Resemblance between animal taxa may be due to convergence
rather than to recent common
ancestry. Constraints on biological materials and adaptation to
particular habits or habitats
will produce widespread convergence. How may we distinguish the
two causes of resemblance?
The relationship between convergence and taxonomy is discussed,
demonstrating that the
choice of taxonomic method will itself determine the extent to which
convergence is perceived.
In particular, cladistic analysis based on parsimony will tend to
minimise and thus conceal
convergence: neither the resulting cladogram nor a consistency
index derived from it can be
used to assess the prevalence of convergence. With any taxonomic
system, there can be no
substitute for evaluation of the morphological characters used. Complementary
use of
molecular characters shows promise: we wait further understanding
of constraints in genetic
evolution and of the possibilities of convergence at this level also.
These general principles are illustrated with a range of examples
from within and between
invertebrate phyla: the phylogeny of Cnidaria and Platyhelminthes
cannot be traced with
certainty, but where the fossil record allows clear rooting, as for
the echinoderms and in
particular the echinoids, combination of morphological and molecular methods
has made
much progress. Sub-groups within a phylum, for example opisthobranch molluscs
and the
dipteran Phoridae, may show an uncontested phylogeny, and here studies
have precisely
identified convergence and shown that it may be the commoner cause of resemblance.
Adaptation to exacting environments shown by terrestial and freshwater
nemertines may also
result in a predominance of convergent resemblance.
Traditional grouping of phyla breaks down on re-examination of
supposedly key characters,
such as segmentation, body cavities, germ layers and symmetry, each of
which must have had
multiple origins: nor are developmental stages (especially not
larvae) a reliable guide to
relationships. Demarcation of phyla may be difficult, as with
arthropods, and location of phyla
is even more difficult, due to their early and rapid radiation.
Over-simplified definition of
characters has bedevilled invertebrate classification and
the use of molecular data has not yet
resolved the major controversies.
The question ‘How common is convergence?’ remains
unanswered and may be
unanswerable. Our examples indicate that even the minimum detectable
levels of convergence
are often high, and we conclude that at all levels convergence has
been greatly underestimated.