Species: A Modest Propasal 35

to be filled (or not) by suitable organisms (a classic example is Elton

1927). A niche imposes demands on its occupants, and these demands
explain similarities within and differences across species. But ecolo-
gists no longer think that we can assume that communities, built from
different species, nonetheless have a common organization, that, say, a
temperate rainforest in British Columbia will make available the same
array of occupations as one on New Zealand’s west coast. And so niches
are now defined by the organisms that occupy them, by vectors of their
resource requirements (Griesemer 1992). Moreover, organisms alter
both their own physical and biological environment and those of oth-
ers. Trees stabilize soils; moderate storm impacts; and provide shelter,
resources, and concealment for a host of other organisms (Jones et al.
1997; Lewontin 1985; Odling-Smee et al. 2003).
So the niches of some species depend largely on the geology and
meteorology of their habitat and the ecological milieu that determines
their place in the food webs and nutrient cycles of which they are a part.
In other species, selection for “ecological engineering” plays a much
stronger role. Earthworms, for example, are adapted to an aquatic envi-
ronment. They only survive out of water because they have evolved the
ability to alter their own environment. They reduce surface litter; aggre-
gate soil particles; and increase levels of organic carbon, nitrogen, and
polysaccharides, which enhances plant yields and improves porosity,
aeration, and drainage. In so doing they co-opt the soils they inhabit and
the tunnels they build to “serve as accessory kidneys and compensate
for their poor structural adaptation” (Odling-Smee et al. 2003, 375).
Thus niche occupation appears much more active in some species than
others. Furthermore, the idea that each species has a unique niche is
more plausible in some cases than in others. Scavenging generalists
battle it out in a widely contested and “open to all comers” niche. By
contrast, figs and fig wasps have coevolved; each provides an essential
and specific service to the other.
The idea that a species lives in, and is shaped by, a unique niche
turns out to glide over a complex and variable set of relationships be-
tween species and environments. The same is true of the apparently
straightforward idea of reproductive isolation. It turns out to be a
cover-all label for a large variety of prezygotic and postzygotic interac-
tions that largely keep lineages separate. The great strength of the bio-
logical species concept is that reproductive isolation and the resulting
restriction in gene flow are real and important facts in the evolution
of populations and metapopulations. However, there are many ways in
which gene flow can be restricted. Australian immigrant species that
are wind-borne to New Zealand shores face the blustery Tasman Sea