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Biodiversity hotspots
Walter V. Reid
Copyright 1998, Elsevier Science Ltd. All rights reserved. 0169-5347/98/$19.00 PII: S0169-5347(98)01363-9
275
REVIEWS
North America
160
r = 0.375
n = 208
120
80
40
10
15
20
Australia
25
r = 0.531
n = 67
200
150
100
10
15
20
India
25
r = 0.726
n = 61
450
300
150
12
24
36
48
60
276
REVIEWS
surprising, therefore, that
studies are showing that at
fine scales of resolution hotspots of certain taxa do not
reflect hotspots of other
taxa. For indicator taxa to be
useful in setting priorities,
a careful balance needs to
be struck between the geographic and taxonomic scale
of resolution, and the type of
conservation question being
addressed.
Higher-taxon diversity as a
surrogate for species
diversity
Fish
Frogs
Tortoises
Snakes
Birds
Mammals
0.24 (110)
0.54 (98)
0.19 (118)
0.67 (3)
0.19 (144)
0.35 (31)
0.61 (128)
0.25 (153)
1.00 (2)
0.22 (172)
0.00 (5)
0.00 (9)
0.62 (133)
0.88 (32)
0.65 (155)
0.35 (31)
0.49 (47)
0.10 (10)
0.47 (19)
0.34 (177)
0.48 (31)
0.72 (47)
0.00 (12)
0.59 (61)
0.41 (39)
0.36 (28)
0.62 (47)
0.08 (12)
0.42 (59)
0.47 (68)
0.00 (21)
0.00 (7)
0.20 (10)
0.14 (14)
0.25 (24)
0.00 (10)
0.18 (22)
0.42 (33)
0.20 (15)
0.24 (25)
0.19 (16)
0.44 (18)
0.22 (9)
0.16 (19)
0.40 (20)
Instead of relying on indicator taxa to reflect disaNumbers in parentheses are total possible overlaps.
tribution patterns in lesserknown taxa, other studies
have examined the potential
value of other surrogate measures of species richness or endemism, including vegetation Setting conservation priorities
classes, land classes, environmental variables (e.g. pre- Setting global priorities
cipitation and net primary productivity) and richness at
Biodiversity hotspot analysis was originally used to
higher taxonomic levels. A number of these surrogates have identify large regions, typically the size of an entire nation,
been shown to have useful predictive power, but higher that deserved conservation attention, such as Madagascar,
taxon-diversity appears to be most closely correlated to Northern Borneo, or the Philippines. The 18 global hotpatterns of species diversity20. Most of this research has spots defined by Norman Myers have since been used by the
focused on patterns of species richness.
MacArthur Foundation to target its grantmaking. OrganAt both coarse and fine geographic scales, richness of izations including the World Wide Fund for Nature-India and
genera and families have proved to be relatively good pre- Conservation International also set priorities among coundictors of species richness21,22. For example, at a continental tries or regions using Myers rankings.
scale (using 611 000 km2 grid cells), 99% of the variation in
The studies discussed here provide reasonable support
bird species richness in North America can be explained by for the utility of hotspots that are defined by using species
genus richness, and 91% of variation by family richness20. richness and endemism at a global or continental scale of
Similarly, at a fine scale in 35 forest reserves ranging in size analysis. At this coarse scale, patterns of richness and enfrom 1830 000 ha in Sri Lanka, 96% of woody-plant species demism tend to correspond reasonably well across taxa. By
richness can be explained by genus richness, and 86% of its nature, a continental or global hotspot analysis tends to
variation by family richness22.
identify regions with relatively small overlap in species comCan higher-taxon richness patterns be combined to cre- position Madagascar is unlikely to share many species with
ate even better surrogates for overall species diversity? The Borneo, for example.
challenge here is to determine how best to weight numbers
of genera or families in different taxa, because higher taxo- Using hotspots to guide conservation decisions
nomic groups may be defined in different ways, particularly
At more fine-grained geographic scales, which tend to be
where the taxa are distantly related. In one coarse-grained the scales at which conservation and development decisions
(global) study examining different methods for combining are made, the application of hotspot analysis is more chalfamily richness in plants, reptiles, amphibians and mammals lenging19,24,25. To begin with, the poor correspondence be(absolute number of families; proportional family richness; tween species hotspots in different taxa implies that priorand proportional family richness weighted for total species ities determined on the basis of either richness or endemism
richness in each group), all three methods gave fairly simi- in a few taxonomic groups cannot be relied upon to capture
lar results23. At other scales of analysis or using other taxo- similar patterns in other groups.
nomic groups, however, the different methods would be exJust as important, the choice of method for defining a hotpected to produce different patterns. Which method to use spot whether it is based on richness, endemism, threat or a
depends on the objective of the analysis and the availability combination of these factors significantly influences which
of data (e.g. good estimates of total species richness might regions or sites are identified as conservation priorities.
not be available for some taxa).
Site selection algorithms based on identifying hotspots
Patterns of higher taxon richness can therefore serve as of species richness tend to be the least efficient in maximizsurrogates for species richness. It remains to be seen whether ing the protection of species diversity. This is because
this same correspondence holds for patterns of endemism. hotspots of species richness do not often include relatively
The utility of this surrogate measure for conservation plan- rare species hotspots that are ranked highest for richness
ning hinges on the seemingly reasonable but untested as- often contain overlapping sets of common species, while
sumption that if only a partial survey of a region has been failing to capture rarer species. In the UK, 43% of rare bird
undertaken, the cumulative list of higher taxa encountered species do not occur in the top 5% of species richness hotwill converge on the total number of taxa more rapidly than spots18. In South Africa, the top 5% of richness hotspots for
will the cumulative list of species.
fish only include 66% of the total diversity of fish species 19.
TREE vol. 13, no. 7 July 1998
277
REVIEWS
(a)
(b)
Number of plants
Number of birds
Zero/no data
One
Two
3 to 4
5 to 7
8 to 18
19 to 77
(c)
Zero/no data
One
Two
Three
Four
5 to 13
(d)
Number of fish
Zero/no data
One
Two
Three
Four
5 to 7
Number
of molluscs
Zero/no data
One
Two
Three
4 to 5
6 to 14
Fig. 2. The geographic distribution of four groups of endangered species in the United States. (a) Plants, (b) birds, (c) fish and (d) molluscs. The maps illustrate the
number of Federally listed endangered species in each country. Alaska and Hawaii are shown in the bottom left-hand corner of the maps (not to scale). From Ref. 13,
with permission.
278
REVIEWS
This objective helps to reduce the potential overlap in species
composition if only a limited number of protected areas are
to be established. For example, if we were to identify the top
two hotspots of species richness in a region encompassing
two biogeographic regions, both hotspots could fall within a
single region and the overlap in species composition could
be quite high. If, instead, the top hotspot in each region was
selected that is, representative samples of the ecosystems
were chosen then overlap in species lists would decrease,
and the total number of species protected could increase.
When only richness or endemism hotspots are used in the
analysis, protection of representative samples using this
method helps to increase the complementarity of site selection. World Wildlife Fund-USA, for example, recently identified 232 conservation priorities by first stratifying data into
major habitat types and biogeographic realms and then examining species richness and endemism (and other factors)
to identify relatively rich ecoregions within each group32.
No matter which approach to priority setting is used,
however, if the areas conserved capture the diversity of the
indicator taxa only and not that of the more poorly-known
taxa, then the use of the method for conserving biodiversity
is questionable. Recent studies, demonstrating the failure of
hotspots of richness and endemism to correspond across
different taxa, seem to undermine the utility of hotspots in
priority setting. However, on closer examination the message
of these studies is not so bleak: these same studies lend support to the hypothesis that a set of areas in which one major taxon is well represented can also represent diversity in
unrelated taxa.
For example, in the UK, a recent study found that the
greatest overlap in hotspots among five taxonomic groups
was only 34%. However, if it were possible to protect every
hotspot designated for birds (that is, the top 5% of grid cells
with the most species of birds), then 87% of birds, 100% of
butterfly species and over 90% of dragonflies, liverworts and
aquatic plants would be encompassed18. Similarly, it appears
that a set of areas in Oregon, USA, which completely represents one major taxon does a good job of representing diversity in others24. This result is not surprising. Any site selection approach, whether it is based on richness, rarity or
complementarity, that captures most of the diversity in one
taxon is likely to include a diversity of habitats and, therefore, capture a substantial amount of the diversity of other
taxa as well. This is the case even if the areas of high richness or endemism do not correspond.
279
CORRESPONDENCE
23 Williams, P.H., Gaston, K.J. and Humphries, C.J. (1997) Mapping
biodiversity value worldwide: combining higher-taxon richness
from different groups, Proc. R. Soc. London Ser. B 264, 141148
24 Csuti, B. et al. (1997) A comparison of reserve selection algorithms
using data on terrestrial vertebrates in Oregon, Biol. Conserv. 80,
8397
25 Williams, P. et al. (1996) A comparison of richness hotspots, rarity
hotspots, and complementary areas for conserving diversity of
British birds, Conserv. Biol. 10, 155174
26 Curnutt, J. et al. (1994) Hotspots and species diversity, Nature 367,
326327
27 Ceballos, G. and Brown, J.H. (1995) Global patterns of mammalian
diversity, endemism, and endangerment, Conserv. Biol. 9,
559568
28 Blackburn, T.M. and Gaston, K.J. (1996) Spatial patterns in the
species richness of birds in the New World, Ecography 19, 369376
Extra-pair paternity in
birds: good-genes and
something else
In a recent TREE article, Petri and Kempenaers1
reviewed determinants and variation of extra-pair
paternity (EPP) in birds, emphasizing the
good-genes hypothesis: females paired to
relatively poor-quality males will seek extra-pair
copulations (EPCs) from males of better quality to
improve the fitness of some of their offspring1.
This is a widely accepted hypothesis for EPP in
birds and assumes a genetic basis for ranking
quality traits and related fitness (good genes). If
the task of females is to select the highest ranking
male possible, they should not seek EPP when
their social mate is a good-quality mate.
However, increasing evidence indicates that
females often do so2,3, which suggests that
(according to this hypothesis) they can still
discriminate some differences. This is unlikely
because the amount of phenotypic variability
resulting from geneenvironment covariance and
interaction is not negligible4,5 and should mask
small genetic differences. In addition, male quality
is often not correlated with their ability to obtain
extra-pair fertilizations2 and males bearing
relatively large ornaments can be cuckolded3. The
authors also attributed low proportions of EPP in
bottleneck populations to the good genes
hypothesis because the proportion of EPP and the
amount of variation in male quality are correlated.
However, there are alternative explanations for
Letters to TREE
Correspondence in TREE may address topics raised in very recent issues of
TREE, or (occasionally) other matters of general current interest to ecologists and
evolutionary biologists. Letters should be no more than 500 words long with a
maximum of 12 references and one small figure; original results, new data or new
models are not allowed. Letters should be sent by e-mail to TREE@elsevier.co.uk.
The decision to publish rests with the Editor, and the author(s) of any TREE article
criticized in a Letter will normally be invited to reply. Full-length manuscripts in
response to previous TREE articles will not be considered.
280
Pedro J. Cordero
Museo Nacional de Ciencias Naturales,
Consejo Superior de
Investigaciones Cientficas,
Jos Gutierrez Abascal 2,
28006 Madrid, Spain
(mcnctr@fresno.csic.es)
References
1 Petrie, M. and Kempenaers, B. (1998) Trends
Ecol. Evol. 13, 52 58
2 Bridgot, J.M. et al. (1997) Behav. Ecol. Sociobiol.
40, 119 126
3 Cordero, P.J., Wetton, J.H. and Parkin, D.T.
J. Avian Biol. (in press)
4 Bailey, R.C. (1997) Genetica 99, 125 133
5 Veiga, J.P. and Puerta, M. (1996) Proc. R. Soc.
London Ser. B 263, 229 234
6 Charlesworth, D. and Charlesworth, B. (1987)
Annu. Rev. Ecol. Syst. 18, 237268
7 Brown, J.L. (1997) Behav. Ecol. 8, 60 65
8 Svensson, E. and Skarstein, F. (1997) Trends
Ecol. Evol. 12, 9293
9 Jennions, M.D. (1997) Trends Ecol. Evol. 12,
251253
10 Potts, W.K. and Wakeland, E.K. (1990) Trends
Ecol. Evol. 5, 181187