Species: A Modest Propasal 37

2.3 the effect of speciation

Speciation is problematic, as we have just seen, because of the variety of

mechanisms through which one lineage can become two. All that these
mechanisms have in common is their effect; dividing lineages acquire
independent evolutionary trajectories. It is also true that independence
is a matter of degree. Moreover, the transitivity of gene flow makes it
possible for there to be gene flow between two groups of species even
though members of one group are not able to interbreed with mem-
bers of the other group. In some cases, such as domestic dogs, we have
been happy to accept such heterogeneous metapopulations as single
species. In other cases (generally when gene flow is a little better be-
haved) taxonomists have accepted that populations are distinct species
that are nonetheless genetically linked metapopulations (an example is
described in Box 2.2.)

b o x 2 . 2 : Reproductive Isolation

Vicariance events leading to prolonged geographical isolation are a major

cause of speciation. However, when an existing population expands to sur-
round a large uninhabitable region, strange partial speciation events can
occur. The standard-bearer for the group of so-called ring species has until
recently been the herring gull (Larus argentatus) complex, which has a circum-
polar distribution in the Northern Hemisphere. However, recent work sug-
gests that this much loved example is not in fact a ring species at all (Liebers
et al. 2004). Thankfully, an understudy to the role exists in the form of the Eur-
asian greenish warbler complex whose range expanded around the margins
of the arid Tibetan Plateau. The flanks of the initial population now overlap on
the northern edge of the plateau where they differ in both plumage and song,
but do not interbreed (Irwin et al. 2005). The western flank is now classified
as Phylloscopus viridanus and the eastern as Phylloscopus plumbeitarsus.

figure 2.1. Speciation by circular overlap. After Helbig (2005).