Phylogeny of the Waxwings and Allied Birds

Part 1

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Phylogeny of the Waxwings and Allied Birds

M. DALE ARVEY

University of Kansas Publications Museum of Natural History

Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables October 10, 1951

UNIVERSITY OF KANSAS LAWRENCE 1951

UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: E. Raymond Hall, Chairman, Edward H. Taylor, A. Byron Leonard, Robert W. Wilson

Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables Published October 10, 1951

University of Kansas Lawrence, Kansas

PRINTED BY FERD VOILAND, JR., STATE PRINTER TOPEKA, KANSAS 1950 [Illustration: union label] 23-1019

Phylogeny of the Waxwings and Allied Birds

by M. DALE ARVEY

CONTENTS

PAGE Introduction 476 Acknowledgments 476 Nomenclatural History 477 Materials 478 Diagnoses 478 Coloration 485 Courtship 489 Nest Building 491 Food 493 Skeleton 494 Skull 494 Humerus 499 Pygostyle 502 Sternum 505 Relative Lengths of Bones 505 Leg-trunk Percentages 509 Arm-trunk Percentages 511 Musculature 514 Caudal Muscles 514 Pectoral Muscles 517 Hind Limb Musculature 517 Digestive Tract 517 Origin of the Species 519 Conclusions 521 Summary 524 Bibliography 525

INTRODUCTION

A small family of passerine birds, the Bombycillidae, has been selected for analysis in the present paper. By comparative study of coloration, nesting, food habits, skeleton and soft parts, an attempt is made to determine which of the differences and similarities between species are the result of habits within relatively recent geological time, and which differences are the result of inheritance from ancient ancestral stocks, which were in the distant past morphologically different. On the basis of this information, an attempt is made to ascertain the natural relationships of these birds. Previous workers have assigned waxwings alone to the family Bombycillidae, and a question to be determined in the present study is whether or not additional kinds of birds should be included in the family.

It has generally been assumed that the nomadic waxwings originated under boreal conditions, in their present breeding range, and that they did not undergo much adaptive radiation but remained genetically homogeneous. Also it is assumed that the species were wide ranging and thus did not become isolated geographically to the extent that, say, the Fringillidae did. The assumption that waxwings originated in the northern part of North America or Eurasia may be correct, but it is more probable that the origin was more southerly, perhaps, in northern Mexico, of North America (see p. 519.) Subsequent to the differentiation of this stock in the south, there was a northerly movement, while certain populations remained behind and underwent an evolution different from the northern group. Since the fossil record does not permit us to say when in geological time the family originated, we must rely on anatomical evidence and the distributional evidence of present-day species to estimate when the family stock had diverged from some unknown group sufficiently to merit the status of a separate family.

ACKNOWLEDGMENTS

It is with pleasure that I acknowledge the guidance received in this study from Professor E. Raymond Hall of the University of Kansas. I am indebted also to Dr. Herbert Friedmann of the United States National Museum for the loan of certain skins, skeletons, and alcoholic material; to Mr. Alexander Skutch, for notes on certain Central American birds; and to Dr. Henry W. Setzer, Mr. George H. Lowery, Jr., Mr. Victor E. Jones, Mr. Victor Housholder, Mr. Alvaro Wille-Trejos, and Mr. Morton F. Davis, for gifts of specimens that have been used in this work. Suggestions and critical comments from Professors Worthie H. Horr, Charles G. Sibley and Edward H. Taylor are gratefully acknowledged. I wish also to thank Mrs. Virginia Unruh for the preparation of the drawings used in this work.

NOMENCLATURAL HISTORY

The oldest name available for any species of the waxwings is Lanius garrulus Linnaeus (1758). Lanius garrulus and Lanius garrulus variety B carolinensis were described as conspecific. The description has been associated with the first of the two names. The latter name is a nomen nudum since it was not accompanied by a separate description. The generic name Lanius was originally applied to both shrikes and waxwings by Linnaeus. Since that name is applied to the shrikes only, the next available generic name that may be applied to the generically different waxwings must be used. This is Bombycilla, a name originally proposed by Brisson (1760) for the Cedar Waxwing. In the 12th Edition of the Systemae Naturae (1766) Gmelin proposed the generic name Ampelis for the Bohemian Waxwing, and combined it with the specific name garrulus, the Cedar Waxwing being termed variety B. Vieillot (1807) proposed the generic name Bombycilla and combined it with a new specific name, cedrorum, for the Cedar Waxwing. Vieillot has been cited as the author of Bombycilla since that time, although Brisson used Bombycilla 33 years before. Oberholser (1917) did not cite Brisson's work in his discussion of the proper generic name for the waxwings, and Bombycilla should be ascribed to Brisson and not Vieillot, since Opinion 37, rendered by the International Zoological Committee on Nomenclature, states that generic names used by Brisson (1760) are valid under the Code. In consequence, the specific name available for the Cedar Waxwing, since Brisson is ruled not to be a binomialist, is Bombycilla cedrorum Vieillot (1807).

Most workers prior to 1900 utilized the family name Ampelidae to include waxwings, silky flycatchers, and palm-chats. Ridgway (1904:113) elevated the silky flycatchers to family rank under the name Ptilogonatidae, and assigned the palm-chats to a separate family, the Dulidae.

MATERIALS

The following specimens, numbering 238, and representing each currently recognized species and subspecies, were used in the study, and were supplemented by observation in 1947 on specimens in the United States National Museum.

==================================================================== Species or Subspecies | Skin | Skeleton| Alcoholic ----------------------------------------+------+---------+---------- Phainoptila melanoxantha melanoxantha | 8 | 1 | 2 Phainoptila melanoxantha minor | 2 | | Ptilogonys cinereus cinereus | 13 | 3 | 4 Ptilogonys cinereus molybdophanes | 6 | | Ptilogonys caudatus | 16 | 3 | 4 Phainopepla nitens nitens | | 1 | 5 Phainopepla nitens lepida | 12 | 5 | 4 Bombycilla cedrorum | 53 | 27 | 8 Bombycilla garrula garrula | 4 | 3 | Bombycilla garrula centralasiae | 9 | 2 | Bombycilla garrula pallidiceps | 7 | 3 | 2 Bombycilla japonica | 10 | | Dulus dominicus dominicus | 9 | 5 | 2 Dulus dominicus oviedo | 4 | 1 | |--------------------------- Totals | 153 | 54 | 31 --------------------------------------------------------------------

DIAGNOSES

Family Bombycillidae

Part 2

Diagnosis.--Bill short, flat, somewhat obtuse, minutely notched near tip of each maxilla, flared at base; gape wide and deeply cleft; culmen convex; nasal fossa broad, exposed, or filled with short, erect or antrorse, close-set velvety feathers; nostril narrowly elliptical; rictal vibrissae long, short, or absent; lacrimal bone free, articulating at two points; wings long and pointed, or short and rounded; primaries ten, tenth reduced in some species; tail short, narrow, even, two thirds or less length of wing, or much longer and forked or rounded; feet weak (except in Dulus and Phainoptila); tarsus generally shorter than middle toe and claw, distinctly scutellate with five or six divisions, the lateral plate subdivided (except in Phainoptila); lateral toes of nearly equal length; hallux approximately as long as inner lateral toe, or shorter; basal phalanx of middle toe more or less united to that of outer and inner toes; body stout; head generally conspicuously crested; plumage soft, smooth and silky (except in Dulus); eggs spotted; nest in trees; three subfamilies, five genera, eight species.

Subfamily Ptilogonatinae

Diagnosis.--Rictus with conspicuous bristles; nasal fossa almost entirely exposed; tail long and rounded, graduated, or square; caudal muscles and pygostyle well developed; wings rounded and short, first primary a half to a third as long as second; second primary shorter than third; humerus long, with small external condyle; plumage soft and silky, less so in Phainoptila; sexes dissimilar, young like adult female; three genera, four species.

Genus =Phainoptila= Salvin

Phainoptila Salvin, Proc. Zool. Soc. London, 1877:367, April 17, 1877. Type Phainoptila melanoxantha Salvin.

Diagnosis.--Without crest; tarsus longer than middle toe and claw, and booted or very slightly reticulate; tail shorter than wing, rounded; nostril exposed, ovate; rictal bristles distinct; first primary well developed; plumage normal, bill flared slightly at base.

Range.--Costa Rica and Panama.

=Phainoptila melanoxantha melanoxantha= Salvin

Phainoptila

Phainoptila melanoxantha melanoxantha Salvin, Proc. Zool. Soc. London, 1877:367; April 17, 1877.

Diagnosis.--Coloration of adult males: Pileum, hindneck, back, scapulars, and upper tail coverts Black (capitalized color terms after Ridgway, Color Standards and Color Nomenclature, Washington, D. C., 1912), with Bluish Gray-Green gloss; rump Lemon Yellow tinged with Olive; lower breast and abdomen Gull Gray or Slate Gray; sides and flanks clear Lemon Yellow; lower chest, upper breast, and under tail coverts Yellowish Olive-Green, extending to patch on sides and flanks of same color; bill and feet Black or Blackish Brown. Coloration of adult females: Most of upper parts Olive-Green, with Yellowish Olive on rump; thighs Olive-Gray, as are sides of head; rest of coloration as in male. Coloration of young: As in adult female, but duller throughout.

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