Phylogeny of the Waxwings and Allied Birds — Context and Discussion
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Phylogeny of the Waxwings and Allied Birds
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
Phylogeny of the Waxwings and Allied Birds
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
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.
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
M. Dale Arvey's 1951 monograph opens with a clear methodological aim: to distinguish traits shaped by recent habits from those inherited from ancient ancestors. The study focuses on the small passerine family Bombycillidae, comparing coloration, nesting, food habits, skeleton, and soft parts across genera. Arvey's approach is notably systematic, using proportional measurements—such as humerus length relative to thoracic vertebrae—to account for size differences. The text is dense with osteological detail, describing pygostyle shapes, sternal keel depths, and muscle attachment areas, all illustrated with 49 figures and 13 tables. This is a specialist work, written for ornithologists and evolutionary biologists, with a precise, technical voice that prioritizes morphological evidence over speculation.
Comparative Osteology as Phylogenetic Evidence
Arvey devotes substantial space to skeletal comparisons, particularly the pygostyle and sternum. He notes that the pygostyle in Phainoptila is the most generalized, while Bombycilla has a proportionately small but complex shape. The sternum's carina is deepest in Bombycilla garrula, and the manubrium is more extended in that species. In the Ptilogonatinae, the centra of pygostyles offer more area for muscle attachment, correlating with less specialized caudal muscles in Bombycilla. These observations are grounded in measurements and cross-sectional descriptions, such as the "winged" lateral edges of the centrum in Ptilogonys cinereus. Arvey uses these skeletal differences to infer functional and evolutionary divergence, though he does not overstate conclusions from limited material.
Proportional Measurements and Functional Inference
A key analytical tool in this monograph is the use of relative bone lengths expressed as percentages, with the thoracic vertebral column as a stable denominator. Arvey explains that raw lengths can mislead when animals differ in overall size, so ratios provide clues to functional differences. For example, he compares humerus length relative to trunk length across genera, though the excerpt does not reveal the outcomes. This method reflects a careful, quantitative approach typical of mid-century comparative anatomy. The text also includes leg-trunk and arm-trunk percentages, suggesting a focus on locomotion and wing function. Arvey's language is methodical: he describes the sternum's lateral processes flaring outward in adult Ptilogonatines, while in Bombycilla and Dulus they remain closer to the body. Such details hint at adaptive differences in flight or perching.
Voice and Structure: A Technical Monograph
The narrative voice is impersonal and data-driven, with sentences like "The centrum in Ptilogonys shows great width, and the spine is in a large expanded tip as in Bombycilla." Arvey avoids speculation, instead presenting observations and letting the evidence suggest relationships. The structure follows a logical progression: introduction, nomenclatural history, materials, then systematic comparisons of coloration, courtship, nesting, food, skeleton, musculature, and digestive tract. Each section is concise, with frequent references to figures and tables. The prose is dense but clear, aimed at readers familiar with avian anatomy. Notably, the monograph does not include a narrative conclusion in the excerpts, but the summary and bibliography suggest a rigorous synthesis. This is a work of empirical science, not popular natural history.
Readers approaching this monograph should expect a technical, data-rich analysis rather than a broad survey of waxwing natural history. Arvey's evidence is drawn from museum specimens and published literature, with careful attention to measurement and comparison. The 49 figures and 13 tables are integral to the argument, so consulting them alongside the text is essential. This work is best suited for those with a background in vertebrate anatomy or ornithology, offering a detailed snapshot of mid-20th-century phylogenetic methodology.
Reading Arvey’s meticulous bone sketches, I remembered my grandfather’s field journal—same patient penciling of wing feathers, same quiet awe at how structure reveals kinship. Wild Spain (España agreste) Records of Sport with Rifle, Rod, and Gun, Natural History Exploration — Inside the Classic once handed me that identical hush: a man kneeling in reeds, counting vertebrae in a heron’s throat, knowing each one mattered.
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