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The origin of the flightless ratite birds of the southern continents has been debated for over a century. Whether dispersal or vicariance (continental breakup) best explains their origin depends largely on their phylogenetic relationships. No consensus has been reached on this issue despite many morphological and molecular studies. To address this question further we sequenced a 2.8-kb region of mitochondrial DNA containing the ribosomal genes in representative ratites and a tinamou. Phylogenetic analyses indicate that Struthio (Africa) is basal and Rhea (South America) clusters with living Australasian ratites. This phylogeny agrees with transferrin and DNA hybridization studies but not with sequence analyses of some protein-coding genes. These results also require reevaluation of the phylogenetic position of the extinct moas of New Zealand. We propose a new hypothesis for the origin of ratites that combines elements of dispersal and vicariance.

Old World starlings have been thought to be related to crows and their allies, to weaverbirds, or to New World troupials. New World mockingbirds and thrashers have usually been placed near the thrushes and/or wrens. DNA-DNA hybridization data indicated that starlings and mockingbirds are more closely related to each other than either is to any other living taxon. Some avian systematists doubted this conclusion. Therefore, a more extensive DNA hybridization study was conducted, and a successful search was made for other evidence of the relationship between starlings and mockingbirds. The results support our original conclusion that the two groups diverged from a common ancestor in the late Oligocene or early Miocene, about 23-28 million years ago, and that their relationship may be expressed in our passerine classification, based on DNA comparisons, by placing them as sister tribes in the Family Sturnidae, Superfamily Turdoidea, Parvorder Muscicapae, Suborder Passeres. Their next nearest relatives are the members of the Turdidae, including the typical thrushes, erithacine chats, and muscicapine flycatchers.

We present a classification of the living birds of the world based on the results of DNA-DNA hybridization studies. Several family-group names are presented formally for the first time; some of these names have appeared earlier in various publications.

Twenty-two species of Hawaiian honeycreepers (Fringillidae: Carduelinae: Drepaninini) are known. Their relationships to other groups of passerines were examined by comparing the single-copy DNA sequences of the Apapane (Himatione sanguinea) with those of 5 species of cardueline finches, 1 species of Fringilla, 15 species of New World nine-primaried oscines (Cardinalini, Emberizini, Thraupini, Parulini, Icterini), and members of 6 other families of oscines (Turdidae, Monarchidae, Dicaeidae, Sylviidae, Vireonidae, Corvidae). The DNA-DNA hybridization data support other evidence indicating that the Hawaiian honeycreepers shared a more recent common ancestor with the cardueline finches than with any of the other groups studied and indicate that this divergence occurred in the mid-Miocene, 15-20 million yr ago.The colonization of the Hawaiian Islands by the ancestral species that radiated to produce the Hawaiian honeycreepers could have occurred at any time between 20 and 5 million yr ago. Because the honeycreepers captured so many ecological niches, however, it seems likely that their ancestor was the first passerine to become established in the islands and that it arrived there at the time of, or soon after, its separation from the cardueline lineage. If so, this colonist arrived before the present islands from Hawaii to French Frigate Shoal were formed by the volcanic "hot-spot" now under the island of Hawaii. Therefore, the ancestral drepaninine may have colonized one or more of the older Hawaiian Islands and/or Emperor Seamounts, which also were formed over the "hot-spot" and which reached their present positions as the result of tectonic crustal movement. The cardueline-drepaninine lineage probably diverged from the Fringilla lineage in the late Oligocene, from the New World nine-primaried oscines in the early Oligocene, and from the other oscines in the early Eocene.During this study we also obtained evidence that the vireos (Vireonidae) are not closely related to the New World nine-primaried oscines.

of Hubby and Lewontin (1966) and Lewontin and Hubby (1966), there has been considerable interest in determining the percentage of the genome that may be either heterozygous in individuals or polymorphic in populations. Comparable data for natural populations of many species, both vertebrates and invertebrates being represented, are now available. However, the extent of protein polymorphism and individual heterozygosity in natural avian populations is known for relatively few species (Bush 1967; Sibley and Corbin 1970; Nottebohm and Selander 1972). Several other studies of protein polymorphism in bird populations have dealt with only a few loci (Stratil and Valenta 1966; Brush 1968, 1970; Bush et al. 1970; Ferguson 1971; Brush and Scott 1972). Most of the work on the specific variation of bird proteins involved domestic species. In this category, proteins found to be polymorphic include the serum esterases of the Japanese Quail (Coturnix coturnix) (Kaminski 1964; Manwell and Baker 1969), several species of pigeons (species of Columba and ALAN H. BRUSH

The living species of flamingos form a small, easily defined group of three genera, Phoenicopterus, Phoeniconaius, and Phoenicopawus. Phoenicopterus is usually divided into three forms, considered to be species by some authors while others treat them as races of the Greater Flamingo, Phoenicopterw Tuber. These are P. T. Tuber of the New World tropics and subtropics, P. r. chilen&.s of temperate South America, and P. T. roseus (= antiquorum) of the Old World. The Lesser Flamingo; Phoeniconaius minor, occurs in Africa and India and Phoenicoparrus an&us and P. jamesi are confined to the Andes in Peru, Argentina, and Chile. These four (or six) living species are the survivors of a long and well-documented lineage extending at least to the early Tertiary and possibly into the Cretaceous. In spite of this unusually good fossil record the relationships of the flamingos to other groups of birds have been difficult to determine beyond doubt. A large body of evidence, mainly anatomical, suggests an alliance to the Ciconiiformes (herons, storks, ibises), but the bill and feet, development of the young, the voice, and the mallophagan parasites have been interpreted as indicating an origin from the Anseriformes ( ducks, geese). The question is, are the flamingos most closely related to the herons and storks and merely convergent to the anseriform birds in certain characters or were they derived from the ducks and geese and later converged toward the ciconiiform birds? A third possibility is that they were derived from some other group and are similar to both geese and herons only by convergence. In this paper we will review various opinions about the classification of the flamingos, the fossil history, and the anatomical, behavioral, and other evidence which has been presented. We will then present some new data from our studies of the egg-white proteins and hemoglobins of birds which bear upon this problem.

relationships has led to the study of many characters. In addition to comparative morphology some systematists have utilized ecology, behavior, serology, and biochemistry as clues to the degrees of genetic relatedness among organisms. Because protein molecules are primary gene products it is logical to assume that comparisons among homologous proteins from different organisms should provide useful systematic data. The rationale behind this approach to systematics has been discussed by several authors including Sibley (1960, 1962, 1964, 1965, 1967), Zuckerkandl and Pauling (1965), and Dessauer (1969). The plasma proteins are an obvious choice for investigation because they are easy to collect and because a great deal is known about their properties and functions. Plasma is the fluid portion of blood in which the blood cells are suspended. It is a complex mixture of proteins, carbohydrates, lipids, steroids, and free ions whose composition varies with sex, age, starvation, season, etc. (Moore 1948; Clegg et al. 1951; Vanstone et al. 1955; Dessauer and Fox 1956; Saito 1957b). The protein constituents of plasma, while often quantitatively variable, usually show a high degree of qualitative species specificity when examined by any standard biochemical technique (Morris and Courtice 1955; Zweig and Crenshaw 1957; Drilhon et al. 1958; Woods et al. 1958; Sulya et al. 1961). Some of these studies were based upon serum, the fluid portion of the plasma which is extruded from a blood clot. Plasma thus contains the blood proteins involved in clotting while serum lacks them. Most of the research on plasma proteins has dealt with human material although there has been a great deal of work on other mammalian species and the domestic fowl, Gallus gallus. The major protein components of plasma are albumin, the alpha-, beta-, and gamma-globulins, and various subfractions thereof. The nomenclature of the various components is usually determined by their electrophoretic mobilities with reference to normal human plasma. Thus, albumin is the fastest fraction, alpha-globulin the next fastest and gammaglobulin the slowest. The identification of plasma proteins under different conditions can be difficult (Espinosa 1961; Beaton et al. 1961). Up to 70 different proteins have been found in normal human plasma (Dessauer and Fox 1964) while Baker et al. (1966) found 40 electrophoretic bands in pheasant serum, 14 of which were identifiable. The review by Putnam (1960) provides information on the chemical composition of plasma. In paper electrophoresis at pH 8.6 the fastest c mponent in human plasma is albumin. It has a molecular weight of about 69,000 and is is electric at pH 4.7 (Phelps and Putnam 1960). It may be assumed that the values for other mammals and for birds are similar (Phelps and Putnam 1960:169). Plasma albumin has the same chemical properties as alpha livetin of egg yolk (Williams 1962a) but it has no known specific biological functions (Foster 1960). There are several alpha-globulins, which presumably have different functions. The best known are the hemoglobin-binding haptoglobins and the copper-binding ceruloplasmin. Human haptoglobin has a molecular weight of 85,000 and it is isoelectric at pH 4.1, while ceruloplasmin has a molecular weight of 151,000 and is isoelectric at pH 4.4 (Phelps and Putnam 1960). There are several discrete beta-globulins, most of which have unknown functions. The best known of these are the transferrins. Tra sferrin, also called siderophilin, is an ironbinding protein with a molecular weight of approximately 90,000; it is isoelectric at pH 5.9 (Phelps and Putnam 1960). It has been found that the protein moiety of the transferrin molecule is identical to that of the conalbumin of egg white. They differ only in their 1 Present address: Department of Biology, University of North Carolina at Greensboro, Greensboro, North Carolina 27412.

Since 1951, the alpine habitat of the Black Rosy Finch (Leucosticte tephrocotis atrata) has been visited in many mountain ranges for the purpose of determining the breeding range of this species. The distribution of this species has long been uncertain. Available information is scattered and some of it remains unpublished in the form of museum specimens. This report adds new findings and summarizes existing data on distribution.

In 1858 Baird (Pac. R. R. Repts., vol. 9, p. 336) designated the warbling vireos of the western United States as Vireo swainsonii. The Great Basin form was separated as Vireosylva gilva leucopolia by Oberholser in 1932 (Sci. Publ. Cleveland Mus. Nat. Hist., vol. 4, p. 9). From the mountains of Chihuahua, Ridgway described Vireosylva gilva brewsteri (Proc. Biol. Soc. Wash., vol. 16, 1903, p. 107). Within the past year van Rossem (Trans. San Diego Soc. Nat. Hist., vol. 9, 1940, pp. 77-78) has named Vireo gilvus connectens from the state of Guerrero in southern Mexico, and Sutton and Burleigh (Auk, vol. 57, 1940, pp. 398-400) have described Vireo gilvus eleanorae from Hidalgo in central eastern Mexico. These three races from the mainland of Mexico constitute an integrating series that establishes Ridgway's belief that the leucophrys and amauronotus groups of vireos of Central and South America are conspecific with gilvus. Breeding birds from the Cape Region of Lower California prove to be another link in this chain. Relationship to the southern forms is shown by the admixture of brown in the back, and that to the races of the United States by the almost immaculate underparts. The differences are sufficient to warrant separation as a distinct race.

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Single-copy nuclear DNAs (scnDNAs) of eight species of arvicoline and six species of murine rodents were compared using DNA-DNA hybridization. The branching pattern derived from the DNA comparisons is congruent with the fossil evidence and supported by comparative biochemical, chromosomal, and morphological studies. The recently improved fossil record for these lineages provides seven approximate divergence dates, which were used to calibrate the DNA-hybridization data. The average rate of scnDNA divergence was estimated as 2.5%/Myr. This is approximately 10 times the rate in the hominoid primates. These results agree with previous reports of accelerated DNA evolution in muroid rodents and extend the DNA-DNA hybridization data set of Brownell.

first specimens to be recognized as hybrids were described by Sutton (1938), and the zone of hybridization has been studied by Sibley and Short (1964), Rising (1968, 1969, 1970, 1973), Anderson (1971), and Misra and Short (1974). The specimens collected during the 1950's by Sibley and Short (1964) indicated that the parental forms were behaving biologically as subspecies. As clarified by Short (1969), the parental types were allopatric and separated by a hybrid zone. At that time there was no zone of overlap and hybridization, within which both parental types existed in sympatry and in a combined frequency greater than 5% of the total population. That is, there was no apparent reinforcement of isolating mechanisms. In 1973 the A.O.U. Check-list Committee accepted the recommendation by Sibley and Short (1964) that the western forms (bullockii) be treated as subspecies of I. galbula. This decision was based primarily on birds collected along the Platte River, extending from Greeley, Colorado to Blair, Nebraska. However, Rising (1968, 1970) believed that the "Bullock's Oriole" and "Baltimore Oriole" should not be

The relationships of the Wrentit (Chamaea fasciata) have been debated since its discovery in 1845. The titmice (Parus), babblers (Timaliini), Old World warblers (Sylviini), bushtits (Psaltriparus), wrens (Troglodytidae), and mockingbirds (Mimini) have been suggested as its closest relatives. Many authors have placed the Wrentit in a monotypic family, Chamaeidae. DNA-DNA hy- bridization comparisons between the homologous nucleotide sequences of the single-copy DNAs of the Wrentit and other passerines indicate that Chamaea is most closely related to the babblers and the Old World warblers, and that the latter two groups are ecotypes of a single, varied, monophyletic assemblage. The Wrentit is annroximatelv eauidistant from the sylviines and the timaliines, but ecologically it is a babbler. _ The Wrentit (Chamaea fasciata) occurs in chaparral and other dense, brushy, plant for- mations in western Oregon, California, and northwestern Baja California. The question of its affinities to other passerines has been de- bated since the 1850's. Sibley (1970:57-60) reviewed the taxonomic history of the Wrentit and presented electrophoretic comparisons between its egg white proteins and those of many other oscine taxa. He concluded that the total evidence did not offer a clear choice be- tween the sylviine warblers and the timaliine babblers as the closest relatives of Chamaea and that the question of the "relationships of the wren-tit remains open and further evidence should be presented before the affinities of the wren-tit are considered to have been proved beyond doubt" (p. 60). More recently, Sibley and Ahlquist (1980) concluded that Cha- maea is a babbler. In the present paper we offer additional data showing that the sylviine warblers and timaliine babblers are actually ecotypes of a single varied assemblage and that the Wrentit probably was derived from an Asiatic babbler. METHODS

The Sharpbill (Oxvruncus cristatus) is a Neotropical suboscine passerine whose affinities have been unclear. The tyrant flycatchers, cotingas, and manakins of the superfamily Tyrannoidea have been suggested as the closest relatives of the Sharpbill. DNA-DNA hybridization comparisons between the radioiodine-labeled single-copy DNA of the Sharpbill and the DNAs of cotingas, manakins, tyrant flycatchers, and representatives of the other Neotropical superfamily, the Furnarioidea, indicate that Oxyruncus is a cotinga. We place it in the subfamily Cotinginae, family Tyrannidae. The Sharpbill (Oxyruncus cristatus) is a medium-sized (ca. 16 cm) suboscine passerine with a straight, pointed bill and short rictal bristles. In the adult male the crested crown has a crimson center bordered by olive feathers tipped with black. The back is bright olivegreen, wings and tail blackish, underparts white with black spots, and the flanks pale greenishyellow. The plumage colors of the adult female are less intense than those of the male and the coronal stripe is inconspicuous. The Sharpbill apparently has a discontinuous range in the humid montane forests of Costa Rica, Panama, southeastern Venezuela, southern Guyana and Surinam, northeastern and southeastern Brazil, southern Paraguay, and central Peru (Mees 1974, Traylor, 1979). Extensive field observations of the Sharpbill have not been recorded but Sick (197 1) reviewed the sparse literature and reported on his encounters with the species during his many years of field studies in Brazil. He found that Sharpbills tend to occur most often in dense tall forest, although they sometimes feed on fruits at the forest edge or in solitary trees. Fruits are the principal food but insects have been found in the stomachs of specimens and Sick observed Sharpbills hanging upside down from twigs to capture insect larvae. The vocalizations are relatively simple and similar to those of certain cotingas. Sick found the birds most often in pairs and in mixed flocks of tanagers, furnariids, troupials, cotingas, woodpeckers, etc. feeding on fruiting trees. A captive Sharpbill ate fruits and cast hard pellets composed of the skins. Bangs and Barbour (1922) also observed Sharpbills feeding in fruiting trees with cotingas; Mees (1974) found them solitary, in pairs, and in mixed bird swarms in Surinam. Wetmore (1972:605) saw “four or five” together in Panama and recorded spiders, ants, and seeds from stomach contents. Brooke et al. (1983) found the first recorded nest of the Sharpbill near the top of a 30-m tree in the montane forest of the Serra do Tingua, 50 km north of Rio de Janeiro, Brazil. The nest, which was built by the female, was “saddled onto a slender (ca. 3 cm) horizontal branch” and was a “simple . . . shallow cup . . . composed of roughly interlaced leguminous petioles . . . and a very few dry leaves.” The outer surface “was a thin coat of mosses, liverworts, and spider’s web. . . with a texture akin to thin cardboard, possibly resulting from the application and subsequent drying of saliva.” The female fed the young by regurgitation, as do cotingas, rather than as in insectivorous tyrannids. Brooke et al. (1983) observed adult Sharpbills feeding in the ways described by Sick (1971).