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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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frontiers of biogeography aims to be a forum for biogeographers and a way to disseminate research in biogeography to the general public; our scope includes opinions, perspectives, and reviews, symposia proceedings, letters to the editor, book reviews, research upda‐tes, interviews, and articles on how to teach, disseminate and/or apply biogeographical knowledge. Letters to the editor and symposium proceedings may include novel analyses of original datasets (see editorial instructions). Manuscripts should be submitted to frontiersof‐[email protected]. Editorial enquiries should be made to the Editor‐in‐Chief at [email protected].
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you can find information about the International Biogeography Society at http://www.biogeography.org/; for the latest job announce‐ments and other news please visit also the IBS blog (http://biogeography.blogspot.com/), and the IBS facebook group (http://www.facebook.com/group.php?gid=6908354463).
President: Lawrence R. Heaney President Elect: Rosemary Gillespie VP for Conferences: Daniel Gavin VP for Public Affairs & Communications: Michael N Dawson VP for Development & Awards: George Stevens Secretary: Richard Field Treasurer: Lois F. Alexander Director‐at‐large: Catherine Graham Director‐at‐large: Kathy Willis Student‐at‐large: Ana M. C. Santos
First Past President: James H. Brown Second Past President: Mark V. Lomolino Third Past President: Brett R. Riddle Fourth Past President: Vicki Funk Fifth Past President: Robert J. Whittaker Upcoming meeting host (ex officio): Kenneth Feeley Past Graduate student representative (ex officio): Matthew Heard
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
news and update ISSN 1948‐6596
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: [email protected];
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: [email protected];
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: [email protected];
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: [email protected];
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: [email protected];
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: [email protected]‐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
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One of the benefits open to IBS members is the opportunity to have job openings posted on the IBS blog (http://biogeography.blogspot.com/). If you have a position you would like to have ad‐vertised, please contact Karen Faller ([email protected]) or Michael Dawson ([email protected]) with details.
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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: [email protected]; 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 [email protected] and [email protected].
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: [email protected]
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 [email protected]. We
welcome offers to review books for Frontiers of Biogeography, but will not accept an offer to review a speci‐
fic book. Anyone wishing to review books should send a brief curriculum vitae, description of competencies,
and a statement of reviewing interests to the Book Review Editor. Reviews should be in an essay style, ex‐
pressing an opinion about the value of the book, its focus and breadth, setting it in the context of recent
developments within the field of study. Textbook reviews should consider their utility as resources for tea‐
ching and learning. Avoid describing the book chapter by chapter or listing typographical errors. The length
should normally be 1000 words (1500 words for joint reviews of related texts) including a maximum 10 refe‐
rences. Authors may suggest a short heading for the review, followed by the title of the book(s), the aut‐
hors/editors, publisher, publication date, price, hbk/pbk, pages, ISBN and website (where available). Figures
or tables will not ordinarily be included. Authors of reviews must verify that they have not offered (and will
not offer) a review of the same book to another journal, and must declare any potential conflict of interest
that might interfere with their objectivity. This may form a basis for editorial decisions and such disclosures
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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: [email protected]; http://www.biogeografia.org/
Abstract. This article outlines the approaches to modeling the distribution of threatened invertebrates
using data from atlases, museums and databases. Species Distribution Models (SDMs) are useful for esti‐
mating species’ ranges, identifying suitable habitats, and identifying the primary factors affecting species’
distributions. The study tackles the strategies used to obtain SDMs without reliable absence data while
exploring their applications for conservation. I examine the conservation status of Copris species and
Graellsia isabelae by delimiting their populations and exploring the effectiveness of protected areas. I show that the method of pseudo‐absence selection strongly determines the model obtained, generating
different model predictions along the gradient between potential and realized distributions. After assess‐
ing the effects of species’ traits and data characteristics on accuracy, I found that species are modeled
more accurately when sample sizes are larger, no matter the technique used.
Keywords: Environmental niche modeling, Iberian Peninsula, invertebrates, predictive accuracy, species
distribution models
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: [email protected]; 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.
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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: [email protected]; http://www.geog.ox.ac.uk/staff/rladle.html
Abstract. The rediscovery of a species that was putatively considered to be extinct can provide valuable
data to test biogeographical hypotheses about population decline and range collapse. Moreover, such
rediscoveries often generate much‐needed publicity and additional funds for the conservation of rare
species and habitats. However, like extinction, rediscovery is challenging to define. In this perspective
we argue that the ‘loss’ of a species and its subsequent rediscovery can be understood in terms of the
interplay among four socio‐ecological factors: (1) the state of knowledge of species loss and rediscovery;
(2) the presence of people and/or organizations with the interest, motivation, resources, skills and tech‐
nology to find target species; (3) the accessibility of the areas, habitats or sites where the species are
thought to survive; and (4) the ease with which a species can be located when it is present within a habi‐
tat. Thus, species are ‘lost’ from scientific knowledge for different reasons and, consequently, not all
rediscoveries are equally significant for biogeographical research or conservation. Indeed, rediscoveries
of species that underwent a well documented decline and disappearance – and are therefore of greatest
potential importance for both conservation and biogeographical research – appear to be poorly repre‐
sented in the literature compared to rediscovered species that were only known from a handful of mu‐
seum specimens. Thus, carefully distinguishing between the causes of temporal gaps in zoological re‐
cords is essential for improving the utility of rediscovery data for biogeographical research and conser‐
vation practice.
Keywords: extinction, range collapse, rarity, critically endangered, monitoring
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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matic large‐billed reed warbler Acrocephalus orinus rediscovered after 139 years. Journal of Avian Biology, 38, 133–138.
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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.
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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.
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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: [email protected]
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
Dan Gavin IBS Vice‐President for Conferences;
Department of Geography, University of Oregon, USA.
e‐mail: [email protected]
If you want to announce a meeting, event or job offer that could be of interest for (some) bio‐geographers, or you want to make a call for manuscripts or talks, please contact us at [email protected] and [email protected].
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.