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Taxonomy of True Finches (Fringillidae, Passeriformes): a Review of Problems1 V

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Taxonomy of True Finches (Fringillidae, Passeriformes): a Review of Problems1 V ISSN 10623590, Biology Bulletin, 2015, Vol. 42, No. 8, pp. 713–723. © Pleiades Publishing, Inc., 2015. Original Russian Text © V.A. Payevsky, 2015, published in Zoologicheskii Zhurnal, 2015, Vol. 94, No. 2, pp. 221–232. Taxonomy of True Finches (Fringillidae, Passeriformes): a Review of Problems1 V. A. Payevsky Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia email: [email protected] Received February 10, 2014 Abstract—A number of issues in phylogeny and classification of true finches is reviewed. Emphasis is made on poor knowledge of their relationships, in spite of intensive research. The main historic stages of develop ment of true finches classification are reviewed together with the results of molecular studies of their phylog eny. Most recent revisions at the level of subfamilies and genera of true finches are discussed. Keywords: birds, passerines, true finches, Fringillidae, taxonomy, phylogeny, classification DOI: 10.1134/S1062359015080051 1 Avian phylogeny and classification currently state that the limits of the true finch family are uncer remain hotly debated topics. This debate became tain, and that different and often strongly varying especially acute in conjunction with the advent of views on this topic exist (Sharpe, 1888; Sushkin, 1924; molecular methods in taxonomy. The recent reviews Dementiev, 1937; Tordoff, 1954; Stallcup, 1954; Wet of classification of passerines, Passeriformes (Sibley more, 1960; Kartashev, 1974; Sibley and Ahlquist, and Ahlquist, 1990; Koblik et al., 2014; Payevsky, 1990; Sibley and Monroe, 1990; Cramp and Perrins, 2014), which comprise ca. 60% of extant birds (Sibley 1994; Yuri and Mindell, 2002 etc.). In spite of varying and Monroe, 1990), highlighted ongoing difficulties opinions on individual genera and species, based on that taxonomists encounter in defining the family lim both morphological and molecular data, the main dis its in this order. agreements on the limits of taxa have been on the level The family of true finches (Fringillidae) tradition of subfamilies. During the entire history of taxonomy ally includes mainly birds with coneshaped bill, i.e. of true finches, different authors included into or typical seedeaters, with short and stout bill and nine excluded from this family such groups, in whole or functional primary feathers. Structure of palate sur partly, as buntings (Emberizidae), Darwin’s or Gal face with oblong crests, typical of conebilled birds, is apagos finches (Geospizidae), a number of weavers perfected in true finches. This form makes it possible (Ploceidae), in particular true sparrows (genera Passer to shell the seeds by longitudinal movements of sharp Briss., Petronia Kaup, Montifringilla Brehm), cardi mandible edges. True finches are distributed nearly nals (Cardinalidae, or Pyrrhuloxiidae), Hawaiian globally (Eurasia, Africa, North and South America), honeycreepers (Drepanididae), tanagers (Thraupi and their lifestyle is mainly connected with forest veg dae), icterids (Icteridae) and New World warblers etation and scrubland. However, desert and mountain (Parulidae). Consequently, the number of species in ous species also exist. the family could vary from 137 (e.g. Böhme and Flint, Some true finches are well familiar not only to biol 1994) to 993 (Sibley and Ahlquist, 1990). ogists, but to the general public as well. This family includes such species as chaffinches, goldfinches, The situation is further complicated by semantic siskins, bullfinches, crossbills, canaries etc. Many true ambiguities. The English word ‘finch’ and the German finches inhabit parks and gardens close to humans, word ‘Fink’ denote not only true finches, but is used have beautiful ornate plumage and superb vocal quali for more than one thousand species from other fami ties. It is thus somewhat puzzling that in spite of this lies: estrildid finches (Estrildidae), weavers, cardinals, popularity of true finches, their taxonomy and rela Darwin’s finches and some buntings. For this reason, tionships remain poorly studied and in a need of a the English vernacular name for Fringillidae is true revision. finches, and the German authors write of ‘eigentli Most handbooks on birds and taxonomic studies, chen Finken’, i.e. proper finches. At the same time, both from the 19th20th centuries and the recent ones, New World buntings (family Emberizidae) are called American sparrows, which sometimes leads to the 1 The article was translated by the author. ambiguous statement that sparrows are finches. 713 714 PAYEVSKY All that defines the aims of the current contribu anatomic studies in the 19th century, in particular the tion. They are to follow the controversial history of classic studies by E.H. Haeckel and T.H. Huxley, led to research into phylogeny and classification of true a significant progress in the field of avian systematics. finches and on the basis of recent molecular, morpho An important milestone was the Catalogue of Birds in logical, and biogeographic studies to define the main the British Museum by R.B. Sharpe, which diagnosed points of the current taxonomy of this group. families, subfamilies and genera. The first volume was published in 1874, and in the 12th volume (Sharpe, 1888) true finches included 97 genera and were MAIN STAGES OF HISTORY OF TRUE divided into three subfamilies: Fringillinae, Emberiz FINCHES CLASSIFICATION BEFORE inae and Coccothraustinae. However, the author THE PERIOD OF MOLECULAR STUDIES emphasized that no one had proposed a satisfactory In his classification of bird, Aristotle (4th century system of true finches. The subfamilies by Sharpe were B.C.) mentioned both insectivorous and herbivorous criticized by Ridgway (1901) as nonnatural groups. birds, and also true finches. Further attempts to con The latter author divided true finches into 18 groups: struct a system of birds, finches including, were based Fringillae, Loxiae, Pyrrhulae, Coccothrausteae etc. on external characters and similarities in body struc In the classic 19th century works on avian system ture, diet and behavior. They were made in publica atics by M. Fürbringer (1888) and H. Gadow (1893), tions of U. Aldrovandi, P. Belon, J. Ray, F. Willoughby, most passerines were considered closely related due to M. Brisson, G. Cuvier, B. Merrem, C.L. Nitzsch, their morphological similarity. Fürbringer lumped C.J. Temminck, J. Cabanis, C.L. Bonaparte, J.G. Gme them into just two families, but Gadow recognized lin, J.B. Lamarck, G.L. Buffon (the reviews: Karta among songbirds (Oscines) 28 families, true finches shev, 1974; Sibley and Ahlquist, 1990). A new era in among them. In his monograph on the birds of the systematics, based on binary taxonomy and hierarchi world, M. A. Menzbier (1909) considered true finches cal system of taxa, was started by the work of Carolus to be close to buntings, however mentioning signifi Linnaeus. In the 10th edition of his ‘Systema Naturae’ cant difference between these two families in jaw (Linnaeus, 1758) class birds and order passerines structure. Following Sharpe, Menzbier included all (Passeres) among species of the Old and the New true sparrows into true finches, but split hawfinches World include genus Fringilla L. with species like into a separate family Coccothraustidae, which also chaffinch, brambling, goldfinch, linnet, redpoll, siskin included greenfinches and cardinals. and serin, currently F. coelebs L., F. montifringilla L., A significant contribution to the systematics of true Carduelis carduelis (L.), Linaria cannabina (L.), finches was made by the studies of P.P. Sushkin. On the Acanthis flammea (L.), Spinus spinus (L.), Serinus basis of structure of syrinx, bony palate and external canaria (L.), respectively; and genus Loxia L. with morphology of horny palate Sushkin (1924, 1925) species crossbill, greenfinch, bullfinch, hawfinch and divided true finches into three large groups, Cardue pine grosbeak, currently L. curvirostra L., Chloris chlo line, Passerine and Emberizine. Cardueline consist of ris (L.), Pyrrhula pyrrhula (L.), Coccothraustes cocco three lineages with genera: 1—Fringilla L., 2—genera thraustes (L.), Pinicola enucleator (L.), respectively. of cardinals, 3—Carduelis Briss., Carpodacus Kaup After the introduction of a new taxon rank, family, and Coccothraustes Briss. Cardueline are closer related in the early 19th century, the name Fringillidae Leach to PloceoPasserine than to Emberizine, and Hawai 1820 starts to denote true finches. Different classifica ian honeycreepers are particularly close to Cardueline tions of North American birds in 1858–1872 included (Sushkin, 1929). As a result, Sushkin proposed the fol into this family true finches, American buntings lowing taxonomic decisions: superfamily Fringilloidei (= American sparrows), true sparrows and parrotbills consists of families Fringillidae (subfamilies Fringilli (Paradoxornis Gould). Species were distributed across nae, Carduelinae, Cardinalinae), Ploceidae (subfami subfamilies and cohorts mainly on the basis of their lies Passerinae, Viduinae, Ploceinae) and Drepanid external morphology: bill shape, tongue structure, idae, whereas superfamily Emberizoidei consists of number of primaries, form of wings, tail and legs families Icteridae, Coerebidae, Tanagridae and (Kartashev, 1974; Sibley and Ahlquist, 1990). At the Emberizidae. same time, the classification of oscine passerines by By the mid 20th century E. Stresemann (1959) Wallace (1874) was based solely on wing structure, believed that genuine phylogeny could never be stud with true finches
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