The genealogical history of organisms, both living and extinct. Phylogeny represents the historical pattern of relationships among organisms which has resulted from the actions of many different evolutionary processes. Phylogenetic relationships are depicted by branching diagrams called cladograms, or phylogenetic trees. Cladograms show relative affinities of groups of organisms called taxa. Such groups of organisms have some genealogical unity, and are given a taxonomic rank such as species, genera, families, or orders. For example, two species of cats—say, the lion (Panthera leo) and the tiger (Panthera tigris)—are more closely related to each other than either is to the gray wolf (Canis latrans). The family including all cats, Felidae, is more closely related to the family including all dogs, Canidae, than either is to the family that includes giraffes, Giraffidae. The lion and tiger, and the Felidae and Canidae, are called sister taxa because of their close relationship relative to the gray wolf, or to the Giraffidae, respectively.
Cladograms thus depict a hierarchy of relationships among a group of taxa (illus. a). Branch points, or nodes, of a cladogram represent hypothetical common ancestors (not specific real ancestors), and the branches connect descendant sister taxa. If the taxa being considered are species, nodes are taken to signify speciation events. The goal of the science of cladistics, or phylogenetic analysis, is to discover these sister-group (cladistic) relationships and to identify what are termed monophyletic groups—two or more taxa postulated to have a single, common origin.
hierarchical pattern of sister-group relationships (those taxa descended from a common ancestor). (b) Relationships are determined by identifying derived characters; in this case the condition of five digits is primitive, whereas the loss of a digit is derived and unites taxa 1 and 2.">
Phylogenetic trees. (a) A tree representing a hierarchical pattern of sister-group relationships (those taxa descended from a common ancestor). (b) Relationships are determined by identifying derived characters; in this case the condition of five digits is primitive, whereas the loss of a digit is derived and unites taxa 1 and 2.
The acceptance of a cladogram depends on the empirical evidence that supports it relative to alternative hypotheses of relationship for those same taxa. Evidence for or against alternative phylogenetic hypotheses comes from the comparative study of the characteristics of those taxa. Similarities and differences are determined by comparison of the anatomical, behavioral, physiological, or molecular [such as deoxyribonucleic acid (DNA) sequences] attributes among the taxa. A statement that two features in two or more taxa are similar and thus constitute a shared character is, in essence, a preliminary hypothesis that they are homologous; that is, the taxa inherited the specific form of the feature from their common ancestor. However, not all similarities are homologs; some are developed independently through convergent or parallel evolution, and although they may be similar in appearance, they had different histories and thus are not really the same feature. In cladistic theory, shared homologous similarities are either primitive (plesiomorphic condition) or derived (apomorphic condition), whereas nonhomologous similarities are termed homoplasies (or sometimes, parallelisms or convergences). This distinction over concepts and terminology is important because only derived characters constitute evidence that groups are actually related.
As evolutionary lineages diversify, some characters will become modified. Examples include the enlargement of forelimbs or the loss of digits on the hand. Thus, during evolution the foot of a mammal might transform from a primitive condition of having five digits to a derived form with only four digits (illus. b). Following branching at node 1, the foot in one lineage undergoes an evolutionary modification involving the loss of a digit (expressed as character state A). A subsequent branching event then produced taxa 1 and 2, which inherited that derived character. The lineage leading to taxon 3, however, retained the primitive condition of five digits (character state a). The presence of the shared derived character, A, is called a synapomorphy, and identifies taxa 1 and 2 as being more closely related to each other than either is to taxon 3. Distinguishing between the primitive and derived conditions of a character within a group of taxa (the ingroup) is usually accomplished by comparisons to groups postulated to have more distant relationships (outgroups). Character states that are present in ingroups but not outgroups are postulated to be derived. Systematists have developed computer programs that attempt to identify shared derived characters (synapomorphies) and, at the same time, use them to construct the best phylogenetic trees for the available data.
Knowledge of phylogenetic relationships provides the basis for classifying organisms. A major task of the science of systematics is to search for monophyletic groups. Some groups, such as birds and mammals, are monophyletic; that is, phylogenetic analysis suggests they are all more closely related to each other than to other vertebrates. However, other traditional groups, such as reptiles, have been demonstrated to be nonmonophyletic (some so-called reptiles, such as dinosaurs and their relatives, are more closely related to birds than they are to other reptiles such as snakes). Classifications based on monophyletic groups are termed natural classifications. Phylogenies are also essential for understanding the distributional history, or biogeography, of organisms. Knowing how organisms are related to one another helps the biogeographer to decipher relationships among areas and to reconstruct the spatial histories of groups and their biotas. See also Animal evolution; Animal systematics; Biogeography.