Are Ecological And Evolutionariy Rules Being Dismissed Prematurely?

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DDI_115.fm Page 289 Thursday, October 25, 2001 9:04 PM

Diversity and Distributions (2001) 7, 289– 295

B I O D I VE R S I T Y V I E W P O I N T S

Are ecological and evolutionary rules being dismissed
prematurely?
Blackwell Science Ltd

K YLE G. ASHTON Department of Environmental, Population and Organismic Biology, University
of Colorado, Boulder, CO 80309, U.S.A.

Abstract.Recently, it has been suggested that
the group of patterns known as the ecological
and evolutionary rules are invalid and only of
interest historically. Here, I briefly review evidence for the four most prominent patterns:
Bergmann’s rule, Cope’s rule, Rapoport’s rule,
and the latitudinal gradient in species richness.
Although more comprehensive reviews exist for
these patterns, the purpose of thispaper is to
highlight recent work illustrating the validity of
each pattern, to question the conclusion that

I NTRODUCTION
A general goal of science is to document broad
patterns and understand the mechanisms that
generate those patterns (MacArthur, 1972). This
is particularly true for the fields of ecology and
evolution. In fact, ecology has been described
as ‘the study of patterns innature, how those
patterns came to be, how they change in space
and time, why some are more fragile than others
(Kingsland, 1995).’ Documenting and understanding the processes that generate broad
patterns in ecology and evolution are important
steps towards understanding biodiversity.
Recently, a group of patterns in ecology and
evolution, the ecological and evolutionary rules,
have beencalled into question (Geist, 1987;
Gould, 1997; Gaston et al., 1998a). Clearly,
many ‘rules’ have been proposed without the

Present address: Archbold Biological Station, 123 Main Drive,
Venus, Florida 33960, USA. E-mail: kashton@archboldstation.org

these patterns are only of historical interest, and
to briefly explore possible connections between
each of these patterns. I also recommendthat
meta-analytical techniques, a group of statistical
methods rarely used in ecology and evolution,
need to be incorporated in future tests of general
trends.
Key words. Ecological rules, evolutionary rules,
Bergmann’s rule, Rapoport’s rule, Cope’s rule,
species diversity gradient.

necessary supporting evidence. However, there
appears to be an attitude in recent literature suggestingthat the study of ecological and evolutionary rules is of primarily historical interest,
and that many of the proposed patterns are
invalid (e.g. Gould, 1997; Gaston et al., 1998a).
For instance, Gaston et al. (1998a, p. 70) wrote:
The list of evolutionary and ecological ‘rules’
and ‘laws’ is brief, and primarily of historical
rather than current interest. Of those that have
been postulated,most have subsequently
proven not to exhibit the generality which
was initially claimed, or have been found to
be artifactual (e.g. Bergmann’s Rule, Cope’s
Rule ... ).
However, the studies cited by Gaston et al.
(1998a) to substantiate their argument that these
rules are invalid and primarily of historical interest are problematic. In particular, none of the
references they cite forBergmann’s rule (Geist,
1987; Currie & Fritz, 1993; Hawkins & Lawton,
1995) reviewed general intraspecific patterns of
body size variation for endothermic vertebrates,

© 2001 Blackwell Science Ltd. http://www.blackwell-science.com/ddi

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DDI_115.fm Page 290 Thursday, October 25, 2001 9:04 PM

290

K. G. Ashton

t he common definition of Bergmann’s rule
(Rensch, 1938; Mayr, 1956).Rather, they concern
arguments about the mechanisms responsible for
Bergmann’s rule (Geist, 1987), interspecific
patterns for a group of ectothermic organisms
(Hawkins & Lawton, 1995), or interspecific
patterns based on unclear methodology and
taxonomic sampling (Currie & Fritz, 1993).
Additionally, the one study cited by Gaston et al.
(1998a) to dismiss Cope’s rule (Jablonski, 1997)
has...
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