Phenetics

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Phenetics should not be confused with phonetics, the study of speech sounds, despite the similarity in pronunciation.

In biology, phenetics, also known as numerical taxonomy, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary relation.

Phenetics has largely been superseded by cladistics for research into evolutionary relationships among species. However, certain phenetic methods, such as neighbor-joining, have found their way into cladistics, as a reasonable approximation of phylogeny when more advanced methods (such as Bayesian inference) are too computationally expensive.

Phenetic techniques include various forms of clustering and ordination. These are sophisticated ways of reducing the variation displayed by organisms to a manageable level. In practice this means measuring dozens of variables, and then presenting them as two or three dimensional graphs. Much of the technical challenge in phenetics revolves around balancing the loss of information in such a reduction against the ease of interpreting the resulting graphs.

Difference from cladistics

Phenetic analyses do not distinguish between plesiomorphies - traits that are inherited from an ancestor (and therefore phylogenetically uninformative) - and apomorphies - traits that evolved anew in one or several lineages. Consequently, phenetic analyses are liable to be misled by convergent evolution and adaptive radiation.

But the two methodologies need not be mutually exclusive. In general, phenetics is is today recognized to provide little if any information about the evolutionary relationships among taxa. But there is no reason why e.g. species identified using phenetics cannot subsequently be subjected to cladistic analysis, to determine its evolutionary relationships.

Phenetic methods can be superior to cladistics when only the distinctness of related taxa is important, as the computational requirements are lower. On the other hand, whenever information on the evolutionary history of taxa is needed for a study, cladistic methods are used today.

Phenetics today

Traditionally there was a great deal of heated debate between pheneticists and cladists, as both methods were initially proposed to resolve evolutionary relationships. Perhaps the "high-water mark" of phenetics were the DNA-DNA hybridization studies by Charles G. Sibley, Jon E. Ahlquist and Burt L. Monroe Jr., from which resulted the 1990 Sibley-Ahlquist taxonomy for birds. Highly controversial at its time, some of its findings (e.g. the Galloanserae) have been vindicated, while others (e.g. the all-inclusive "Ciconiiformes") have been rejected. However, with computers growing increasing powerful and widespread, more refined cladistic algorithms became available and could put the suggestions of Willi Hennig to the test; as it turned out, the results of cladistic analyses turned out to be superior to those of phenetic methods - at least when it came to resolving phylogenies.

Many systematists continue to use phenetic methods, particularly in addressing species-level questions. While the ultimate goal of taxonomy includes finding the 'tree of life' - the evolutionary path connecting all species - in fieldwork one needs to be able to separate one taxon from another. Classifying diverse groups of closely-related organisms that differ by very subtle differences is difficult using a cladistic approach. Phenetics provides numerical tools for examining overall patterns of variation, allowing researchers to identify discrete groups that can be classified as species.

Modern applications of phenetics are common in botany, and you'll find some examples in most issues of the journal Systematic Botany. Indeed, due to the effects of horizontal gene transfer, polyploid complexes and other peculiarities of plant genomics, phenetic techniques in botany - though less informative altogether - are also less prone to errors compared cladistic analysis of DNA sequences.

In addition, many of the techniques developed by phenetic taxonomists have been adopted and extended by community ecologists, due to a similar need to deal with large amounts of data.

References

Phenetics was developed by many people, but the most influential are Sneath and Sokal. Their book is still the primary reference for this sub-discipline, although it is now somewhat dated and out of print.

• Sneath, P. H. A. & R. R. Sokal. 1973. Numerical taxonomy — The principles and practice of numerical classification. W. H. Freeman, San Francisco. xv + 573 p.

An excellent, recent textbook on numerical techniques used by ecologists and taxonomists is Legendre and Legendre:

• Legendre, Pierre & Louis Legendre. 1998. Numerical ecology. 2nd English edition. Elsevier Science BV, Amsterdam. xv + 853 pages.

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