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Onetouch 4.0 Scanned Documents i7 A CRITIQUE OF CRACRAFT'S CLASSIFICATION OF BIRDS STORRS L. OLSON National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 USA ABSTRACT.•A recently proposed "phylogenetic" classification of birds (Cracraft 1981b) is not constructed according to dadistic principles and contains little information to support most of the taxa proposed in it. That which is presented is frequently misleading or erro- neous. The nomenclature used is inconsistent and ungrammatical. In failing to provide synapomorphies to cluster taxa, in using data that are not presented in a primitive-derived sequence, in citing differences as evidence of nonrelationship, and using convergence to refute phylogenetic hypotheses, Cracraft commits the very methodological transgressions for which he has long criticized other systematists. Received 4 January 1982, accepted 24 April 1982. IN the past decade or so, there has been great are postulated to be strictly monophyletic" controversy over methodology in systematics, (Cracraft 1981b: 682). occasioned by the rise of dadistics or "phylo- Division 1.•^This contains the orders Sphen- genetic systematics." Most of this debate has isciformes, Gaviiformes (including the Podi- taken place outside the field of ornithology, cipediformes and the Cretaceous toothed di- with the principal exception of a continuous vers of the order Hesperornithiformes), series of papers and reviev^^s by Cracraft (e.g. ProceUariiformes, and Pelecaniformes. Cracraft 1971, 1972, 1980, 1981a). Although these have (1981b: 686) states that "These four orders have primarily been discussions of methodology and often been placed near one another, although criticisms of the work of others, Cracraft (1981b) a strong argument for their interrelationships has now put forth a classification of the entire has not yet been made." Not a single synap- Class Aves that permits an examination of how onnorphy is advanced to justify "Division 1" as effectively he has applied his preferred meth- a monophyletic group. odology in actual practice. Division 2.•This contains the paleognathous ratites and tinamous. An increasing amount of CRACRAFT'S "DIVISIONS" evidence, including new data from paleontol- Because Cracraft (1981b; 681) considers that ogy (Houde and Olson 1981), suggests that the "most currently recognized orders and families few chiiracters that can be used to define this [of birds] are probably monophyletic," his group are primitive (plesiomorphous). Cracraft principal innovation would appear to be the (1980, 1981b) has specifically admitted this segregation of the Class Aves into nine major possibility but rationalizes the consequences categories, virhich he terms "Divisions." In- by saying that even if these characters "are deed, it is clear from his abstract that this is to primitive, that in itself does not constitute an be regarded as the most important aspect of his argument against monophyly but merely sig- classificahon. Yet these Divisions are not named nifies that the hypothesis of monophyly is not formally, "because in a number of cases their well corroborated" (Cracraft 1981b: 688). Not status as monophyletic groups is not yet well corroborated at all would be more accurate, be- documented" (Cracraft 1981b: 685). If no doc- cause, in a cladistic phylogeny, monophyletic umentation is to be provided for the focal point taxa are supposed to be defined by shared de- of his classification, then we might fairly ques- rived character states, without which there is tion from the outset why it was presented at no justification for Division 2. To cite neotenic all. Nevertheless, let us examine the basis for features of ratites as shared derived characters, Cracraft's "Divisions," bearing in mind that as Cracraft (1981b: 689) has done, is hardly likely the principal tenet of the cladist school is that to persuade nonomithologists of the mono- "Taxa are clustered on the basis of synapo- phyly of this group. By the same logic, all of morphies (Hennig 1966), and the taxa so formed the various neotenic species of salamanders 733 The Auk 99: 733-739. October 1982 734 STORRS L. OLSON [Auk, Vol. 99 (belonging to each of the eight famihes of Uro- . are placed together because a fossil is said dela) would form a monophyletic group. [emphasis added] to have a head only similar Division 3.•This category includes the Ci- to one taxon and a postcranial skeleton only coniiformes and the Falconiformes, the latter similar to the other taxon. The evidence of including the Strigiformes. One wiU search in comparative morphology and systematics sug- vain for a single synapomorphy that wiU define gests that mosaic evolution does not work as a group that contains both flamingos and owls absolutely as this" (Cracraft 1981b; 696). Thus, as monophyletic within the Class Aves. In- the concrete evidence offered by the fossils is stead, we are told that "The placement of such dismissed, because preconceptions about how disparate orders in the same Division may seem evolution works cannot accommodate it. Cra- unwarranted, and, admittedly, there is no clear craft offers no alternative explanation for the evidence for this .... Their placement in Di- relationships of Presbyornis and agrees with vision 3 is tentative and boldly hypothetical" Olson and Feduccia that a relationship be- (Cracraft 1981b: 690). No synapomorphies are tween Anseriformes and Galliformes "is not mentioned, and thus no evidence is presented well documented" (Cracraft 1981b: 695). that this weird taxon is monophyletic. It is de- Division 5.•In this category Cracraft in- cidedly misleading for Cracraft (1981b: 690) to cludes the Gruiformes, Charadriiformes, and cite Ligon (1967) as supporting the view that Columbiformes. He states that the interrela- the Ciconiiformes and Falconifonnes "may have tionships of these three orders "have not yet a relationship." Ligon argued for a close rela- been resolved satisfactorily" but that "there is tionship only between the Cathartidae and Ci- reason to maintain this association" because coniidae, which he placed in their own order "it stands as a working hypothesis" £md "be- (Ciconiiformes) apart from herons (Ardei- cause no alternative hypothesis of interordinal formes) and hawks (Accipitriformes). This is relationships seems better" (Cracraft 1981b: completely contrary to Cracraft, who regards 697). Once again, no synapomorphies are ad- the Ciconiiformes and Falconiformes each to vanced in support of monophyly of this group. be monophyletic. Division 6.•This includes only the order Division 4.•This contains the Galliformes and Psittaciformes, which most systematists con- Anseriformes. Arguments against a close re- sider to be composed of but a single family, lationship between these two orders have been Psittacidae. Never has it been doubted that the discussed elsewhere (Olson and Feduccia 1980a: parrots constitute a natural group, and it should 2). Practically the only morphological character have been possible for Cracraft to provide syn- that has been at all consistently cited as indi- apomorphies here to demonstrate monophyly, cating a possible relationship between Anser- but none is offered. Cracraft (1981b: 699) ig- iformes and Gcdliformes is the supposed sim- nores the evidence that suggests a possible close ilarity in the large, rounded "basipterygoid" relationship between the Psittaciformes and processes. Olson and Feduccia (1980a: 5) doc- Columbiformes (Sibley and Ahlquist 1972) and, umented that these structures are not really instead, says only that parrots "are so mor- similar between the two groups, that they have phologically distinct from other birds that their a different developmental history, and that they relationships to other groups have remained are not homologous. But Cracreift (1981b: 696) unresolved." Considering that he criticizes simply ignores this and continues to cite ba- other avian systematists for bestow^ing "high sipterygoid morphology as possibly indicating rank on taxa that are niorphologically divergent a relationship between Anseriformes and Gal- compared to their closest relatives" (p. 683), liformes. would it not have been preferable to reflect un- The fossil bird Presbyornis, which is known certainty about the relationships of parrots by from thousands of specimens, has, in essence, considering them incertae sedis, rather than the head of a duck and the body of a charad- raising the family Psittacidae to the same level riiform. It provides strong evidence for a der- as a taxon that contains penguins, loons, pel- ivation of the Anseriformes from the Charad- icans, grebes, and Hesperornis, among others? riiformes (Olson and Feduccia, 1980a). Cracraft, Division 7.•This division also consists of a however, seems to regard Presbyornis as a hy- single order, the Cuculiformes. Cracraft's pothetical creature and finds "it disquieting for (1981b: 699) justification for this taxon is that theoretical reasons , . when two higher taxa "There is reasonably strong evidence for ac- October 1982] Critique of Phyhgenetic Classification 735 cepting a sister-group relationship between the Grues. This deficiency has still not been re- turacos (Musophagidae) and cuculids, but the medied, even though Cracraft (1981b) changed affinities of both families to other birds remain the composition of the taxon radically in the uncertain." What the "reasonably strong evi- meantime. Nor will one find any mention of dence" may be, or if it consists of synapomor- synapomoiphies that define the suborder Ralli, phies, is never stated.
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