 18). These reductions have been widely

criticized by environmental NGOs and the local

scientific community1, mainly due to the absence

of a rigorous scientific process in its development.

Although certainly the first version of the cata‐

logue could be improved, the main reasons be‐

hind the new revisions were not conservation is‐

sues 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 forbid‐

den due to their impacts on protected species in‐

cluded in the original version of the Canarian cata‐

logue.

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 scien‐

tific community.

Within the new revised catalogue a com‐

pletely new criterion for protection has emerged

“especies de interés para los ecosistemas canar‐

ios” (literally: “species of interest for Canarian eco‐

systems”), comprising 152 species (see Box 1). The

phrase is poorly chosen. It is supposed to apply

only to endangered species, consequently the fre‐

quent and abundant species which usually struc‐

ture and dominate the ecosystems are explicitly

not listed, leading to a curious paradox: the Ca‐

narian pine (Pinus canariensis) is not a species of

interest for the Canarian pine forest, the

Macaronesian Laurel (Laurus novocanariensis) is

Abstract. The outstanding nature of the Canary Islands has been recognized by European, national and

regional administrations since the arrival of democracy in Spain. Forty‐five per cent of its emerged terri‐

tory has been declared as Natural Protected Areas, four Canarian National Parks were included within the

Spanish network, more than 200 endemics were listed in the Spanish catalogue of endangered species,

and 450 species were listed in the Canarian catalogue of protected species. However, in recent years, po‐

litical decisions have started dismantling this splendid conservation network, which impedes construction

of large infrastructure, golf courses and resorts, despite the advice of the scientific community. Canarian

nature is now facing two threats: delisting and downgrading of numerous endangered species, and trans‐

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

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

tion, scientific community, species delisting

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

106 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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 desig‐

nated Natural Protected Area (NPA). (In the Ca‐

naries, that means in either the Canarian Network

of NPAs or the European Union Natura 2000 Net‐

work, which overlap extensively). If a listed spe‐

cies, for instance the woodcock (Scolopax rusti‐

cola) or the coot (Fulica atra) which are both in‐

cluded 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 in‐

consistency affects, for instance, ca. 10 endemic

species of sea lavenders (Limonium spp.) pro‐

tected in certain ravines, but not in others.

The new law could have negative implica‐

tions for conservation biogeography, and this can

be illustrated with some examples of the Canarian

flora and fauna. The endemic legume Cicer ca‐

nariensis, previously considered as vulnerable in

the 2001 Canarian catalogue, is now included un‐

der the criterion species of interest. From its 12

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

the six populations in the North of La Palma2 are

outside NPAs and therefore unprotected accord‐

ing to the new law. Metapopulation dynamics in

this species could be affected by this new criterion

if source populations within these northern loca‐

tions are threatened, endangering sink popula‐

tions included in NPAs. The same could apply to

the Abalone or Canarian clam (Haliotis tuberculata

ssp. coccinea) or the Sea Horse (Hippocampus hip‐

pocampus). Both are marine species with sparse

populations in the meso‐ and infra‐littoral, which

do not always coincide with the geographical loca‐

tion of the marine Special Areas for Conservation,

which occupy mainly leeward fringes on the Archi‐

pelago’s coasts. Collection and capture of both

species is prohibited by the Regulation of the Fish‐

José María Fernández‐Palacios and Lea de Nascimento

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.

Box 1

Law 4/2010, June 4, of the Canarian Catalogue of Protected Species (see the original Spanish text at

http://www.gobiernodecanarias.org/boc/2010/112/)

Article 3. Canarian protected species

2) Species of interest for Canarian ecosystems

The Canarian Catalogue of Protected Species will also include “species of interest for Canarian ecosys‐

tems" which are those that, without being listed in the threatening situations above (endangered or vul‐

nerable), are worthy of particular attention for its ecological significance in areas of the Canarian Network

of Natural Protected Areas or Natura 2000 network.

2. Effects of inclusion in the Catalogue

b) The legal regime for protection of “species of interest for Canarian ecosystems" will be applicable only

in the territory of the Canarian Network of Natural Protected Areas or Natura 2000 Network. To this end,

applicable measures shall be provided by the management plans of Natural Protected Areas and Habitats

of the Natura 2000 Network in which they are located. Such plans shall include the determinations, con‐

trol and monitoring to ensure effectiveness of protection, or where applicable, the justification that there

is no need for plans. (...) In the case of actions promoted by reasons of public interest and priority affect‐

ing the “species of interest for Canarian ecosystems" these actions could be possible as long as they do

not affect the ecosystem substantially, under the terms in paragraphs 4 to 7 of the Article 45 of the Law

42/2007, December 13, of Natural Heritage and Biodiversity.

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

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 In‐

terest 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 com‐

munity from an ecological point of view as they

play an important role in the carbon cycle, stabi‐

lize sandy soils, export biomass and act as a fish

nursery area (Barberá et al. 2005). The latter char‐

acteristic is also very important for the sustainabil‐

ity 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 there‐

fore 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 pro‐

posal submitted by the Canarian Government, the

Port Authority and the Canarian Company of Gas

Transportation, to delist C. nodosa3. 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 infrastruc‐

tures of already existing harbours), that were hid‐

den from the European Commission by Spain’s

National Government.

This controversial criterion — especies de

interés para los ecosistemas canaries — is an ad‐

aptation 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 down‐

graded threatened species. However in the former

criterion there were no restrictions in the protec‐

tion, 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, frag‐

mented or limited. The previous criterion for pro‐

tection 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 metapopula‐

tion dynamics, genetic diversity or viability of

populations, but simply in protecting less de‐

graded remnants of communities that were still

available. As in many parts of the world, reserves

were not designed to meet the principles of sys‐

tematic conservation planning needed to achieve

representativeness and persistence of biodiversity

(Margules and Pressey 2000). The situation fur‐

ther 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 distri‐

bution throughout the windward slope of the is‐

land. From the point of view of mitigating the ef‐

fects of global change, vulnerability of certain spe‐

cies 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 prece‐

dent in Spain). The Catalonian Plan of Areas of

Natural Interest includes species of flora and

fauna strictly protected in designated areas. To

our knowledge no cases of the failure of these

practices or public disapproval have been re‐

ported there, but we suspect that the species with

restricted protection in the Catalonian NPA Net‐

108 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

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

José María Fernández‐Palacios and Lea de Nascimento

work were not demoted from higher protection.

In theory, the main aim of the existence of re‐

gional catalogues is ensuring the protection of

particular species that are not considered by the

National Catalogue. On the other hand several

authors have questioned and analysed the effec‐

tiveness of NPAs Networks in biodiversity conser‐

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

and concluded that reserve networks are geo‐

graphically and taxonomically unbalanced leaving

a big proportion of endemic and threatened spe‐

cies 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 com‐

petently in the Canaries’ Marine Reserves with

Fishery Interest, but is nonsensical when the aim

of the declaration is to protect a threatened spe‐

cies. 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 numer‐

ous public protests and the clear opposition of the

majority of the Canarian scientific community, the

new catalogue was presented by the leading po‐

litical force in the Regional Parliament. These

kinds of conflicts are not exclusive to the Canary

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 con‐

cern, 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), Ti‐

manfaya (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 Constitu‐

tional Court now has determined that NPs man‐

agement is exclusively a matter for the Regional

Governments. Consequently the Central Govern‐

ment has transferred all management to the re‐

gions. 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 Govern‐

ment expressed her intention to discuss again this

transfer and to limit the management of the is‐

land 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 conserva‐

tion, have been charged with environmental

crimes5. Although some implications of decentrali‐

zation should be positive, for instance the creation

of regional lists and plans considering the particu‐

lars 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 Au‐gust/2011); http://www.eldia.es/2011‐04‐13/CANARIAS/5‐Es‐frecuente‐alcaldes‐esten‐imputados‐delitos‐urbanisticos.html (last accessed August/2011); http://www.elpais.com/articulo/espana/corrupcion/presenta/elecciones/elpepiesp/20110410elpepinac_1/Tes (last accessed August/2011); http://www.europapress.es/islas‐canarias/noticia‐imputados‐canarias‐logran‐mantenerse‐instituciones‐20110524094822.html (last accessed Au‐gust/2011) .

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

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 ac‐

tively 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, to‐

gether with other potential emerging threats. De‐

spite the capacity and good work of environ‐

mental technicians, who struggle with budget cuts

every year, the Councils have demonstrated a tra‐

jectory of inefficiency and lack of commitment to

the management of NPAs. Within the new Ca‐

narian NPs framework, the rabbits will receive the

responsibility of taking care of the lettuces.

Acknowledgements

We would like to thank Rafael Loyola and three

anonymous reviewers for their comments on the

manuscript. We are also grateful to the editorial

board of Frontiers in Biogeography for their help

improving this paper.

References

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Barberá, C., Tuya F., Boyra C., Sanchez‐Jerez P., Blanch I. & Haroun R.J. (2005) Spatial variation in the structural parameters of Cymodocea nodosa seagrass meadows in the Canary Islands: a mul‐tiscaled approach. Botanica Marina, 48, 122–126.

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Servicio de Biodiversidad (2009). Evaluación de espe‐cies catalogadas de Canarias: Cicer canariensis [Ciccan 06/2009]. Consejería de Medio Ambien‐te y Ordenación Territorial, Gobierno de Cana‐rias, Las Palmas de Gran Canaria. Available at h t t p : / / w w w . g o b c a n . e s / c m a y o t /medioambiente/medionatural/biodiversidad/especies/especies_protegidas_amenazadas/

Edited by Joaquín Hortal & Michael N Dawson

You can find information about the International Biogeography Society at http://www.biogeography.org/, and contact with other biogeographers at the IBS blog (http://biogeography.blogspot.com/), the IBS facebook group (http://www.facebook.com/group.php?gid=6908354463) and the IBS twitter channel (https://twitter.com/biogeography).

110 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

Canarian conservation network dismantled

Introduction

Rediscoveries of putatively extinct species are of

great potential interest to both conservationists

and biogeographers (Crowley 2011). For the for‐

mer, ‘rediscovery’ can be a considerable conserva‐

tion policy and publicity asset (Ladle and Jepson

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

global initiatives: in 2009 BirdLife International

launched a “global bid to try to confirm the con‐

tinued existence of 47 species of bird that have

not been seen for up to 184 years” (BirdLife Inter‐

national 2009). The following year Conservation

International launched its “Search for lost Frogs”

which involves a dedicated campaign and expedi‐

tions to 18 countries seeking to locate 40 species

not seen for a decade or more (Conservation In‐

ternational 2010) – at the time of writing 12 spe‐

cies have been rediscovered. Moreover, since re‐

discovered species are typically exceedingly rare

and geographically localized, new knowledge on

population status and distribution supports effec‐

tive conservation interventions. Finally, rediscov‐

eries remove uncertainty from extinction risk as‐

sessments; a confirmed new record moves the

species from ‘extinct’ or ‘probably extinct’ and

into an IUCN threat (or data deficient) category.

For biogeographers, species rediscovery has both

a practical and conceptual significance. From the

ISSN 1948‐6596

perspective

The causes and biogeographical significance of species’ rediscovery Richard J. Ladle1,2,*, Paul Jepson2, Ana C. M. Malhado1,

Steve Jennings3 and Maan Barua2

1. Institute of Biological and Health Sciences, Federal University of Alagoas, Maceió, AL, Brazil. 2. School

of Geography and the Environment, University of Oxford, South Parks Road, Oxford, OX1 3QY, United

Kingdom. 3. Oxfam GB, Oxfam House, John Smith Drive, Oxford, United Kingdom.

*Author for correspondence: Dr Richard J. Ladle, Institute of Biological and Health Sciences, Federal Uni‐

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

e‐mail: 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 tech‐

nology to find target species; (3) the accessibility of the areas, habitats or sites where the species are

thought to survive; and (4) the ease with which a species can be located when it is present within a habi‐

tat. Thus, species are ‘lost’ from scientific knowledge for different reasons and, consequently, not all

rediscoveries are equally significant for biogeographical research or conservation. Indeed, rediscoveries

of species that underwent a well documented decline and disappearance – and are therefore of greatest

potential importance for both conservation and biogeographical research – appear to be poorly repre‐

sented in the literature compared to rediscovered species that were only known from a handful of mu‐

seum specimens. Thus, carefully distinguishing between the causes of temporal gaps in zoological re‐

cords is essential for improving the utility of rediscovery data for biogeographical research and conser‐

vation practice.

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

opinion and perspectives

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

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 ad‐

dress 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 de‐

clared 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 fre‐

quently 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 redis‐

covered species relative to its historical range re‐

flects the pattern of range collapse. Thus, if an‐

thropogenic pressures (e.g. unsustainable exploi‐

tation) are strongest at the periphery (Channel

and Lomolino 2000) the rediscovery will most

likely be made near the centre of the historic

range. Diana 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 redis‐

coveries 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 ele‐

vations can sometimes provide ecological refugia

(Towns and Daugherty 1994) and fits with the fre‐quently 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 surpris‐

ing since rediscovery and extinction are conceptu‐

ally intertwined; extinction is the permanent ab‐

sence of current and future records while redis‐

covery reflects the temporary absence of such

records. Moreover, rediscovery is the proof re‐

quired to refute a hypothesis of extinction. Given

the close conceptual linkage between the con‐

cepts of rediscovery and extinction it is interesting

that, until recently, there have been so few stud‐

ies linking patterns of rediscovery to contempo‐

rary 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 signifi‐

cance, and which can be subject to meaningful

analysis. Here, we propose a conceptual frame‐

work for understanding and analyzing species re‐

discovery, based on the social, institutional and

ecological factors that created the temporal gap in

occurrence data. We believe that formalizing the

concept of rediscovery in this way has the poten‐

tial to create new measures of the state of knowl‐

edge of the world’s rarest species, provide a quan‐

tifiable metric to support existing endangerment

categorizations, and would help to maintain the

culture of biogeographical exploration that con‐

tributes to the datasets that underpin global con‐

servation target‐setting, advocacy and monitoring.

Conceptual framework

The ‘loss’ of a species and its subsequent rediscov‐

ery can be conceptualized as a result of the inter‐

play among four socio‐ecological aspects of redis‐

covery (schematically illustrated in Figure 1): (1)

the state of knowledge of species loss and redis‐

covery; (2) the presence of people and/or organi‐

zations 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 qual‐

ity, funding and the culture of scientific explora‐

tion, rediscovery research has focused almost ex‐

clusively on herptiles, birds and mammals (cf.

Scheffers et al. 2011).

rediscoveries in biogeography

112 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 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 appar‐

ently ‘lost’. For example, BirdLife International has

made significant investments in compiling new

and authoritative assessments of threatened spe‐

cies using information from a variety of sources

including amateur and university‐led research ex‐

peditions 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 profes‐

sional and amateur ornithologists (Tobias et al.

2006, Butchart 2007).

The knowledge of what is ‘lost’ is compli‐

cated, as rediscoveries can logically be split into

four categories that reflect different degrees of

uncertainty (and authority) about the continued

existence of a target species (Table 1). An addi‐

tional category could potentially be added to this

typology to account for cases where an unre‐

corded 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 estab‐

lished 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 White‐

eye (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 re‐

discovery could seriously influence research find‐

ings. 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 cen‐

tre of their historic range as are better‐known

species. Moreover, all four categories of rediscov‐

ery 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, Schef‐

fers et al. (2011) found that the majority of re‐

cently claimed amphibian, bird and mammal re‐

discoveries represent first documentations since

their original scientific description. It should also

be noted that such rare species may have re‐

mained 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 rediscov‐

ery (see McCarthy 2008; Fisher and Blomberg

2011).

Figure 1. The four major dimensions of species rediscovery (see text).

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

Perhaps the most important type of redis‐

covery 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 redis‐

covery in 1992 (in the stomach of a snake) con‐

firmed that the species now has “a dramatically

reduced geographical range” (Milne and Bull

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

Woodpecker (Campephilus principalis) (Fitzpatrick

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

cept that this rediscovery is increasingly looking

like a case of mistaken identity (Dalton 2005,

2010, Stokstad 2007). The apparent scarcity of

such rediscoveries (cf. Scheffers et al. 2011)

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 ex‐

ist 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 19th 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 inappropri‐

ate vocalizations”. Thus, ‘lost’ species can some‐

times turn up thousands of kilometres away from

where they were last seen, or in completely differ‐

ent 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

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

Type Rediscovery of… Example

1. a species declared extinct by an authori‐

tative 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

The Sao Tome Grosbeak (Neospiza concolor) was

described as probably extinct by Greenway (1967)

and rediscovered in 1991 (Sergeant et al. 1992)

3. a species believed to be still extant but

for which substantive searches over dec‐

ades have drawn a blank.

According to the NGO BirdLife International the

Madagascar Serpent Eagle (Eutriorchis astur) was

not definitely recorded between 1930 and 1993 de‐

spite considerable search‐effort within its habitat.

4. a species whose habitat or range had not

been surveyed for a significant period of

time, but for which there is no real rea‐

son to assume has become extinct

The Chestnut‐bellied Flowerpiercer (Diglossa glorio‐

sissima) was unrecorded for 38 years: since 2003 it

has been recorded from three locations (Tobias et

al. 2006)

114 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

rediscoveries in biogeography

level, has varied considerably over time and space

in response to various cultural and ecological fac‐

tors. Most notably, the mainstreaming of biodiver‐

sity 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 conserva‐

tion organizations (Jepson and Ladle 2010). More

generally, increased funding of expeditions by in‐

ternational NGOs has probably been the driving

force behind the increasing frequency of rediscov‐

eries of various taxa (Scheffers et al. 2011).

Other trends within science and conserva‐

tion also help determine the capacity and motiva‐

tion that enables rediscoveries, especially the in‐

troduction of new technology. For example, ad‐

vances in molecular biology have made it much

easier to genetically compare preserved type

specimens in museums with contemporary mate‐

rial collected directly or acquired from hunters or

from rural markets. This has opened the way for

completely new ways of rediscovering lost spe‐

cies, where a fragment of hair or a faecal sample

may be sufficient to prove the continuing exis‐

tence of a species that has still not been physically

observed.

An excellent example of such a technology‐

aided 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 se‐

quences derived from old museum specimens

with samples extracted from dung samples re‐

cently collected in the field. Such remotely col‐

lected DNA evidence can also be used to discount

presumed discoveries or rediscoveries. For exam‐

ple, Hennache et al. (2003) used a range of tech‐

niques, including captive hybridization experi‐

ments and analysis of mitochondrial DNA and mi‐

crosatellites, to conclusively demonstrate the hy‐

brid origin of the imperial pheasant (Lophura im‐

perialis). This mysterious bird had first been cap‐

tured 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 avail‐

ability of sophisticated audiovisual equipment has

been especially important in the evolution of bird

surveying. Two such technological advances, the

increased availability of less expensive sound‐

recording and playback equipment in the late

1990s and the more recent internet‐based bird‐

sound 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 re‐

corders, 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 19th Cen‐

tury, 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 proba‐

bly 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), rediscov‐

ered in 2003 in Angola (Ryan et al 2004).

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

Richard J. Ladle et al.

A closely related factor is a lack of commu‐

nication with remote and isolated rural communi‐

ties who may already have knowledge of the con‐

tinued 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 communica‐

tion with remote tribes and communities whose

knowledge of local biodiversity may extend con‐

siderably beyond that of conservationists. How‐

ever, Fisher and Blomberg (2011) found that hu‐

man population overlap did not predict rediscov‐

ery rate in mammals, possibly because expedi‐

tions 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 exis‐

tence 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 pheno‐

typic 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 be‐

haviour were not significantly associated with re‐

discovery – although smaller rediscovered mam‐

mals 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 ap‐

proached will only flush at close quarters, then fly

low over short distances before plunging back into

cover (Forshaw and Cooper 2002). Perhaps unsur‐

prisingly, 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 sys‐

tem (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 pre‐

viously reported as globally extinct or possibly ex‐

tinct. It should be noted, however, that this ap‐

proach 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 – conceiva‐

bly the same individual.

Using a simple time‐based criterion would

provide a single, objective definition of rediscov‐

ery – whatever the cause of the gap in zoological

records. Conservation bodies could potentially use

this definition to periodically produce lists of spe‐

cies 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 re‐

corded (e.g. <25 years ago, 25–49 years ago, 50–

100 years ago, >100 years ago, etc.). One advan‐

tage of such a system would be to maintain and

extend the practice of biogeographical expedi‐

tions to remote areas. It would also help guard

against the overuse or misrepresentation of redis‐

116 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

rediscoveries in biogeography

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 repre‐

sents a real opportunity for conservationists to

reassert core values and raise funds that may help

protect poorly known habitats. Moreover, redis‐

coveries provide a unique source of information

about the rarest and least‐known species (for cer‐

tain taxa) that can be used to investigate bio‐

geographic theories about range loss and extinc‐

tion. Both of these important agendas would

benefit from a greater systematization of the con‐

cept 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 pro‐

vides a wonderful opportunity to assess both the

subtle ecological and biogeogeographical charac‐

teristics of exceptionally rare species of well stud‐

ied 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 interact‐

ing factors (Fisher 2011a,b; Fisher and Blomberg

2011, Scheffers et al. 2011), while the recent tar‐

geting of ‘lost species’ by international conserva‐

tion NGOs is generating considerable amounts of

valuable new data. Nevertheless, the lack of redis‐

covered species that were previously well known

and which had undergone a well documented

process of population decline, fragmentation and

local extinction (Scheffers et al. 2011) remains a

worrying trend for global conservation.

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Channell, R. & Lomolino, M.V. (2000) Trajectories to extinction: spatial dynamics of the contraction of geographical ranges. Journal of Biogeography, 27, 169–179.

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Fisher, D.O. & Blomberg, S.P. (2011) Correlates of redis‐covery and the detectability of extinction in mammals. Proceedings of the Royal Society B: Biological Sciences, 278, 1090–1097.

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Hennache, A., Rasmussen, P., Lucchini, V., Rimondi, S. & Randi, E. (2003) Hybrid origin of the imperial pheasant Lophura imperialis (Delacour and Jabouille, 1924) demonstrated by morphology, hybrid experiments, and DNA analyses. Biologi‐cal Journal of the Linnean Society, 80, 573–600.

Jepson, P.J. & Ladle, R.J. (2010) Conservation: a begin‐ner’s guide. One World, Oxford.

Ladle, R.J. & Jepson, P.R. (2008) Toward a biocultural theory of avoided extinction. Conservation Let‐ters, 1, 111–118.

Ladle, R.J., Jepson, P., Jennings, S. & Malhado, A.C.M. (2009) Caution with claims that a species has been rediscovered. Nature, 461, 723.

Lomolino, M.V., Riddle, B.R. & Brown, J.H. (2006) Bio‐geography. 3rd edn. Sinauer, Sunderland, MA.

Mace, G.M., Collar, N.J., Gaston, K.J., Hilton‐Taylor, C., Akçakaya, H.R., Leader‐Williams, N., Milner‐Gulland, E.J. & Stuart, S.N. (2008) Quantification of extinction risk: IUCN's system for classifying threatened species. Conservation Biology, 22, 1424–1442.

McDougall, A., Porter, G., Mostert M., Cupitt R., Cupitt S., Joseph, L., Murphy S., Janetzki H., Gallagher A. & Burbidge A. (2009) Another piece in an Aus‐tralian ornithological puzzle –a second Night Parrot is found dead in Queensland. Emu, 109, 198–203.

McCarthy, M.A. (1998) Identifying declining and threat‐ened species with museum data. Biological Con‐servation, 83, 9–17.

Milne, T. & Bull, C.M. (2000) Burrow choice by individu‐als of different sizes in the endangered pygmy blue tongue lizard Tiliqua adelaidensis. Biologi‐cal Conservation, 95, 295–301.

Pitra, C., VazPinto, P., O’Keeffe, B.W.J., Willows‐Munro, S., Jansen van Vuuren, B. & Robinson, T.J. (2006) DNA‐led rediscovery of the giant sable antelope in Angola. European Journal of Wildlife Research, 52, 145–152.

Rasmussen, P.C., Wardill, J.C., Lambert, F.R., & Riley, J. (2000) On the specific status of the Sangihe White‐eye Zosterops nehrkorni, and the taxon‐omy of the Black‐crowned White‐eye Z. atrifrons complex. Forktail, 16, 69–80.

Riddle, B., Ladle, R.J., Lourie, S. & Whittaker, R.J. (2011) Basic biogeography: estimating biodiversity and mapping nature. In: Ladle, R.J. & Whittaker, R.J. (Editors) Conservation Biogeography. Oxford University Press, Oxford, pp. 47–92.

Round, P.D., Hansson, B., Pearson, D.J., Kennerley, P.R. & Bensch, S. (2007). Lost and found: the enig‐

matic large‐billed reed warbler Acrocephalus orinus rediscovered after 139 years. Journal of Avian Biology, 38, 133–138.

Rozendaal, F.G. & Lambert, F.R. (1999) The taxonomic and conservation status of Pinarolestes sanghirensis Oustalet 1881. Forktail, 15, 1–13.

Ryan, P.G., Sinclair, I., Cohen, C., Mills, M.S.L., Spottis‐woode, C. & Cassidy, R. (2004) The conservation status and vocalisations of threatened birds from the scarp forest of the Western Angola Endemic Bird Area. Bird Conservation Interna‐tional, 14, 247–260.

Scheffers, B.R., Yong, D.L., Harris, J.B.C., Giam, X. & Sodhi, N.S. (2011) The World’s rediscovered species: back from the brink? PLoS ONE 6, e22531.

Sergeant, D.E., Gullick, T., Turner, D.A. & Sinclair, J.C. (1992) The rediscovery of the São Tomé Gros‐beak Neospiza concolor in south‐western São Tomé. Bird Conservation International, 2, 157–159.

Stephenson, B.M., Flood, R., Thomas, B. & Saville, S. (2008) Rediscovery of the New Zealand storm petrel (Pealeornis maoriana Mathews 1932): two sightings that revised our knowledge of storm petrels. Notornis, 55, 77–83.

Stresemann, E. (1931) Die Zosteropiden der indo‐australischen Region. Mitteilungen aus dem Zoologischen Museum Berlin, 17, 201–238.

Stokstad, E. (2007) Gambling on the ghost bird. Sci‐ence, 317, 888–892.

Svensson, L., Prŷs‐Jones, R., Rasmussen, P.C. & Olsson, U. (2008) Discovery of ten new specimens of large‐billed reed warbler Acrocephalus orinus, and new insights into its distributional range. Journal of Avian Biology, 39, 605–610.

Timmins, R.J., Mostafawi, N., Rajabi, A.M., Noori, H., Ostrowski, S., Olsson, U., Svensson, L. & Poole, C.M. (2010) The discovery of Large‐billed Reed Warblers Acrocephalus orinus in north‐eastern Afghanistan. BirdingASIA, 12, 42–45.

Tobias, J.A., Butchart, S.H.M. & Collar, N.J. (2006). Lost and found: a gap analysis for the Neotropical avifauna. Neotropical Birding, 1, 4–22.

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Edited by Jan Beck

118 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

rediscoveries in biogeography

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 bio‐

geographers are also becoming increasingly in‐

volved within the IBS. From 2002 to 2010, the pro‐

portion 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 cul‐

minated in the first IBS Early Career conference

that was held at Oxford University from 23 to 25

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

html/Meetings/index.html).

With the intention of getting to know its

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 in‐

vited 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 institu‐

tions (50% and 33% respectively; total of 42 an‐

swers), they represent a total of 24 nationalities;

26% are from North America, 17% from Central

and South America, 15% from Northern Europe,

28% from Southern Europe, and the other 12%

from Australia/New Zealand, the Middle East, Af‐

rica and Asia. ECM work on a very broad range of

topics, from species distribution patterns (the

most mentioned topic), to evolutionary biogeog‐

raphy, dispersal and colonization, biogeography of

species’ traits, island biogeography, phylogeogra‐

phy, 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, con‐

servation, paleontology, geography, botany, mam‐

malogy, 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; re‐

sponses to open‐ended questions emphasized the

need for more off‐year meetings (regional meet‐

ings, workshops, etc.), more jobs/grant announce‐

ments, more travel grants, online teaching re‐

sources, more talks at the IBS meetings by

younger researchers and more opportunities to

meet other researchers. The IBS is already work‐

ing towards improving the services it provides to

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 biogeogra‐

phers who have finished their PhDs in the past five

years). Almost ninety young researchers partici‐

pated and had the chance to present their work,

and to interact with each other and with the IBS

board members. This conference was organized

into ten different sessions that covered several

aspects of macroecology, island biogeography,

phylogeography, paleobiogeography, evolutionary

biogeography and conservation biogeography.

Second, we are also working towards in‐

creasing regular communication among IBS mem‐

bers. One way of doing this is through online so‐

cial networks, such as Facebook, and other web‐

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

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

has a Facebook group with ~590 members, where

ISSN 1948‐6596

from the society

Getting to know IBS Early Career Members

membership corner

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

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% actu‐

ally 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 mem‐

bers, is the online journal Frontiers of Biogeogra‐

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

This journal has a section especially devoted for

this purpose – the membership corner – of which

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

cent 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 ab‐

stracts, 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 organ‐

ized 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 pref‐

erence). 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 discus‐

sion 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 spe‐

cific research topics and methodologies (31% and

29%, respectively; there was a tie between meet‐

ings on specific geographic realms and on a broad

scope within biogeography – 20% each). Some

respondents also called for workshops and semi‐

nars, online courses, cross‐society ventures to

boost interaction between similarly oriented aca‐

demics and excursions into biogeographically in‐

teresting regions covering a broad range of taxa.

There was also a significant interest in having a

showcase at the next IBS meeting of funding agen‐

cies from different countries (70%), with most re‐

spondents being willing to provide information on

this matter (55%).

The long‐term success of any growing soci‐

ety 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,

membership corner

Did you know that any member of the IBS may raise an issue or appeal a decision of the gover‐ning Board of Directors by placing a matter before the Board of Directors for discussion?

If there is a matter you would like discussed at the next Board meeting, write to the society's Secretary (check current list of officers at http://www.biogeography.org/).

120 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

to help organizing off‐year activities, and to sub‐

mit manuscripts to Frontiers of Biogeography. The

IBS wants to hear and share more of the early ca‐

reer 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 Biogeog‐

raphy 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 soci‐

ety. 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. Fron‐tiers of Biogeography, 3, 71‐73.

Edited by Matthew Heard

membership corner

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

from the society

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

We are seeking proposals for hosting the 7th bien‐

nial conference of the International Biogeography

Society to be held in early January 2015. Proposals

should be submitted by individuals who are inte‐

rested in chairing the local (host) committee. The

duties of the local host include conducting contra‐

ct negotiations with the venue and the hotel as

well as all local logistics including field trip organi‐

zation and production of the abstract bo‐

ok. 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 ca‐

pacity 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 Euro‐

pe. 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 ins‐

titution or organization.

2. What would be the benefit of hosting the con‐

ference at this location?

3. Actual site of the meeting and the capacity of

the auditorium.

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

cation relative to the auditorium.

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

nue. A specific quote is not needed, but evi‐

dence 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 orga‐

nizing 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;

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) bio‐geographers, or you want to make a call for manuscripts or talks, please contact us at ibs@mncn.csic.es and frontiersofbiogeography@gmail.com.

Erratum ‐ 11 Feb 2011: The original article published on 09 Feb 2011 incorrectly identified Dan Gavin as the "IBS Student‐at‐Large".

membership corner

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 frag‐

mentation

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/

21st 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/

3rd European Congress of Conservation Biol‐

ogy

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/

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 facul‐

ty positions: Ecology (Full or Associate with tenu‐

re, or Assistant tenure‐track), Systems Biology

(Assistant tenure‐track), and Biostatistics

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

logy Lecturer. For the Ecology position, we seek

outstanding individuals with research interests in

any ecological field using experimental, field, com‐

putational, and/or theoretical approaches and

working at population to global scales. The Sys‐

tems Biology position includes research areas that

use comprehensive datasets and multiple types of

analysis to relate overall biological function to un‐

derlying biochemical or biophysical processes for

predictive understanding. The Biostatistics rese‐

arch areas of interest include statistical methods

for experimental design, epidemiology, medical

informatics, evolutionary biology, sequence bioin‐

formatics, genomics, evolution of microbial sys‐

tems and pathogens, and systems biology. The

Lecturer position closely parallels a tenure‐track

Assistant Professor but with an emphasis on un‐

dergraduate 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

F912A3B023DA4F1F80?seriesId=1

Interested applicants should submit mate‐

rials online. Applications will be considered star‐

ting 05 December 2011 (Biostatistics, Systems Bio‐

logy professorships), or 16 December 2011

(Ecology professorship and Biology Lecturer). UC

Merced is an AA/EOP employer.

122 © 2011 the authors; journal compilation © 2011 The International Biogeography Society — frontiers of biogeography 3.3, 2011

Job announcements

table of contents

ISSN 1948‐6596

news and update

update: Species–area curves and the estimation of extinction rates, by J. Beck 81

update: Extinct or extant? Woodpeckers and rhinoceros, by R. Ladle 83

update: Climate wars, by J. Beck 84

update: Emerging research opportunities in global urban ecology, by F.A. La Sorte 85

update: Beyond taxonomical space: large‐scale ecology meets functional and phylogenetic diversity, by M.V. Cianciaruso

87

book review: A mangrove compendium, by U. Berger 91

book review: A comprehensive foundation for the application of biogeography to conservation, by T. Newbold 93

book review: A new encyclopedia for biological invasions, by R.A. Francis 95

book review: A piscine history of the Neotropics, by A.E. Magurran 97

books noted with interest 99

thesis abstract: Applying species distribution modeling for the conservation of Iberian protected invertebra‐tes, by R.M. Chefaoui

101

opinion and perspectives

opinion: Political erosion dismantles the conservation network existing in the Canary Islands, by J.M. Fernán‐dez‐Palacios & L. de Nascimento

106

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

membership corner

from the society: Getting to know IBS Early Career Members, by A.M.C. Santos 119

Job announcements 122

Upcoming meetings 122

from the society: Call for proposals for hosting 7th Biennial Conference of the IBS, by D. Gavin 121

frontiers of biogeography the scientific magazine of the International Biogeography Society volume 3, issue 3 ‐ November 2011

frontiers of biogeography copyright notice

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