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OIKOS 75: 303-309. Copenhagen 1996
Abundance-body size relationships: the area you census tells
you more
Tim M. Blackburn and Kevin J. Gaston
Blackburn, T. M. and Gaston, K. J. 1996. Abundance-body size relationships: the
area you census tells you more. - Oikos 75: 303-309.
mil ihe
d Piper
15-109.
hemical
cos 68:
The interspecific relationship between abundance and body size in animals is often
claimed to be strongly negative, with species abundance limited by energetic require
ments. This view has been criticized for a number of reasons, but is still widely
accepted. Here, we provide evidence of further fundamental difficulties with this
relationship as derived from compendium studies. We suggest that there is a potential
artcfactual component to these relationships resulting from variation in the areas
over which the densities of species of different body size are censused, and differences
in the ways species use these areas. While the interspecific relationship between body
size and abundance is still likely to be negative after accounting for the artefactual
component, the slope of the relationship is unlikely to support energetic equivalence
arguments.
Carbon
tebrate
Owcnf planl
avanna
id leaf
cfcnse.
T. M. Blackburn, NERC Centre for Population Biology, Imperial College at Silwood
Park, Ascot, Berkshire, U.K. SL5 7PY. - K. J. Gusiun, Dept of Entomology. Natural
History Museum. Cromwell Rd. London, U.K. SW7 5BD (present address: Dept of
Biology, Imperial College at Silwood Park, Ascot, Berkshire, U.K. SL5 7P Y).
-•%urce
}30:
1990.
i cnvima of
-335.
y: The
ido. -
cmical
•n balale of
C.L.,
id the
source
eneral
. Phy•cand
-715.
Considerable attention has been focused on the inter
specific relationship between abundance and body size
(Damuth 1981, 1987, 1993, Peters 1983, Peters and
Wasscnberg 1983, Peters and Raelson 1984, Juanes
1986, Brown and Maurer 1987, Marquet et al. 1990,
Cotgreave and Harvey 1991, 1992, 1994, Nee et al.
1991, Blackburn et al. 1993a, b. Cotgreave 1993, 1994,
Currie 1993, Blackburn and Lawton 1994). Initial stud
ies revealed a strong, negative interaction, with body
mass generally explaining 60-70% of the variation in
animal abundance (with both variables log-trans
formed; Damuth 1981, 1987, Peters 1983, Peters and
Wassenberg 1983, Peters and Raelson 1984). Further,
the slope of this relationship typically approximated to
-0.75 within trophic groups (e.g. mammalian herbi
vores), using ordinary least squares regression. Since
body mass scales with metabolic rate to the 0.75 power
(Kleiber 1962), the -0.75 exponent between size and
abundance was taken as evidence that a species' abun
dance is limited by its energetic requirements, and that
equal amounts of energy are available to each species in
a community: the so-called 'energetic equivalence rule'
(Damuth 1981. 1987, 1991, Nee et al. 1991).
The view that a species' abundance is determined by
its body size has been criticised (Brown and Maurer
1986, 1987, Lawton 1989, 1991, Blackburn et al. 1993a,
b, Blackburn and Lawton 1994). The principal difficulty
is that the data used to establish the -0.75 relationship
between abundance and body size are not derived from
samples of whole communities, but tend to be compendia front the literature. They maytheretore"undercstimatc thcTnumber of rare species, and small-bodied rare
species in particular (Brown and Maurer 1987, Morse
et al. 1988, Lawton 1989, 1991). Subsequent studies
sampling whole assemblages of taxonomically similar
animals have revealed polygonal relationships between
/
Accepted 5 July 1995
Copyright © OIKOS 1996
ISSN 0030-1299
ff
^GVILK
Printed in Ireland all rights reserved
(19%)
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densities; our own statistical treatment (see above)
highlights the problem, but does not solve it. Even
aside from the problem of co-linearity, controlling for
area essentially assumes that intraspecific density-area
relationships have similar trajectories, and that biologi
cally comparable densities can be generated by standar
dising study area (the same problem occurs in whole
assemblage studies where all densities are estimated
using a common study area). The first assumption
seems likely to be erroneous; intraspecific density-area
relations certainly differ markedly (Fig. 4). The second
assumption seems equally dubious. A common measure
of ecological density is unlikely to equate to measure
ment of density over the same sized area: elephants and
dik-diks, for example, will certainly make different use
of a standardised study area. The veiling of density
estimates in whole assemblage studies is another conse
quence of a fixed sample area. That species density
_ (IWft)
OIKOS 75:2 |l-%)
Fig. 4. The relationship between log,,, density and log,,, census
area in a) Peromyscus maniculatus (Rodcnlia). slope = —0.818.
r2 = 0.766. p < 0.0005, n — 11; b) Aepyccms mulampus (Artiodactyla), slope = -0.021. r2 = 0.001, p> 0.9. « = 9 (excluding
the obvious outlier, slope = -0.44, r2 = 0.458, />< 0.065,
n = 8); and c) Loxodonta africana (Proboscidea),
slope = —0.155. r2 = 0.287, p <0.06, n = 13. Each point is the
mean density (km-2) and mean area (km2) in one study.
musl be correlated lo some extent with the area over
which density is measured, because the latter is used in
the calculation of the former, is irrelevant to these
arguments. The problem of which density of Per
omyscus maniculatus to compare with which density of
Loxodonta africana remains.
This said, it is noteworthy that when controlling for
the size of study area, the resulting body size: abun
dance relationship (Fig. 3) seems broadly intermediate
in form between relationships documented in com
pendium studies (Damuth 1981, 1987, 1993, Peters and
Wassenberg 1983, Peters and Raelson 1984) and rela
tionships documented in whole assemblage studies
(Brown and Maurer 1987, Gaston 1988, Morse et al.
1988, Blackburn et al. 1993a), with characteristics
of both patterns (see also Gregory and Blackburn
1995). For example, the low correlation is similar to
that seen in assemblage studies, but the lack of veiling
307
at low abundances is characteristic of compendium
studies.
Conclusion
There is a growing list of criticisms of studies describing
a strong negative relationship between body size and
ecological density based on compendium data, and
attributing that relationship to energetic equivalence
(see reviews in Cotgreave 1993, Blackburn and Lawton
1994), including problems of undersampling rare spe
cies, likely inequalities in energy distribution between
species (Lawton 1989, 1991), and combining exponents
excluding error terms (see e.g. Sugihara 1989, Black
burn and Gaston 1994. Medel et al. 1995). Problems
arising from the confounding effect of census area can
now be added to this list. It seems reasonable to
conclude that a strong interspecific relationship between
the body size of a species and the area over which its
density is measured results from a combination of
biolocical and non-biological factors. This results in
differences in the kinds of densities calculated for spe
cies of different body sizes, and contributes to a strong
negative relationship between the density and body size
of species. While the density size relationship is still
likely to be negative after accounting for area effects,
the suggestion that this relationship provides evidence
for energetic equivalence seems unlikely.
Acknowledgements We thank Bob Molt for helpful discus
sion Peter Coigreave, David Currie. Jeremy Greenwood and
John Lawlon for comments on earlier versions oT this
manuscript, and the staff of the library at the Natural History
Museum. London, for their co-operation wiih l.terah.rc
searches. T.M.B. was supporled by NhRC grant GR3/8029.
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