The most exciting advances in science often
involve scientific conflict and debate. These can be friendly and cordial
exchanges, or they can be acrimonious and personal. Scientists often wed
themselves to their ideas and can be quite reluctant to admit that their
precious idea was wrong. Students in ecology often learn about some of these
classic debates (Clements v. Gleason; Diamond v. Simberloff and Connor), but
often other debates fade from our collective memory. Scientific debates are important things to study, they tell
us about how scientists function, how they communicate, but more importantly by
studying them we are less likely to repeat them! Take for example the debate
over species per genus ratios, which happened twice, first in the 1920s, then
again in the 1940s. The second debate happened in ignorance of the first, with
the same solution being offered!
To understand the importance of testing
species-genus ratios we can start with a prediction from Darwin:
As species of the same genus have
usually, though by no means invariably, some similarity in habits and
constitution, and always in structure, the struggle will generally be more
severe between species of the same genus, when they come into competition with
each other, than between species of distinct genera (Darwin 1859)
To test this hypotheses, the Swiss
botanist, Paul Jaccard (1901) created a ‘generic coefficient’ to describe biogeographical
patterns and to measure the effects of competition on diversity. The generic
coefficient was a form of the species-genus ratio (S/G), calculated as G/S x
100, and he interpreted a low S/G ratio (or high coefficient) to mean that
competition between close relatives was high, and a high ratio (low
coefficient) meant that there was a high diversity of ‘ecological conditions’
supporting closely related species in slightly different habitats (Jaccard 1922). At the same time as Jaccard was working on his generic coefficient,
the Finnish botanist, Alvar Palmgren, compiled S/G patterns across the Aland
Islands and inferred the low S/G values on distant islands to reflect random
chance (Palmgren 1921). Over several years, Jaccard and Palmgren had a heated exchange in
the literature (across different journals and languages!) about interpreting
S/G ratios (e.g., Jaccard 1922, Palmgren 1925). Palmgren’s contention was that the S/G ratios he observed were
related to the number of species occurring on the islands –an argument which
later work vindicates. A few years after their exchange, another Swiss
scientist, Arthur Maillefer, showed that Jaccard’s interpretation was not
supported by statistical inference (Maillefer 1928, 1929). Maillefer created what is likely one of the first null model in
ecology (Jarvinen 1982). He calculated the expected relationship between Jaccard’s generic
coefficient and species richness from ‘chance’ communities that were randomly
assembled (Fig. 1 –curve II). Maillefer rightly concluded that since the
number of genera increase at a slower rate than richness, the ratio between the
two couldn’t be independent of richness.
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Jaccard’s generic coefficients plotted by
Maillefer showing the relationship between the coefficients (calculated as
Genera/Species x 100) and species richness (Maillefer 1929). The four curves depict different scenarios.
Curve I shows the maximum values possible, and curve IV is the minimum. Curve
III is when coefficients are calculated on sampled values from a flora, which
stays on a mean value. Curve II represents the first null model in ecology,
where species are randomly sampled (‘hasard’ is translated as chance or luck)
and the coefficient was calculated from the random assemblages.
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This example is especially poignant because
it foreshadowed another debate 20 years later –and not just in terms of using a
null expectation, but that S/G ratios cannot be understood without comparison
to the appropriate null. Elton (1946) examined an impressive set of studies to show that small
assemblages tended to have low S/G ratios, which he thought indicated
competitive interactions. Mirroring the earlier debate, Williams (1947), showed that S/G ratios were not independent of richness and that
inferences about competition can only be supported if observed S/G values
differed from expected null values. However, the error of inferring competition
from S/G ratios without comparing them to null expectations continued into the
1960s (Grant 1966, Moreau 1966), until Dan Simberloff (1970) showed, unambiguously, that, independent of any ecological
mechanism, lower S/G are expected on islands with fewer species. Because he
compared observationed values to null expectations, Simberloff was able to show
that assemblages actually tended to have higher S/G ratios than one would
expect by chance (Simberloff 1970). So not only is competition not supported, but the available
evidence indicated that perhaps there were more closely related species
on islands, which Simberloff took to mean that close relatives prefer the same
environments (Simberloff 1970).
Darwin, C. 1859. The origin of the species by means of
natural selection. Murray, London.
Elton, C. S. 1946. Competition and the
Structure of Ecological Communities. Journal of Animal Ecology 15:54-68.
Grant, P. R. 1966. Ecological Compatibility
of Bird Species on Islands. The American Naturalist 100:451-462.
Jaccard, P. 1901. Etude comparative de la
distribution florale dans une portion des Alpes et du Jura. Bulletin de la
Societe Vaudoise des Sciences Naturelle 37:547-579.
Jaccard, P. 1922. La chorologie selective
et sa signification pour la sociologie vegetale. Memoires de la Societe
Vaudoise des Sciences Naturelle 2:81-107.
Jarvinen, O. 1982. Species-To-Genus Ratios
in Biogeography: A Historical Note. Journal of Biogeography 9:363-370.
Maillefer, A. 1928. Les courbes de Willis:
Repar- tition des especes dans les genres de diff6rente etendue. Bulletin de la
Societe Vaudoise des Sciences Naturelle 56:617-631.
Maillefer, A. 1929. Le Coefficient
générique de P. Jaccard et sa signification. Memoires de la Societe Vaudoise
des Sciences Naturelle 3:9-183.
Moreau, R. E. 1966. The bird faunas of
Africa and its islands. Academic Press, New York, NY.
Palmgren, A. 1921. Die Entfernung als
pflanzengeographischer faktor. Series Acta Societatis pro Fauna et Flora
Fennica 49:1-113.
Palmgren, A. 1925. Die Artenzahl als
pflanzengeographischer Charakter sowie der Zufall und die säkulare Landhebung
als pflanzengeographischer Faktoren. Ein pflanzengeographische Entwurf, basiert
auf Material aus dem åländischen Schärenarchipel. Acta Botanica Fennica 1:1-143.
Simberloff, D. S. 1970. Taxonomic Diversity
of Island Biotas. Evolution 24:23-47.
Williams, C. B. 1947. The Generic Relations
of Species in Small Ecological Communities. Journal of Animal Ecology 16:11-18.
*This text has been modified from a forthcoming book on ecophylogenetics authored by Cadotte and Davies and published by Princeton University Press