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diff --git a/34556-8.txt b/34556-8.txt new file mode 100644 index 0000000..1a4512e --- /dev/null +++ b/34556-8.txt @@ -0,0 +1,3313 @@ +The Project Gutenberg EBook of Phylogeny of the Waxwings and Allied Birds, by +M. Dale Arvey + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Phylogeny of the Waxwings and Allied Birds + +Author: M. Dale Arvey + +Release Date: December 3, 2010 [EBook #34556] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK PHYLOGENY OF THE WAXWINGS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper, The +Internet Archive for some images and the Online Distributed +Proofreading Team at https://www.pgdp.net + + + + + + + + + + Phylogeny of the Waxwings + and Allied Birds + + + BY + + 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 Zoölogical 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 + +_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. Zoöl. 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 Panamá. + + +=Phainoptila melanoxantha melanoxantha= Salvin + +Phainoptila + + _Phainoptila melanoxantha melanoxantha_ Salvin, Proc. Zoöl. 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. + +_Measurements._--Wing 99.0, tail 88.5, culmen 15.2, tarsus 28.4. + +_Range._--Highlands of Costa Rica and extreme western Panamá (Volcán +de Chiriquí). + + +=Phainoptila melanoxantha minor= Griscom + +Phainoptila + + _Phainoptila melanoxantha minor_ Griscom, Amer. Mus. Novitates, + 141:7, 1924. + +_Diagnosis._--Coloration as in _P. m. melanoxantha_, but female with +hindneck more extensively gray and of slightly darker shade; rump, +upper tail coverts, and edgings to tail feathers slightly greener, +less yellow; average size smaller than in _P. m. melanoxantha_. + +_Range._--Highlands of westeran Panamá (Cerro Flores and eastern +Chiriquí). + + +Genus =Ptilogonys= Swainson + + _Ptilogonys_ Swainson, Cat. Bullock's Mex. Mus., App. 4, 1824. + Type _Ptilogonys cinereus_ Swainson. + +_Diagnosis._--Tail much longer than wing, even or graduated; head with +bushy crest; nostril large, rounded and fully exposed, bordered by +membrane; rictal bristles well developed; tarsus shorter than middle +toe with claw; plumage soft, blended. + +_Range._--Southwestern United States to Costa Rica. + + +=Ptilogonys cinereus cinereus= Swainson + +Ashy Ptilogonys + + _Ptilogonys cinereus cinereus_ Swainson, Cat. Bullock's Mex. Mus., + App. 4, 1824. + +_Diagnosis._--Coloration of adult male: Frontals, supralorals, malars, +and chin White; orbital ring White; auriculars and nape grayish brown; +rest of head smoke gray; back, scapulars, wing coverts, rump, and +upper tail coverts plain Bluish Black; rectrices (except middle pair) +with large patch of White midway between base and tip, rest plain +Bluish Black; chest, breast, and anterior parts of sides plain Bluish +Gray-Green, much lighter than back, and fading into paler Gray on +throat; abdomen and thighs White; flanks and posterior part of sides +Olive-Yellow or Yellowish Olive; under tail coverts Lemon Yellow; +bill, legs and feet Black. Coloration of adult females: Head plain +Smoke Gray, passing into White on frontals, malars, and chin; back, +scapulars, wing coverts, and rump Hair Brown; upper tail coverts Dark +Gull Gray; remiges and rectrices Black with faint Dusky Green gloss, +edged with Gull Gray; chest Dark Grayish Brown lightening to Wood +Brown on sides and flanks; abdomen White; under tail coverts Yellow +Ocher. Coloration of young: As in adult female, but paler throughout. + +_Measurements._--In adult male, wing 94.0, and tail 104.2; in adult +female, wing 93.3, and tail 94.8; both sexes, culmen 11.1, and tarsus +18.7. + +_Range._--Mountainous districts of central and southern Mexico, in +states of Durango, Zacatecas, Hidalgo, México, Oaxaca, Colima, +Morelos, Veracruz, San Luís Potosi, Guerrero and Michoacán. + + +=Ptilogonys cinereus molybdophanes= Ridgway + +Ashy Ptilogonys + + _Ptilogonys cinereus molybdophanes_ Ridgway, Man. N. American Birds, + 464 (footnote), 1887. + +_Diagnosis._--Coloration of adult male: Upper parts darker bluish than +in _P. c. cinereus_; venter paler; flanks Olive-Green rather than +Olive as in _P. c. cinereus_. Coloration of adult female: Like female +of _P. c. cinereus_ but colors darker throughout; dorsum more +olivaceous. + +_Measurements._--In adult male, wing 89.4, and tail 97.1; in adult +female, wing 89.4, and tail 93.3; both sexes, culmen 11.7, and tarsus +17.3. + +_Range._--Western Guatemala, in subtropical and temperate zones. + + +=Ptilogonys caudatus= Cabanis + +Costa Rican Ptilogonys + + _Ptilogonys caudatus_ Cabanis, Jour. für Orn., 1866:402, Nov. 1866. + +_Diagnosis._--Coloration of adult male: Forehead and crown Pale +Grayish Blue, slightly paler anteriorly; orbital ring Lemon Yellow; +rest of head and neck, including crest, Olive-Yellow; throat paler and +tinged with Light Gull Gray; back, scapulars, rump, upper tail coverts +and wing coverts uniform Bluish Slate-Black; chest and breast similar +but paler; sides and flanks Yellowish Olive-Green; thighs, lower +abdomen, and under tail coverts Lemon Yellow; remiges, primary coverts, +and tail Black, glossed with Bluish Black and edged with Gull Gray; +inner webs of rectrices (except two middle pair) with large middle +patch of White; bill, legs, and feet Black. Coloration of adult +female: Forehead and crown Pale Gull Gray, becoming paler anteriorly; +rest of head, together with neck, back, scapulars, rump, and wing +coverts plain Yellowish Olive Green; chest and breast similar but more +grayish; lower abdomen and flanks White tinged with Yellowish Olive; +under tail coverts Olive-Gray; remiges, primary coverts, and rectrices +Black with Gull Gray edges. Coloration of young: Dorsum plain Light +Grayish Olive; upper tail coverts Brownish Olive; underparts Grayish +Olive anteriorly, becoming more Yellowish Olive on abdomen; under tail +coverts pale Yellowish Olive with Grayish Olive base; bill and feet +Brownish Drab. + +_Measurements_--In adult male, wing 96.2, and tail 135.7; in adult +female, wing 93.9, and tail 113.7; both sexes, culmen 12.6, and tarsus +19.1. + +_Range._--Highlands of Costa Rica and extreme western Panamá. + + +Genus =Phainopepla= Sclater + + _Phainopepla_ Sclater, Proc. Zoöl. Soc. London, 26:543, 1858. Type + _Phainopepla nitens_ (Swainson). + +_Diagnosis._--Tail almost as long as wing; head with pointed crest of +narrow, separated feathers; rectrices without white; bill narrow, +compressed terminally; conspicuous white patch under wing; nostril +small, exposed; rictal bristles distinct; tail slightly rounded. + + +=Phainopepla nitens nitens= (Swainson) + +Phainopepla + + _Phainopepla nitens nitens_ (Swainson), Anim. in Menag., 1838:285, + Dec. 31, 1837. + +_Diagnosis._--Coloration of adult male: Uniform glossy Bluish Black; +inner webs of primaries except innermost pair with middle portion +White; bill, legs, and feet Black. Coloration of adult female: Plain +Olivaceous Black, longer feathers of crest Black, edged with Gull +Gray; remiges and rectrices Dusky Drab to Black; rectrices and coverts +margined by White; bill and feet Brownish Drab to Dusky Brown. +Coloration of young: Like adult female but more Brownish Drab. + +_Measurements._--No specimens examined; larger than _P. n. lepida_ +(Van Tyne, 1925). + +_Range._--Central and southern Mexico, in states of Coahuila, San Luís +Potosi, Durango, Guanajuato, México, Puebla, and Veracruz. + + +=Phainopepla nitens lepida= Van Tyne + +Phainopepla + + _Phainopepla nitens lepida_ Van Tyne, Occ. Pap. Bost. Soc. Nat. + Hist., 5:149, 1925. + +_Diagnosis._--Coloration same as _P. n. nitens_; separated by smaller +size. + +_Measurements._--Wing 91.0, tail 90.3, culmen 11.5, tarsus 17.6. + +_Range._--Southwestern United States, from central California, +southern Utah, and central western Texas southward to Cape San Lucas +in Baja California, and into northwestern Mexico (Sonora and +Chihuahua). + + +Subfamily =Bombycillinae= + +_Diagnosis._--Wings long and pointed, reaching almost to tip of tail; +first primary spurious; second primary longest; tail short and even; +rictal vibrissae few and short; secondaries generally, and sometimes +also rectrices, tipped with red, corneous appendages; nasal fossa +partly filled with short, antrorse, close-set velvety feathers; +plumage soft, silky; tail tipped with yellow band (red in _B. +japonica_); sexes alike; humerus short with large external condyle; +caudal muscles and pygostyle not well developed; bill flared widely at +base; one genus, three species. + +_Range of subfamily._--Holarctic breeding area; wanders nomadically +south in winter to Central America and West Indies, southern Europe +and Asia. + + +Genus =Bombycilla= Brisson + + _Bombycilla_ Brisson, Orn. ii, 1760:337. Type _Bombycilla garrula_ + (Linnaeus). + +_Diagnosis._--As described for the subfamily. + + +=Bombycilla cedrorum= Vieillot + +Cedar Waxwing + + _Bombycilla cedrorum_ Vieillot, Hist. Nat. Amer., 1:88, Sept. 1, 1807 + +_Diagnosis._--Coloration of adults: Shading from Saccardo's Umber on +dorsum to Bister on top of head; upper tail coverts and proximal +rectrices Gull Gray; underparts shade through pale Lemon Yellow wash +on belly into White on under tail coverts; forehead, lores, and +eye-stripe Black; chin same, soon shading into Blackish Mouse Gray and +into color of breast; side of under jaw with sharp White line; narrow +line bordering forehead, and lores, White; lower eyelid White; quills +of remiges Dark Mouse Gray, darkening at tips; inner quills tipped +with red horny wax appendages; tail feathers like primaries, but +tipped with Lemon Yellow, and occasionally showing also red horny wax +appendages; bill and feet Black. Coloration of young: Dorsum as in +adult, but lightly streaked with White; head concolor with dorsum; +forehead White; lores Black; eye stripe Black anterior to eye and +White posterior to eye; throat Light Buff; belly with alternate +streaks of Dresden Brown and light Ochraceous Buff but posteriorly +White; tail tipped with Lemon Yellow bar; bill black at tip, shading +to Sepia at base. + +_Measurements._--Wing 92.9, tail 55.5, culmen 10.9, tarsus 16.8. + +_Range._--Breeds from central British Columbia, central Alberta and +Manitoba, northern Ontario, southern Quebec and Cape Breton Island +south to northwestern California, northern New Mexico, Kansas, +northern Arkansas, North Carolina, and northern Georgia. Winters south +to Louisiana, Mississippi, Texas, Arizona, Colorado, Florida, +Honduras, Costa Rica, Jamaica, Little Cayman Island, Haiti, and +Panamá. + + +=Bombycilla garrula= (Linnaeus) + +Bohemian Waxwing + + _Bombycilla garrula_ (Linnaeus), Syst. Nat., 10th Ed., 1758:55. + +_Diagnosis._--Coloration of adults: General color Olive-Brown, shading +insensibly from clear Smoke Gray of upper tail coverts and rump to +Cinnamon-Drab anteriorly, heightening on head and forehead to Hazel; +narrow frontal line, lores, broader mask through eye, chin, and upper +throat, Sooty Black; under tail-coverts Cinnamon-Brown; tail Smoke +Gray, deepening to Blackish Mouse Gray distally, and tipped with Lemon +Yellow; wings Blackish Mouse Gray; primaries tipped with sharp spaces +of Lemon Yellow or White, or both; secondaries with White spaces at +ends of outer web, shafts usually ending with enlarged, horny red +appendages; primary coverts tipped with White; bill Blackish Slate and +paler at base; feet Black. Coloration of young: Much like adult, but +general color duller; some streaking on venter and back; chin, throat, +and malar region dull White. Three subspecies. + + +=Bombycilla garrula garrula= (Linnaeus) + +Bohemian Waxwing + + _Bombycilla garrula garrula_ (Linnaeus), Syst. Nat., 10th Ed., + 1758:55. + +_Diagnosis._--Coloration: As described for the species, but darkest of +the three subspecies; tending to be more Vinaceous dorsally than +either _pallidiceps_ or _centralasiae_. + +_Measurements._--Wing 113.5, tail 63.1, culmen 12.5, tarsus 20.7. + +_Range._--Europe; breeds north to northern Russia and Norway, south to +about 65° N latitude; winters south to England and Ireland, southern +France, northern Italy, and Turkey. + + +=Bombycilla garrula centralasiae= Poljakov + +Bohemian Waxwing + + _Bombycilla garrula centralasiae_ Poljakov, Mess. Orn. vi:137, 1915. + +_Diagnosis._--Coloration: As described for the subspecies _garrula_, +but less Vinaceous dorsally, and more Cinnamon; venter lighter gray +than _garrula_, and much paler than _pallidiceps_. + +_Measurements._--Wing 114.7, tail 63.0, culmen 12.2, tarsus 21.0. + +_Range._--Asia; breeds northern Siberia south to Vladivostok; winters +to Turkestan and central eastern China and Japan. + + +=Bombycilla garrula pallidiceps= Reichenow + +Bohemian Waxwing + + _Bombycilla garrula pallidiceps_ Reichenow, Orn. Monats. 16:191, 1908. + +_Diagnosis._--Coloration: As described for the species, but more +grayish above and below than _B. g. garrula_; darker gray than in +_centralasiae_. + +_Measurements._--Wing 115.1, tail 71.7, culmen 12.6, tarsus 21.1. + +_Range._--Breeds from western Alaska to northern Mackenzie and +northwestern Manitoba south to southern British Columbia, southern +Alberta, northern Idaho, and possibly Colorado (Bergtold 1924) and +Montana (Burleigh 1929); winters east to Nova Scotia and irregularly +over much of Canada, and south irregularly to Pennsylvania, Ohio, +Michigan, Indiana, Kansas, Colorado, California, Arizona, and Texas. + + +=Bombycilla japonica= (Siebold) + +Japanese Waxwing + + _Bombycilla japonica_ (Siebold), Nat. Hist. Jap., St. No. 2:87, 1824. + +_Diagnosis._--Coloration: Dorsum generally Brownish Drab shading to +Light Brownish Drab on lower back, rump, and upper tail coverts; +secondary and tertiary coverts Pale Brownish Drab, washed on outer web +with Carmine; primary coverts Blackish Slate, with White edging; tail +feathers Slate-Gray, broadly tipped with Carmine, bordered anteriorly +by subterminal Black bar; head crested, forehead Chestnut; lores, +frontals, and stripe extending around eye and nape, Black; throat +Black, narrowing on lower throat; breast, sides of flanks Light Drab; +venter pale Sulphur Yellow; thighs Brownish Drab; under tail coverts +Carmine; bill, legs, and feet Black. + +_Measurements._--Wing 108.3, tail 53.6, culmen 11.2, tarsus 19.4. + +_Range._--Breeds eastern Siberia, northern China; winters south in +China, and to Japan (Hokkaido, Kyushu), Taiwan, and Korea. + + +Subfamily _Dulinae_ + +_Diagnosis._--Bill deep and compressed, culmen strongly depressed; +nostrils circular, wholly exposed; tail even, and shorter than wing; +tenth primary less than half length of ninth; under parts streaked; +plumage hard and harsh; rictal bristles minute; wing rounded; humerus +long and with small external condyle; pygostyle and caudal muscles not +well developed; one genus, one species. + +_Range of subfamily._--Islands of Haiti and Gonave, Greater Antilles. + + +Genus _Dulus_ Vieillot + + _Dulus_ Vieillot, Analyse, 1816:42. + +_Diagnosis._--Like the subfamily. + + +=Dulus dominicus dominicus= (Linnaeus) + +Palm-chat + + _Dulus dominicus dominicus_ (Linnaeus), Syst. Nat., 12th Ed., + 1766:316. + +_Diagnosis._--Coloration: Dorsum Olive, back, scapulars, and wing +coverts more Brownish Olive; lower rump and upper tail coverts +Olive-Green; pileum and hindneck with indistinct streaks of Brownish +Olive; tail Brownish Drab, edged with Light Olive Gray; lores, +suborbital region, and auricular regions Dusky Brown; malars Dusky +Brown and streaked with Sooty Black, streaks narrower on abdomen, +broader and paler on under tail coverts, bill Light Brownish Drab; +legs and feet Brownish Drab. + +_Measurements._--Wing 85.0, tail 68.8, culmen 15.0, tarsus 24.7. + +_Range._--Island of Haiti, Greater Antilles. + + +=Dulus dominicus oviedo= Wetmore + +Palm-chat + + _Dulus dominicus oviedo_ Wetmore, Proc. Biol. Soc. Wash., 42:117, + 1929. + +_Diagnosis._--Coloration: Like _D. d. dominicus_, but averaging more +Grayish Olive; rump and tail coverts with less greenish wash. + +_Measurements._--Wing 90.1, tail 71.3, culmen 16.2, tarsus 25.1. + +_Range._--Gonave Island, off Haiti, Greater Antilles. + + + + +COLORATION + + +The general coloration of waxwings is cryptic, that is to say, +concealing or blending. The lighter color of the venter, especially of +the belly, contrasts with the duller, darker vinaceous color of the +dorsum. Several ruptive marks tend to obliterate the outline of the +body. The crest of the head, when elevated, tends to elongate the +body, making the outline less like that of a normal bird. The facial +mask effectively breaks up the outline of the head, and conceals the +bright eye, which would otherwise be strikingly distinct. The white +spots on the distal ends of the secondaries of _B. garrula_ and the +yellow color on the distal ends of the rectrices (red in _B. +japonica_) are also ruptive. These ruptive marks on an otherwise +blending type of plumage might be important to waxwings, and probably +are more effective when the birds remain motionless in either a +well-lighted area or in one that is partly in shadow, rather than in +one that is wholly in shadow. + +The red wax tips on the secondaries of the flight feathers, and +sometimes found on the ends of the rectrices in _Bombycilla_, are +puzzling and no wholly convincing reason has been suggested for their +occurrence. Two instances are known of yellow instead of red-colored +wax tips in _B. cedrorum_ (Farley, 1924). It is well known that many +individuals, especially of _B. cedrorum_, do not possess these tips; +they are absent in a smaller proportion of individuals of _B. +garrula_. Of the 53 skins of _B. cedrorum_ available in the University +of Kansas Museum of Natural History, which might be taken as a +sampling at random of the general population of this species, only 17 +possess wax tips. A few specimens are unilateral, and the tips are of +varying sizes in different individuals. Of these 17 birds, 6 are +female and 7 male, the others being unsexed at the time of skinning. +This proportion is, roughly, half and half. Of the seven skins of _B. +garrula pallidiceps_ in the same Museum, five possess the tips, and +two that are females have no trace of the red tips at all. Of the five +which do have the tips, two are males, two are females, and one is +unsexed. In a series of 13 specimens of the three subspecies of _B. +garrula_, loaned by the United States National Museum, all but two +individuals possess the tips on the secondaries, and, in addition, +four specimens, equally divided between the two sexes, have color on +the rachis of some rectrices, and small appendages of pigment extend +beyond the feathers. Stevenson (1882) found that among 144 specimens +of _B. garrula garrula_ killed by storms in England in the winter of +1866-67, 69 individuals had wax tips. Of these, 41 were males and 27 +were females; the remaining one was of uncertain sex. Among 38 +definitely sexed _B. garrula pallidiceps_ in the California Museum of +Vertebrate Zoölogy, Swarth (1922:276) lists tips in 22 males and 16 +females. These data indicate that the proportion of birds with the wax +tips is higher in _B. garrula_ than in _B. cedrorum_. The potentiality +for wax tips is possibly inherited according to Mendelian ratio. + +_Bombycilla japonica_ is of interest in that the adults, at least, +seldom have the waxy appendages. Nevertheless, in the specimens +observed, the entire distal ends of the feathers normally possessing +the tips in other species are suffused with red color. This may be the +original condition of all waxwings, or perhaps, instead, this species +is in a transitional stage in the development of the tips. Swarth +(1922:277) says concerning the probable derivation of the wax tips in +_B. garrula_ (and in _B. cedrorum_): "the ornamentation, in fact, may +well have begun with the coloring of the shaft, spreading later over +adjoining feather barbs. The last stage would have been the coalescing +of the barbs, forming the waxlike scale as is now seen. Various steps +of this hypothetical development are supplied in the wing and tail +feathers of different birds of this series." _Bombycilla japonica_ +thus may be close to the ancestral condition in the waxwing stock in +the development of the waxy appendage. + +The rectrices of all three species of waxwings seldom possess the wax +tips, unless the secondaries have the maximum number of tips. In these +individuals, the pigment seems to "spill over" onto the tail feathers. +Eight is the maximum number of tips found on the secondaries. +Rectrices with wax tips are more frequently found in _B. garrula_, and +only occasionally in _B. cedrorum_. The pigment in the tip of the tail +of _B. japonica_ is red rather than yellow as it is in the other two +species, and some individuals of the Japanese Waxwing show a slight +amount of coalescence of wax in the tail feathers as well as in the +secondaries. + +If the tips were present in all members of the two species, it could +be postulated, in line with recent investigational work by Tinbergen +(1947), that the tips are in the nature of species "releasers," +facilitating species recognition. Such recognition is now regarded as +of prime importance in the formation of species. It is improbable that +sex recognition may be aided, as there is no evidence to indicate that +the tips are found predominantly in either sex. + +The wax tips are not limited to the adult birds in the species _B. +garrula_. Swarth (_op. cit._) mentions the capture of several young +Bohemian Waxwings, and describes them as "possessing all the +distinctive markings of the most highly developed adult." This +includes wax appendages, and several citations are given (Wolley 1857, +Gould 1862) to indicate that this is the rule rather than the +exception, not only for the American subspecies _pallidiceps_, but at +least for the European subspecies _garrula_ as well. On the other +hand, the young of _B. cedrorum_ lack the wax tips, at least as far as +available data show. + +Some characteristics of living animals are of the "relict" type; that +is to say, they were developed in ancient times when some unknown +ecological factor was operative which is no longer demonstrable, and +the characteristic is now neutral or at least not detrimental, +although of no positive value to the organism. Possibly the wax tips +of waxwings are thus to be explained. I am more inclined to the +opinion that the wax tips are adaptations to present-day ecological +conditions for the birds. + +The wax tips are ruptive in effect, since the birds, especially in +winter, are habitués of bushes and trees that have berries, and the +tips, on the otherwise dull body, suggest berries. The red tips tend +further to disrupt the body outline at the midline, or slightly +posterior to this. Perhaps the wax tips on the rectrices emphasize the +end of the tail, the region of the body that is the least vital and +that may be expendable in times of pursuit by an enemy. + +Any characteristic is of survival value to an organism if in any way +the characteristic enhances the chances of survival up to the time +when the organism can successfully raise even a few young to maturity. +If that character, as for example, the red wax tips on the +secondaries, helps to maintain the individual until it can raise to +independence a greater number than merely a few young, such a +character can be said to be of greater survival value. The character +may be effective for a brief period of time and may be uncommon; it +might be effective for a split second in time, and only at a +particular stage in the life history. + +The winter period probably is the most hazardous for waxwings, in that +they then depend at times upon long flights to find food. The food is +vegetable, and thus is comparatively low in food value; the birds must +ingest large quantities of berries or dried fruits to maintain +themselves. In winter, in northern latitudes at least, predators are +more apt to prey upon those species which, like waxwings, do not +migrate south. The winter months are those in which waxwings frequent +berry bushes, and it may well be that in these months, the wax tips +that appear like berries, are especially valuable to the birds, and +operate selectively. + +It is suggested, therefore, that the wax tips are of positive value to +waxwings, rather than being relict characters. Coalescence of pigment +has taken place in the formation of the wax tips. _B. japonica_ is +closer to the ancestral stock insofar as wax tips are concerned, and +generally lacks the tips. _B. cedrorum_ has the tips in approximately +half of the adults, and not at all in the young. _B. garrula_ has the +tips in almost all the adults, and in a like proportion of the young, +and probably has evolved further in the development and retention of +the wax tips than has either of the other two species. + +The streaked plumage of _Dulus_ is decidedly generalized, and is +probably more nearly like the color of the ancestral stock. In this +connection it is notable that young Cedar Waxwings are streaked, and +young Bohemian Waxwings are streaked to a lesser degree. This +streaking is apparently a recapitulation of the feather color of the +stock. Perhaps the color of _Dulus_ has not changed, as the streaking +would not be a disadvantage to the birds in their environment of light +and shadow. In joining together in groups and in the construction of +large communal nests, _Dulus_ has evidently gained sufficient +protection against predators; other birds solve this problem by +modifying their coloration. + +_Ptilogonys_ is ruptively colored, but in a different fashion than +_Bombycilla_. The tail markings, the distinct yellow on the under tail +coverts, the sharply marked pileum, are all examples of ruptive +coloration. The generally lighter venter (especially under tail +coverts), the crest that may be elevated, and the generally drab +bluish dorsum, are cryptic and serve to hide the animal insofar as is +possible considering its habits. The very conspicuous coloration of +the male, in contrast to the more drab color of the female, however, +would lead one to believe that in _Ptilogonys_, following the pattern +of many passerine birds, the male leads a predator from the nest, +leaving the drab female to incubate the eggs, and thus preserve the +young. + +It is difficult to suggest reasons for the brilliant coloration of the +male _Phainopepla_, unless it is for decoying predators away from the +nest. Possibly some birds survive not because of, but in spite of, +their coloration, and _Phainopepla_ may be a case of this sort. Anyone +who has observed _Phainopepla_ in life will agree, certainly, that the +male makes no attempt at concealment, and flaunts his color to all +comers. + +The coloration of _Phainoptila_, in contrast to _Phainopepla_, is much +more plain, and is suited to its habits of brush dwelling; in a brush +habitat the drab coloration is difficult to detect. The Yellowish +Olive under tail-coverts and the Olivaceous dorsum are all evidences +of cryptic coloration, and undoubtedly, this bird depends upon hiding +for escape from its enemies, since it is a bird of the dense forest +cover. + +Coloration, which varies relatively rapidly in response to differing +ecological conditions, has become more different in the species of +Bombycillidae than is true in many other families of passerine birds. +The explanation lies in early geographical isolation of the three +subfamilies, with consequent radiation in three directions. Waxwings +have become adapted by possessing a thick protective layer of feathers +and drab coloration broken by ruptive marks. They still retain the +streaked plumage, which is probably ancestral, in the juveniles; this +is lost at the first molt in the fall. In its evolution, _Dulus_ has +developed large feet, heavy decurved beak, and the large communal nest +that affords protection from enemies; as a consequence, perhaps +_Dulus_ did not need a plumage different from the primitive and +streaked one. The survival of _Dulus_ may not have depended on either +ruptive marks or on brilliant and outstanding plumage. The large feet +and large bill seem to be responses to particular ecological +requirements, as will be shown later. + +The Ptilogonatinae, with habits paralleling those of the flycatchers, +probably are considerably modified from the ancestral stock; the +coloration probably is more brilliant and conspicuous. Perhaps this +type of coloration and the habit of capturing insects from a perch are +correlated. Some amount of territoriality is characteristic of this +subfamily and dimorphism in color--the plumage of the male is +outstandingly conspicuous--possibly is of selective value to the race. +In a tropical forest community, a duller pattern possibly would be +more visible and thus would be selectively disadvantageous. + + + + +COURTSHIP + + +Waxwings are gregarious birds and individuals establish no +well-defined territories as do many birds. The nest itself is the only +defended territory, and as Crouch (1936) has shown, the Cedar Waxwing +will nest in close proximity to others of the same species. Swarth +(1932:275) mentions that the Bohemian Waxwing is tolerant of the nests +of other pairs near by. The extreme condition is that found in +_Dulus_, in which the territory is not limited even to the nest, but +to the individual compartment of the community nest. _Phainopepla_, a +less gregarious bird than _Dulus_ and waxwings, has a much more +definite territory, although individuals of _Phainopepla_ are tolerant +of others of the same species; no feeding territory is established, +and small flocks of birds feed together at any time of the year. + +In birds whose territories lack well-defined boundaries, it would be +expected that elaborate song would not have evolved, and that most of +the recognition of kind and sex would be dependent upon the behavior +of the birds. This is the fact; song, as such, is lacking in the three +subfamilies Bombycillinae, Ptilogonatinae, and Dulinae. Waxwings utter +(1) notes that serve to keep the flock together, (2) calls used by the +young in begging for food, and (3) some low notes that Crouch (_op. +cit._:2) considered as possibly concerned with courtship. +_Phainopepla_ has various call notes, and in addition, a succession of +notes which are run together. _Ptilogonys_ utters a note which Skutch +(MS) characterizes as a loud, not unmusical "tu-whip" that is used as +the birds "fly in straggling parties which keep in contact by their +constant chatter." _Dulus_ is described by Wetmore and Swales +(1931:349) as having only a variety of rather harsh chattering notes +in chorus. + +The most notable behavior pattern associated with courtship in +Waxwings, in the absence of song, is the so-called "mating dance" +described by Crouch (1936), and observed by me in Lawrence, Kansas, in +the spring of 1948. This consists of one bird of a pair (presumably +the male) hopping along a branch toward the other bird (the female), +then away again, repeating the procedure for some little time. The +female remains motionless until, as the male approaches, mutual +fondling of the head and neck feathers takes place, or the birds may +peck at each other's bill. A berry may be passed from bill to bill, +although generally the berry is not utilized for food, and this can be +interpreted as a nervous reaction of the birds. It may be an instance +of "false feeding" as is seen in many birds, in which the female begs +for food, as a nestling would beg, as a preliminary to the sexual act. +I am of the opinion that these reactions are in the nature of +behavioristic patterns that bring the birds into the emotional balance +for copulation, as copulation follows the "dance." Sometimes, however, +copulation is preceded by a "nuptial flight" around the nesting area, +at which time the birds utter loud calls. Armstrong (1924:183) is of +the same opinion, citing numerous instances in which nuptial flights +and elaborate displays have evolved for just this purpose. The birds +are then in the proper physiological balance to initiate the +complicated sequence of copulation, nesting, incubation, feeding, and +brooding of the young. + +It would be valuable to know more concerning the life histories of the +other birds considered in this paper, since behavior is inherent, and +probably can be cited as evidence of close relationship or the +opposite. All that I have been able to learn is that _Phainopepla_ has +a nuptial flight in which the male chases the female, and that _Dulus_ +(Wetmore and Swales, 1931:347) seeks the company of others of its kind +at all times, and that two birds, presumably paired, will sidle up to +one another when they are perched. + + + + +NEST BUILDING + + +There are numerous papers concerning the nesting of waxwings. _B. +garrula_, owing to its nesting in the far north, where observers are +few, has received less attention than _B. cedrorum_. There is, on the +other hand, no literature that deals with the nesting habits of the +majority of the Ptilogonatines, with the exception of _Phainopepla_, +on which there is considerable literature (Merriam, 1896; Myers, 1907, +1908). No detailed study of the nesting of _Dulus_ has been reported, +although Wetmore and Swales (1931) have described carefully the large +communal nest of this genus. + +In _Bombycilla_, both members of a pair apparently aid in the +construction of the nest (Crouch, 1936; Swarth, 1932). Although the +sexes are alike in plumage and general appearance, most students of +the nesting of waxwings agree that one bird, assumed to be the female, +does most of the arranging of the material, and does the shaping of +the nest, whereas both birds carry materials to the nest site. As is +characteristic of many passerine birds, both members of the pair +gather materials and fly back to the nest site, where the female takes +the more active part in the construction of the nest itself. + +Both species of American waxwings build bulky nests, with the base or +platform composed of a large amount of twigs and sticks, from which +there often trails a mass of sticks and moss or string. Softer +materials such as moss, plant fibers, and string, are placed inside +the platform; moss is readily available to, and preferred by, _B. +garrula_ according to Swarth (_op. cit._:271), and various plant +fibers and string are used by _B. cedrorum_. The inner lining consists +of soft plant fibers or down, dry grasses, and feathers. The nest is +usually unconcealed in a tree either adjacent to a trunk or on a main +side branch, but sometimes in a fork. Nest building by both Cedar and +Bohemian waxwings is rapid, taking from three to five days, and is +followed immediately by egg laying. + +Nesting by waxwings is late in the season; June is the month in which +the nest is usually started. This is readily explainable in Bohemian +Waxwings, since adverse weather would prohibit earlier nesting in the +area in which they spend the summer. Crouch (_op. cit._:1) remarks +that _B. cedrorum_ possibly evolved in the far north where it was +impossible for it to start nesting earlier, and that the habit has +been retained. Perhaps, on the other hand, nesting is delayed until +the berry crop is ripe, to insure sufficient food for the young. + +Desertion of the nest is not uncommon in waxwings, despite the +tolerance to other animals that is shown by the birds. A new nest may +suddenly be begun before the first one is finished, and all the +materials from the first nest may be removed, or the nest may be +abandoned before it is completed. The eggs may be left at any time up +to hatching, and the young may be deserted, especially in the earlier +stages of development. + +The very large and bulky communal nest of _Dulus_ is not radically +different from the nest of waxwings. In the absence of sufficient +nesting sites, a pair of gregarious birds such as _Dulus_ could +combine their nest with those of other pairs, retaining for their own +territory only the nest cavity, and in this way communal nests might +have evolved. The nest of _Dulus_ is communal probably because of the +lack of suitable trees for nesting sites, and only incidentally does +this type of nest afford better protection from natural marauders. +Large numbers of Palm-chats work together in the construction of the +nest platform, and both sexes probably take part in the work. + +In _Phainopepla_ the nest is built mostly by the male (Merriam, 1896; +Myers, 1908), although the female does some of the work, especially in +the shaping and lining of the nest. In this genus, the nest is usually +a compact structure, but exceptional nests are of considerable bulk. +The nest is commonly placed in a fork near the main trunk of a tree, +in a conspicuous location, and generally is 10 to 20 feet from the +ground. In shape and location, the nest closely corresponds to that of +_Bombycilla_, but the materials used for a base are stems of annual +plants, whereas _Bombycilla_ uses more woody twigs. The finer +materials used by _Phainopepla_ are more readily obtainable in the +ecological association inhabited by _Phainopepla_ than would be +heavier twigs such as _Bombycilla_ uses. + + + + +FOOD + + +Waxwings are typically frugivorous; berries are the staple food. The +birds are known to catch insects, especially in the spring and summer, +and their insect gathering technique has been likened to that of +Tyrannid flycatchers. Nice (1941) experimented with a young captive +Cedar Waxwing and found that it had a decided preference for red or +blue berries, and that meal worms were utilized as food only when the +birds became educated by other captive birds of other species as to +the food value of the worms. Post (1916) indicates that the food given +to the nestlings of Cedar Waxwings is entirely animal for the first +three days, and that a mixed diet of berries and insects is +subsequently offered. + +In feeding of the young, regurgitation of partly digested food does +not take place, according to Wheelock (1905). Rather, the adults +"store" food in the form of berries in the expanded esophagus or crop, +feeding them whole to the young. Digestion is an unusually rapid +process, involving merely minutes for the passage of berries and +cherries. This is correlated with a short intestinal tract, which is +unusual for a frugivorous bird. Nice's (1940) experiments with Cedar +Waxwings revealed that cherries would pass through the digestive tract +in 20 minutes, blueberries in 28 minutes, and chokecherries in 40 +minutes. Heinroth (1924) states that berries pass through the +digestive tract of Bohemian Waxwings in the space of a "few minutes." +This rapid digestion is obviously adaptive, since the value of the +food is slight and therefore large quantities of it must be ingested; +the large seeds would hamper further ingestion until they were +eliminated, since they seem not to be regurgitated. + +Members of the subfamily Ptilogonatinae are both insectivorous and +frugivorous insofar as available data show, although again there is +relatively little information available concerning them. Skutch (MS) +has found that the Guatemalan _Ptilogonys cinereus_ catches insects by +repeated sallies into the air from a perch, after the manner of +flycatchers. He notes also that the birds feed on berries of _Eurya +theoides_ and _Monnina xalapensis_. It is well known that +_Phainopepla_ catches insects when these are available, and its liking +for berries is so apparent that in parts of its range, it is known as +the "pepper bird," since it frequents pepper trees (_Schinus molle_) +and feeds on the small red berries. The preserved specimens of +_Ptilogonys_ and _Phainoptila_ available for this study contain only +berries in the digestive tract. _Dulus_ feeds mostly, if not wholly, +on plant food. According to Wetmore and Swales (1931:349), berries, +fruits, and parts of flowers are eaten. + + + + +SKELETON + + +A critical analysis of the skeletons provides evidence that aids the +student in estimating which differences are merely the result of +habits developed in relatively recent geological time as opposed to +those which owe their existence to more ancient heritage. Stresses +caused by the action of different sets of muscles can apparently +stimulate changes in bones to meet new needs, and the evidence from +genetics is that such mutations in wild birds are minute and +cumulative, rather than of large degree and of sudden appearance. Once +adaptive mutations have occurred, if genetic isolation from one source +or another accompanies it, a new population different from the +parental stock may become established. Study of the skeleton of any +species of living bird may indicate those characters identifiable as +modifications fitting it to a particular environment. If no +distinguishing characters are discovered that may be attributed to +environmental factors, such a species can be spoken of as generalized; +the inference then is that such a species is not modified for a +single, particular ecological niche. + +Some parts of the skeleton, obviously, are more adaptable or plastic +than others. The beak seems to be the most adaptable part. Probably +this results from its frequent use; it is the part of the bird to +capture the food. The long bones, meeting the environment as legs +which serve as landing mechanisms or as locomotory appendages, and as +wings which provide considerable locomotion for most birds, probably +come next in order as regards plasticity. In these parts, then, one +may look for the most change in birds, which, within relatively recent +geologic times, have been modified to fit a particular set of +conditions. From the beak and long bones of a species in which habits +are unknown, one can infer the habits and habitat from a comparison +with the skeletal features of species of known habits. + + +_Skull._--The skulls in all three subfamilies have essentially the +same general appearance and structure, the most marked differences +being, as would be expected, in the bills and associated bones. + +The most specialized bill is to be found in _Dulus_; its bill is +decurved, and the associated bones are correspondingly changed for +support of the bill. For example, the palatines and "vomer" are much +wider, the palatines are more concave from below and have longer +posterior processes than the corresponding bones in _Bombycilla_. +Moreover, the "vomer" in _Dulus_ and in _Phainoptila_ is larger and +heavier than in _Bombycilla_, and the quadrate and pterygoid bones are +relatively large for support of the beak. The palatines, however, are +weak in _Phainoptila_. In the Ptilogonatinae, with the exception of +_Phainoptila_, the wings of the palatines flare more than in +_Bombycilla_, but not to the extent that they do in _Dulus_, nor does +the palatine bone present a concave appearance in the Ptilogonatinae. +The premaxilla is a relatively weak bone in _Bombycilla_ and +_Phainopepla_, stronger in _Ptilogonys_, and is notably heavy in +_Phainoptila_ and _Dulus_, and in these latter two genera shows a +sharply-ridged tomium. The maxillae connect to somewhat widened nasal +and naso-lateral processes in all the genera, and the premaxillae +narrow abruptly from this point forward. In the family, _Phainopepla_ +and _Phainoptila_ show the least flaring in this region. + + + [Illustration: Figs. 1-7. Skulls in lateral view of five genera of + Bombycillidae. Natural size. + + 1. _Phainoptila m. melanoxantha_, sex?, MNH no. 26493, 15 mi. + SE Cartago, Costa Rica. + + 2. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 3. _Phainopepla nitens_, male, MNH no. 24752, Pima Co., Arizona. + + 4. _Ptilogonys cinereus_, female, Louisiana State University + no. 297, Xilitla Region, San Luís Potosi, Mexico. + + 5. _Dulus dominicus_, female, USNM no. 292652, Don Don, Haiti. + + 6. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 7. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + + [Illustration: Figs. 8-14. Skulls in ventral view of five genera of + Bombycillidae. Natural size. + + 8. _Phainoptila m. melanoxantha_, sex?, MNH no. 26492, 15 mi. + SE Cartago, Costa Rica. + + 9. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 10. _Phainopepla nitens_, male, MNH no. 24754, Pima Co., Arizona. + + 11. _Ptilogonys cinereus_, female, Louisiana State University + no 297, Xilitla Region, San Luís Potosi, Mexico. + + 12. _Dulus dominicus_, female, USNM no. 292652, Don Don, Haiti. + + 13. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 14. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + + [Illustration: Figs. 15-21. Skulls in dorsal view of five genera of + Bombycillidae. Natural size. + + 15. _Phainoptila m. melanoxantha_, sex?, MNH no. 26493, 15 mi. + SE Cartago, Costa Rica. + + 16. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 17. _Phainopepla nitens_, male, MNH no. 24752, Pima Co., Arizona. + + 18. _Ptilogonys cinereus_, female, Louisiana State University + no. 297, Xilitla Region, San Luís Potosi, Mexico. + + 19. _Dulus dominions_, female, USNM no. 292642, Don Don, Haiti. + + 20. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 21. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + +This flaring, immediately lateral to the antorbital plate, is common +to all Bombycillids and constitutes a major skeletal characteristic +useful for recognition of the members of the family, since the +swelling is easily discernible both externally and on the cleaned +skulls. In _Phainopepla_ there is much variability in this character; +some specimens have a narrower antorbital bridge than others. Only one +skeleton of _Phainopepla n. nitens_ was available. The flaring in the +skull of this specimen is identical with that in _Ptilogonys_. Among +the skulls of _P. n. lepida_ in the University of Kansas Museum of +Natural History, is No. 19228, a juvenile, taken 5 miles south of +Tucson, Arizona. In this specimen, the flaring in the antorbital +region is clearly evident and equal in amount to that in skulls of _P. +n. nitens_, but the bird had not attained full skeletal growth. +However, the flaring of the antorbital region appears to be common in +the nestlings of many species of passerine birds. Other specimens of +the subspecies _lepida_ show a varying amount of flaring, the least +(in the series available) being in No. 24754, MNH, in which the +proportion of the skull (length divided by width) closely corresponds +to that in _Phainoptila_; the skull of No. 24754 is long and thin, and +the base of the bill is only slightly swollen. The skull of +_Phainopepla nitens lepida_ is more generalized than that of +_Phainopepla n. nitens_, having a longer and narrower bill like the +generalized _Phainoptila_. In _Phainopepla n. nitens_ and in members +of the genus _Ptilogonys_, more flaring occurs in the antorbital +region. + +_Phainoptila_, as noted above, has no great amount of flaring in the +antorbital region. When more specimens of _Phainoptila_ are examined, +the base of the bill probably will be found to flare more in some +individuals than in others; this would be expected if we may judge by +the data on _Phainopepla_. The premaxilla and maxilla of _Phainoptila_ +are similar to the same bones in _Dulus_, and there is a well-marked +ridge on the tomium (possibly for cutting flower parts). In +_Phainoptila_, the palatines are narrower than in any other genus of +the family and abut the lacrimals. The entire skull appears to be +modified along different lines from those of the skull of _Dulus_; the +skull of _Phainoptila_ seems to be modified for a frugivorous rather +than an insectivorous diet. The skull of _Phainoptila_ probably is +more nearly similar to the ancestral skull than is that of any other +living species in the family. The wide gape characteristic of some +members of the family is undoubtedly a modification for aiding in the +capture of insects, and _Phainoptila_ has progressed less in this +direction than have other species in the family. + +The mandibles vary somewhat in the shape and proportionate size of the +bones. The mandible is proportionately, as well as actually, highest +in _Dulus_. The medial condyle varies to some extent, being slightly +flattened mediad in _Bombycilla_, and less so in the other genera. The +mandible of _Bombycilla_ narrows to the symphysis much more gradually +than it does in the other genera. + +The antorbital plate is large and divides the orbital chamber from the +nasal chamber. The small lacrimal bone anterior to the plate +articulates with the maxilla and the premaxilla. Shufeldt (1889) +states that the free lacrimal ossicle might be of some taxonomic +importance in the passerines, since it is found in the generalized +Corvids and in nestling Turdids. I find it well developed and +identical, with a double articulation and free ends, in all the +Bombycillids. There is no significant variability in the family, and +this is more evidence of close taxonomic relationship between the +members of the family. + +The size of the crania is somewhat variable, although the differences +seem to be primarily those of proportion. Ptilogonatinae have long +crania, whereas the crania of the Bombycillinae and Dulinae are +shorter but deeper. I regard the longer cranium as primitive, and it +is longest in _Phainoptila_. In order of decreasing relative length of +the cranium, _Phainoptila_ is followed by _Ptilogonys caudatus_, _P. +cinereus_, and _Phainopepla_. _Bombycilla garrula_ has the deepest +cranium in the family. + +The measurements of the lengths and widths of the skulls are given in +Table 9. The relative length of the bill and relative width of the +skull are given in Table 10. These relative measurements are +calculated by using the actual measurements in Table 9 as numerators, +the length of the skull from the lacrimal bone to the posteriormost +end of the skull being used as the denominator. The data indicate that +_Phainoptila_ has a slightly narrower cranium. + + +_Humerus._--Certain families of passerine birds have a noticeable +variation in the characteristics of the humerus; the bone varies in +length, in diameter, and in the complexity of the processes at either +end. In the Bombycillids, however, the amount of variation is +relatively small, and the diaphysis of the bone is somewhat twisted, +especially so in _Dulus_. The deltoid tuberosity is variable, being +shorter but more elevated in _Bombycilla_ than it is in the +Ptilogonatinae and in the Dulinae. The tendon from the pectoralis +major muscle, which inserts on this process, probably finds better +insertion on a higher process than on a lower but longer one. + + + [Illustration: Figs. 22-28. Humeri of five genera of Bombycillidae. + Natural size. + + 22. _Phainoptila m. melanoxantha_, sex?, MNH no. 26493, 15 mi. + SE Cartago, Costa Rica. + + 23. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 24. _Phainopepla nitens_, male, MNH no. 24754, Pima Co., Arizona. + + 25. _Ptilogonys cinereus_, female, Louisiana State University + no. 297, Xilitla Region, San Luís Potosi, Mexico. + + 26. _Dulus dominicus_, female, USNM no. 292652, Don Don, Haiti. + + 27. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 28. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + +Distally, the two major condyles and the intercondylar groove or +olecranon fossa that make efficient articulation with the ulnar +process, are not variable. The external condyle, however, is +significantly variable in the family. This condyle is longest and most +pronounced in birds in which the humerus is short in relation to the +trunk, as for example in _Tachycineta_. In the Bombycillidae the +condyle is smallest in _Phainoptila_, where it is a mere suggestion of +a process. In the remainder of the Ptilogonatinae, the condyle is +larger but rounded, and shows a double process in _Ptilogonys +caudatus_, and a slightly pointed process in _P. cinereus_. The +external condyle in _Dulus_ is not specialized, being low and rounded, +but in _Bombycilla_, it is noticeably elongated, indicating a better +attachment distally for the deltoid muscle. (No measurements are +tabulated for this condyle, as the percentage of error in measuring +this small structure is great.) Table 1 gives lengths of humeri, and +Table 2 gives lengths of the humeri expressed as percentages of the +length of the trunk, a standard measurement. + +The area of insertion of the deltoid muscle is elongated in those +birds with shortened humeri; these birds have also greater flight +power than do birds with longer humeri and therefore a shorter +external condyle. + + + Table 1. Lengths of Arm Bones in cm. + + =========================+=========+========+======+======= + Species | Humerus | Radius | Ulna | Manus + -------------------------+---------+--------+------+------- + Ptilogonys caudatus | 2.39 | 2.57 | 2.79 | 2.25 + Ptilogonys cinereus | 2.24 | 2.48 | 2.78 | 2.38 + Phainopepla nitens | 2.21 | 2.59 | 2.82 | 2.39 + Phainoptila melanoxantha | 2.40 | 2.51 | 2.70 | 2.25 + Dulus dominicus | 2.23 | 2.38 | 2.63 | 2.31 + Bombycilla garrula | 2.35 | 2.58 | 2.88 | 2.67 + Bombycilla cedrorum | 2.06 | 2.34 | 2.60 | 2.38 + -------------------------+---------+--------+------+------- + + + Table 2. Arm-trunk Ratios (in percent) + + =========================+=========+========+======+=======+======= + Species | Humerus | Radius | Ulna | Manus | Total + -------------------------+---------+--------+------+-------+------- + Ptilogonys caudatus | 85 | 92 | 93 | 80 | 2.58 + Ptilogonys cinereus | 84 | 90 | 103 | 89 | 2.76 + Phainopepla nitens | 84 | 98 | 107 | 91 | 2.82 + Phainoptila melanoxantha | 73 | 77 | 82 | 69 | 2.31 + Dulus dominicus | 78 | 83 | 92 | 81 | 2.51 + Bombycilla garrula | 69 | 75 | 87 | 78 | 2.34 + Bombycilla cedrorum | 67 | 76 | 85 | 77 | 2.29 + -------------------------+---------+--------+------+-------+------- + + + Table 3. Arm-trunk Ratios (in percent) + + =========================+=========+========+======+=======+======= + Species | Humerus | Radius | Ulna | Manus | Total + -------------------------+---------+--------+------+-------+------- + Corvus brachyrynchos | 90 | 101 | 111 | 106 | 307 + Dendroica audubonii | 68 | 82 | 90 | 77 | 237 + Setophaga ruticilla | 69 | 82 | 91 | 75 | 235 + Myadestes townsendi | 71 | 84 | 96 | 81 | 248 + Sialia sialis | 72 | 84 | 98 | 86 | 256 + Hylocichla mustelina | 75 | 81 | 92 | 80 | 247 + Parus atricapillus | 85 | 90 | 106 | 81 | 272 + Tachycineta thalassina | 71 | 95 | 107 | 128 | 306 + Myiarchus crinitus | 83 | 105 | 115 | 92 | 290 + Dumetella carolinensis | 76 | 75 | 89 | 78 | 243 + Polioptila caerulea | 85 | 93 | 105 | 71 | 261 + Eremophila alpestris | 91 | 99 | 110 | 95 | 296 + Muscivora forficata | 85 | 111 | 120 | 108 | 313 + -------------------------+---------+--------+------+-------+------- + + +_Pygostyle._--This part of the skeletal system is variable in the +species dealt with, not so much in size as in complexity. It reflects, +of course, the character of the caudal muscles and their size, as well +as the length of the rectrices and the corresponding force necessary +to hold these feathers upright and in a useful position. Firm +attachment is important even in flight, because the tail is used as a +rudder, and in the Ptilogonatinae as a brake. The pygostyle is most +modified in this subfamily. + +In lateral aspect, the pygostyles of the species of the Ptilogonatinae +are similar. The crest of the bone is flattened dorsally, and has a +broad anterior surface that is thin and bladelike. This is widest in +_Ptilogonys caudatus_, and narrowest in _Phainoptila_, in which genus, +however, the entire bone is of small size. The centrum is widest in +_Ptilogonys caudatus_, and is progressively narrower in _P. cinereus_, +_Phainopepla_, and _Phainoptila_. Greater width provides a larger area +of attachment for the larger rectrices and also more area for +insertion of the lateralis caudae muscle, the size of which varies +more than that of the other caudal muscles in the different species of +the Bombycillidae. + + + [Illustration: Figs. 29-35. Pygostyles in posterior view of five + genera of Bombycillidae. × 2. + + 29. _Phainoptila m. melanoxantha_, sex?, MNH no. 26493, 15 mi. + SE Cartago, Costa Rica. + + 30. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 31. _Phainopepla nitens_, male, MNH no. 24754, Pima Co., Arizona. + + 32. _Ptilogonys cinereus_, female, Louisiana State University + no. 297, Xilitla Region, San Luís Potosi, Mexico. + + 33. _Dulus dominicus_, female, USNM no. 292652, Don Don, Haiti. + + 34. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 35. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + +In proportionate size (see Table 7), the pygostyle of _Bombycilla_ is +the smallest in the family. The dorsal spinous portion is acutely +pointed instead of flattened as in the Ptilogonatinae. In _Dulus_, the +spinous portion is extremely thin, and shows a decided curve dorsad +from the centrum, and there is no flattened area anterior to the +spinous portion as is seen in _Ptilogonys_. + +The centrum in cross section varies considerably. In _Bombycilla_ the +walls are indented, with definite terminal knobs; both knobs and +indentations are more pronounced in _B. garrula_ than in _cedrorum_, +however. The spinous portion is enlarged in both species, and the rest +of the neck region is constricted (Figs. 29-35). + +The centrum of _Dulus_ in posterior aspect presents the appearance of +a simple shield; little of the indentation seen in _Bombycilla_ is +present. The spinous portion is plain, with no constriction nor +terminal enlargement in the neck. The centrum in _Phainopepla_ is +similar to that in _Dulus_, but has a small expansion at the base of +the spine, the entire centrum being wider in proportion to its +over-all size than in any of the other species mentioned previously. +The centrum in _Ptilogonys_ shows great width, and the spine is in a +large expanded tip as in _Bombycilla_. The lateral edges of the +centrum in _P. cinereus_ are "winged" and in two separate halves; +whereas the centrum of _P. caudatus_ is fairly plain, its +specialization being reflected primarily in breadth and flatness. In +cross section of the centrum, _Phainoptila_ is similar to +_Phainopepla_, although, in the former, the bone is smaller in +proportion to the size of the animal, and the lateral wings are more +angular than in _Phainopepla_. + + + [Illustration: Figs. 36-42. Pygostyles in lateral view of five + genera of Bombycillidae. × 2. + + 36. _Phainoptila m. melanoxantha_, sex?, MNH no. 26493, 15 mi. + SE Cartago, Costa Rica. + + 37. _Ptilogonys caudatus_, male, MNH no. 24492, 15 mi. SE Cartago, + Costa Rica. + + 38. _Phainoptila nitens_, male, MNH no. 24754, Pima Co., Arizona. + + 39. _Ptilogonys cinereus_, female, Louisiana State University + no. 297, Xilitla Region, San Luís Potosi, Mexico. + + 40. _Dulus dominicus_, female, USNM no. 292652, Don Don, Haiti. + + 41. _Bombycilla cedrorum_, male, MNH no. 15331, Bexar Co., Texas. + + 42. _Bombycilla garrula_, sex?, USNM no. 223895, Bozeman, Montana.] + + +In specialization for muscle attachment, the centra of the pygostyles +of the Ptilogonatinae have more area for muscle attachment than do the +centra in the Bombycillinae and Dulinae; the centrum is wide, the +spinous portion is long, and the bone is flattened anteriorly. The +most generalized pygostyle is in _Phainoptila_, and that of _Dulus_ +differs only slightly. In _Bombycilla_ the pygostyle is +proportionately small, but is complex in shape; there is seemingly not +the need for greatly expanded areas since the caudal muscles are less +specialized in this genus. + + +_Sternum._--The sternum in Bombycillids is typically passerine in +general shape and in having a long and deep carina or sternal crest. +The caudal process of the bone is broad, with the terminal ends +flattened, forming dorsally a graceful V-shaped outline, whereas the +outline of the posterior end of the sternum is broad and convex. + +In lateral aspect, the carina is deeper in _Bombycilla_ than in other +genera of the family, and is deepest in _B. garrula_. In this species, +the manubrium is more extended and comparatively larger than in the +other species of the family. The anterior edge of the keel forms the +sharpest angle in _B. cedrorum_. In _Dulus_, the keel is moderately +deep, the manubrium short, and there is a distinct indented curve +between the manubrium and the anterior angle of the keel. + +In ventral aspect the lateral processes of the sternum tend to flare +outwards in adult Ptilogonatines on almost the same plane as the rest +of the bone, whereas in _Bombycilla_ and _Dulus_ the same process is +closer to the body of the sternum. In _Bombycilla_ the xiphoid process +is more dorsal in position than in other species in the family, and in +_Dulus_ an upward curve is very noticeable. The process in these two +genera is narrower than in the Ptilogonatinae, and lacks the heavy +distal terminal enlargement which is apparent in _Ptilogonys_. + + +_Relative Lengths of Bones._--In instances where the animals being +compared are obviously different in over-all size, it is useful to +express the size of a given part in relation to some other part of the +same individual organism if the aim is to obtain clues as to +differences in functions of the parts being compared. Differences in +actual lengths of corresponding bones in two kinds of animals often, +of course, reflect only the difference in over-all size of the +animals. Consequently, the relative size of the part is expressed as a +percentage in this paper. In computing a percentage it is well, of +course, to select some relatively stable part of the animal to use as +a denominator in the mathematical expression that yields the +percentage. The thoracic region of the vertebral column is thought to +be such a part. For example, the length of the humerus divided by the +length of the thoracic region yields, in _Phainopepla_ and +_Ptilogonys_, respective percentages of .84 and .85. These are roughly +the same, whereas the actual lengths of the humeri are 2.21 and 2.39 +cm. + + + Table 4. Lengths of Leg Bones in cm. + + =========================+=======+=============+================= + Species | Femur | Tibiotarsus | Tarsometatarsus + -------------------------+-------+-------------+----------------- + Ptilogonys caudatus | 2.04 | 3.10 | 1.94 + Ptilogonys cinereus | 1.89 | 2.90 | 1.77 + Phainopepla nitens | 1.76 | 2.78 | 1.72 + Phainoptila melanoxantha | 2.43 | 3.77 | 2.58 + Dulus dominicus | 2.09 | 3.34 | 2.09 + Bombycilla garrula | 2.32 | 3.46 | 1.99 + Bombycilla cedrorum | 1.92 | 2.95 | 1.64 + -------------------------+-------+-------------+----------------- + + + Table 5. Leg-trunk Ratios (in percent) + + ====================+=======+=============+=================+======= + Species | Femur | Tibiotarsus | Tarsometatarsus | Total + --------------------+-------+-------------+-----------------+------- + Ptilogonys caudatus | 73 | 110 | 69 | 252 + Ptilogonys cinereus | 71 | 109 | 66 | 246 + Phainopepla nitens | 69 | 106 | 65 | 240 + Phainoptila | 74 | 115 | 60 | 249 + melanoxantha | | | | + Dulus dominicus | 73 | 119 | 73 | 265 + Bombycilla garrula | 68 | 101 | 59 | 228 + Bombycilla cedrorum | 63 | 96 | 53 | 212 + --------------------+-------+-------------+-----------------+------- + + + Table 6. Leg-trunk Ratios (in percent) + + =======================+=======+=============+=================+====== + Species | Femur | Tibiotarsus | Tarsometatarsus | Total + -----------------------+-------+-------------+-----------------+------ + Corvus brachyrynchos | 71 | 120 | 77 | 268 + Corvus corax | 73 | 139 | 78 | 290 + Dendroica audubonii | 62 | 109 | 81 | 252 + Setophaga ruticilla | 66 | 127 | 94 | 287 + Myadestes townsendi | 61 | 99 | 60 | 220 + Sialia sialis | 66 | 111 | 72 | 249 + Hylocichla mustelina | 75 | 133 | 97 | 305 + Parus atricapillus | 78 | 138 | 99 | 315 + Tachycineta thalassina | 61 | 97 | 56 | 214 + Myiarchus crinitus | 68 | 106 | 74 | 248 + Dumetella carolinensis | 73 | 136 | 94 | 303 + Polioptila caerulea | 75 | 144 | 113 | 332 + Eremophila alpestris | 73 | 113 | 115 | 301 + Muscivora forficata | 62 | 98 | 61 | 221 + -----------------------+-------+-------------+-----------------+------ + + + Table 7. Actual Length and Width in mm. of Pygostyle and Proportionate + Length and Width of Pygostyle in percent of Lacrimal Length + + =========================+========+=======+=========+========= + | | | Length, | Width, + Species | Length | Width | percent | percent + -------------------------+--------+-------+---------+--------- + Ptilogonys caudatus | 9.8 | 3.9 | 45 | 18 + Ptilogonys cinereus | 8.8 | 4.1 | 41 | 19 + Phainopepla nitens | 8.4 | 3.9 | 41 | 19 + Phainoptila melanoxantha | 8.5 | 3.5 | 35 | 14 + Dulus dominicus | 8.5 | 2.9 | 38 | 13 + Bombycilla garrula | 7.0 | 3.5 | 31 | 15 + Bombycilla cedrorum | 7.1 | 2.9 | 35 | 14 + -------------------------+--------+-------+---------+--------- + + + Table 8. Length of Sternum and Depth of Carina expressed as + percentages of the Length of the Trunk + + =========================+=========+======== + Species | Sternum | Carina + -------------------------+---------+-------- + Ptilogonys caudatus | 85 | 28 + Ptilogonys cinereus | 91 | 32 + Phainopepla nitens | 81 | 26 + Phainoptila melanoxantha | 76 | 25 + Dulus dominicus | 107 | 28 + Bombycilla garrula | 88 | 33 + Bombycilla cedrorum | 82 | 31 + -------------------------+---------+-------- + + + Table 9. Skull and Sternum, Length and Width in mm. + + =========================+========+=======+=========+========= + | Length | Width | Length | Width + Species | of | of | of | of + | Skull | Skull | Sternum | Sternum + -------------------------+--------+-------+---------+--------- + Ptilogonys caudatus | 34.9 | 15.6 | 23.9 | 7.8 + Ptilogonys cinereus | 33.4 | 14.7 | 24.3 | 8.5 + Phainopepla nitens | 33.3 | 15.1 | 21.3 | 6.9 + Phainoptila melanoxantha | 39.7 | 16.0 | 24.8 | 8.2 + Dulus dominicus | 36.4 | 16.6 | 30.5 | 8.0 + Bombycilla garrula | 37.0 | 16.8 | 30.0 | 11.2 + Bombycilla cedrorum | 34.0 | 15.5 | 25.3 | 9.6 + -------------------------+--------+-------+---------+--------- + + +The length of the trunk was taken as the distance from the anterior +tip of the neural crest of the last cervical vertebra to the anterior +edge of an acetabulum. The number of free thoracic vertebra was five +in each specimen; consequently, there was no error from this source. +In the cranium, a measurement was taken from the anterior edge of the +lacrimal bone to the posteriormost end of the cranium, and the +resultant figure was employed for a constant in cases in which small +bones were compared. + + + Table 10. Relative Length and Width of Skull (in percent) + + =========================+========+======= + | Length | Width + Species | of | of + | Skull | Skull + -------------------------+--------+------- + Ptilogonys caudatus | 160 | 72 + Ptilogonys cinereus | 158 | 69 + Phainopepla nitens | 162 | 73 + Phainoptila melanoxantha | 161 | 65 + Dulus dominicus | 164 | 75 + Bombycilla garrula | 164 | 74 + Bombycilla cedrorum | 162 | 74 + -------------------------+--------+------- + + + [Illustration: Fig. 43. Part of skeleton of _Bombycilla cedrorum_ + showing method of measuring the length of the trunk. + Natural size.] + + +_Leg-trunk Percentages._--Table 4 shows the relative lengths of the +legs and of the separate bones in the legs of the different species of +the Bombycillids. Table 5 shows corresponding lengths for other +passerine birds. The total length of the leg was computed by adding +the figures obtained for the lengths of the femur, tibiotarsus and +tarsometatarsus. The lengths of the toes were disregarded. Length of +leg was recorded in this same way by Richardson (1942:333), who +thought that only in swimming and running birds do the toes contribute +to the functional length of the hind limb. + +Table 4 shows that of the birds compared in this paper, _Dulus_ has +the longest legs. In order of decreasing length the others are the +Ptilogonatinae, and finally the Bombycillinae, which have the shortest +legs of all. In Waxwings the length of the legs, expressed as +percentages of the body-lengths, are identical with those birds that +are similar in habits, that is to say, birds which do not use the hind +limb except in perching. It can be noted by reference to Table 5 that +_Tachycineta_ and _Myadestes_ fall into this category. This shortness +of limb is obviously adaptive, and each of the segments of the limb +has been correspondingly shortened, with no element reduced at the +expense of the other two. The short leg can be more easily folded +against the body while the bird is in flight, than can a long leg +which is more unwieldy. It may be noted from tables 4 and 5 that birds +which spend much time on the ground, or that hop a great deal in the +underbrush, have longer legs than do birds which spend much time in +flight. Two birds with noticeably long legs are _Hylocichla +mustelina_, a typical ground dweller, and _Parus atricapillus_, which +hops about in the trees and underbrush. + +Insofar as the lengths of the legs show, _Dulus_ and _Phainoptila_ are +the most generalized of the Bombycillidae, since the relative length +of leg is approximately the same as that of more generalized birds +such as warblers, crows and thrushes of similar locomotory habits. In +other words, _Dulus_ and _Phainoptila_ have remained unspecialized, in +contrast to the waxwings in which adaptive changes fitting them for a +perching habit have taken place. _Ptilogonys_ and _Phainopepla_ are +intermediate in length of leg between _Phainoptila_ and _Bombycilla_, +and _Ptilogonys_ and _Phainopepla_ have progressed from life on the +ground toward the perching habit. _Bombycilla cedrorum_ is more +specialized than is _B. garrula_ in shortness of leg, and the +reduction is comparable, as is noted above, to that in the legs of +_Tachycineta_. + +In birds which have the legs much modified for walking or for hopping +in the brush, such as _Polioptila_ and _Eremophila_, it is noteworthy +that the distal segment, the tarsometatarsus, is the longest, whereas +in birds such as _Myiarchus_ and _Tachycineta_, that do not utilize +the limbs in this manner, the tibiotarsus, the middle segment, is the +longest. Mammals much modified for walking or hopping likewise have +the proximal segment, the femur, short, and the distal segment long +(Howell, 1944). The waxwings have all of the segments short; these +birds are modified for strong and sustained flight. Their hind limbs +are used principally for landing devices and for perching. No one +element of the leg has been shortened much, if any, more than any +other. + + + [Illustration: Fig. 44. Graph showing relative lengths of bones of + the leg. The percentage values are shown on the axis + of the ordinates. + + A. _Bombycilla cedrorum_; B. _Bombycilla garrula_; + C. _Dulus dominicus_; D. _Phainoptila melanoxantha_; + E. _Phainopepla nitens_; F. _Ptilogonys cinereus_; + G. _Ptilogonys caudatus_. + a. femur; b. tibiotarsus; c. tarsometatarsus; d. total.] + + +_Arm-trunk Percentages._--Tables 1 and 2 show the total length of the +arm, and lengths of the separate arm elements, relative to the trunk. +Table 3 gives the corresponding lengths for birds other than the +Bombycillidae. Total length of arm was obtained by adding together the +lengths of the humerus, ulna, and manus, and by dividing the figure +thus obtained by the length of the trunk as was done for leg lengths +in tables 4 and 5. The method of adding together the component parts +does not give the entire length of the wing, since the length of the +feathers, which add effectively to the total length, as well as do the +lengths of the small carpal elements, is lacking. + + + [Illustration: Figs. 45-46. Outlines of wings. × 1/2 + + 45. _Ptilogonys caudatus_, showing relation of outline of wing + to bones of arm. + + 46. _Bombycilla cedrorum_, showing relation of outline of wing + to bones of arm.] + + +It may be noted that _Phainoptila_ and _Bombycilla_ have the shortest +arm in the family Bombycillidae. The humerus, radius and ulna are +comparable to the same elements in thrushes and the catbird, and it is +only the extremely short manus in _Phainoptila_ that affects the +total. The manus in _Phainoptila_ is comparatively smaller than in any +other genus of the family Bombycillidae, and this indicates poor +flight power. _Bombycilla_ has a total length corresponding closely to +that in warblers, but the lengths of the distal elements correspond +closely to those in the catbird and thrushes. Of the three segments, +the humerus is, relatively, the most shortened. Next in order of +increasing length of arm is _Dulus_; measurements for it are roughly +the same as those of _Myadestes_. The wing bones of the +Ptilogonatinae, other than _Phainoptila_, are the longest in this +series, and they most nearly resemble the same bones in flycatchers, +Parids, and gnatcatchers. + + + [Illustration: Fig. 47. Graph showing relative lengths of bones of + the arm. The percentage values are shown on the axis + of the ordinates. + + A. _Bombycilla cedrorum_; B. _Bombycilla garrula_; + C. _Dulus dominicus_; D. _Phainoptila melanoxantha_; + E. _Phainopepla nitens_; F. _Ptilogonys cinereus_; + G._ Ptilogonys caudatus_. + a. humerus; b. radius; c. ulna; d. manus; e. total.] + + +It is notable that, in general, birds with long and narrow wings +appear to have relatively the shortest humeri, with the distal bones, +especially the manus, variable in length and seemingly correlated with +the manner of feather attachment. Those birds with rounded and short +wings have the longest humeri. In swallows, for example, the humerus +is short, whereas the other arm bones are long, and the manus is +unusually large and heavy. A short humerus gives better lever action +in the flight stroke than a long humerus does. + + + + +MUSCULATURE + + +Dissections showed the same muscles to be present in all genera of the +Bombycillidae. There are, nevertheless, differences in the size of the +muscles in the various species, and these differences have been +investigated primarily as a check on differences noted in the +structure of the bones. Even slight differences in mass can be +important functionally, but the difficulty in accurately measuring the +mass prevents wholly reliable conclusions. The method first used in +the attempt to determine the mass of a given muscle was that of +immersing the muscle in a liquid-filled graduated tube, and then +measuring the amount of liquid displaced. This method, although +adequate for large muscles, was subject to a great amount of error in +the case of small muscles, and consequently was abandoned. The +technique eventually used was that previously employed by Richardson +(1942). It consisted of dissecting out the muscle, placing it in +embalming solution, leaving it there until a later period, and +finally, weighing the muscle on scales, accurate to a milligram, after +the muscle had been out of the liquid for a period of one minute. +After being weighed, the muscle was measured by the displacement +method in a graduated tube, as a check. The results indicate that, +although the two methods give the same general results, weighing is +accurate to one-hundredth of a gram, whereas the displacement method +was accurate to only a tenth of a gram. + +In determining the percentage of the weight of a muscle in relation to +the total weight of the bird, the weight of the muscle was used as the +numerator, and the weight of the preserved specimen was used as the +denominator. Before weights were taken, all specimens were plucked in +identical fashion. + + +_Caudal Muscles._--The muscles of the caudal area that were used for +comparison were the levator caudae and the lateralis caudae. These +muscles are used by the living bird to maintain the position of the +pygostyle and therefore the rectrices; these muscles are especially +important to those birds that utilize the tail as a rudder in flight +and as a brake. As may be seen by reference to Table 11, the two +muscles are largest in proportion to body weight in the +Ptilogonatinae, in which subfamily the species have long rectrices and +must have correspondingly well-developed muscles in order to utilize +the rectrices to best advantage in flight. The lateralis caudae +differs more according to species than does the levator caudae, +showing that rudder action of the tail is of primary importance in the +adaptation for capturing insects. It will be remembered that the +pygostyle in this subfamily has a flattened lateral surface for +attachment of the levator caudae muscle, and it is therefore to be +expected that this muscle will be larger in the Ptilogonatinae than it +is in either the Bombycillinae or the Dulinae. The levator coccygis, +together with the two muscles mentioned above, is responsible for +elevation of the tail. The levator coccygis is less altered in +different species of the family than is the lateralis caudae. It may +be noted that the caudal muscles of _Dulus_ and _Bombycilla_ +constitute a smaller percentage of the total weight of the bird than +in any of the genera in the subfamily Ptilogonatinae. + + + [Illustration: Fig. 48. Caudal musculature, of _Phainopepla nitens + lepida_, in dorsal view. × 2. + + a. Levator coccygis; b. Levator caudae; c. Lateralis caudae; + d. Lateralis coccygis; e. oil gland; f. dorsal tip of pygostyle.] + + + Table 11. Caudal Muscles (Actual and Relative Weights) + + ============================================= + Species | Levator | Lateralis + ------------------------+---------+---------- + Ptilogonys caudatus | .145g. | .022g. + | .092% | .045% + | | + Ptilogonys cinereus | .030g. | .010g. + | .076% | .026% + | | + Phainopepla nitens | .025g. | .008g. + | .096% | .029% + | | + Phainoptila melanoxantha| .040g. | .015g. + | .063% | .014% + | | + Dulus dominicus | .028g. | .006g. + | .063% | .014% + | | + Bombycilla garrula | .034g. | .010g. + | .048% | .014% + | | + Bombycilla cedrorum | .026g. | .008g. + | .050% | .014% + --------------------------------------------- + + + Table 12. Weights of Muscles (These percentages expressed in terms + of weights of the body) + + Key to Table + A) Deltoid + B) Thigh + C) Peronus + D) Gastrocnemius + + ==================================================================== + Species |P. major|P. minor| A | B | C | D + ---------------+--------+--------+--------+--------+-------+-------- + Ptilogonys | 2.42g. | .29g. | .55g. | | | + caudatus | 4.94% | .59 | 1.12% | .43g. | .15g. | + | | | | .88% | .31% | .96% + Ptilogonys | 2.19g. | .28g. | .53g. | | | + cinereus | 5.57% | .71% | 1.35% | .30g. | .08g. | + | | | .71% | .21% | 1.02% + Phainopepla | 1.30g. | .20g. | .30g. | | | + nitens | 4.99% | .77% | 1.15% | .28g. | .10g. | + | | | | 1.12% | .40% | 1.42% + Phainoptila | 3.93g. | .44g. | .92g. | | | + melanoxantha | 6.18% | .69% | 1.45% | 1.09g. | .48g. | + | | | | 1.61% | .75% | 2.97% + Dulus | 2.09g. | .22g. | .50g. | | | + dominicus | 4.81% | .50% | 1.15% | .73g. | .18g. | + | | | | 1.68% | .41% | 1.01% + Bombycilla | 3.85g. | .45g. | .55g. | | | + garrula | 5.31% | .62% | .76% | .50g. | .15g. | + | | | | .69% | .18% | .59% + Bombycilla | 2.58g. | .35g. | .50g. | | | + cedrorum | 5.00% | .68% | .97% | .37g. | .10g. | + | | | | .73% | .19% | .83% + ---------------+--------+--------+--------+--------+-------+-------- + + +_Pectoral Muscles._--The pectoral set of muscles varies but little in +the family; flight power is seemingly not dependent upon size of +either the pectoralis major or pectoralis minor. The data indicate +that the insertion on the humerus, with consequent changes in the +relative length of that bone, is more significant in type of flight +and over-all flight power than is the actual size of the muscle mass. +The deltoid muscle, for example, is smaller in _Bombycilla_ than in +members of the other two subfamilies. The humerus in _Bombycilla_ is +shortened, and the muscle therefore does not need to be large to +accomplish the same powerful stroke that would be accomplished by a +longer humerus and a larger, more powerful deltoid muscle. In the case +of the deltoid, the shortening of the humerus and the more complex +arrangement of the points of insertion have obviated the necessity of +enlarging the muscle. + + +_Leg Musculature._--The muscles of the thigh are noticeably larger in +birds that have long leg bones. (See Table 12 for size of muscles.) On +the tibiotarsus, the peroneus and gastrocnemius muscles were measured. +When expressed as a percentage of the weight of the bird, the peroneus +has much the same relative weight in all but one of the species, +whereas the gastrocnemius varies much. The peroneus is proportionately +large only in _Phainoptila_, in which genus all the leg muscles are +well developed, but the gastrocnemius is larger in all the +Ptilogonatinae and in _Dulus_ than it is in the specialized +_Bombycilla_, in which it has probably been reduced as the leg bones +and other muscles have been reduced. + +The volume of the muscles of the hind limb changes more readily in +response to saltation and running than do the muscles of the forelimb +to flying. + + + + +DIGESTIVE TRACT + + +The digestive tract is relatively uniform in all genera of the family; +there are only slight differences between the species. The degree of +compactness of the visceral mass varies, _Phainoptila_ and _Ptilogonys +caudatus_ having the folds of the digestive tract loosely arranged, +whereas _Ptilogonys cinereus_ and _Phainopepla_ have folds which +adhere more tightly to the ventriculus and liver. In _Dulus_ and +_Bombycilla_, as compared with the Ptilogonatinae, the visceral mass +(primarily liver and ventriculus) is situated more posteriorly in the +body cavity, and is more compact, and the intestine is more tightly +coiled. + +The coiling of the intestine, if its degree of compactness is +disregarded, is nearly identical in the birds of the family; there are +four major loops between the ventriculus and the anus. The length of +this section of the tract is, however, somewhat variable, as can be +seen by reference to Table 13, in which the actual and relative +lengths of the intestine are given. It may be seen that in +_Bombycilla_ and in _Phainopepla_, the tracts are much shortened. This +is notable, since these are frugivorous birds, and in many frugivorous +birds, the tract is lengthened for better extraction of edible +portions of the food. Possibly the action of the digestive juices is +correspondingly more rapid in _Bombycilla_ and _Phainopepla_, thereby +permitting the necessary nutriment to be extracted by a short +digestive tract. + +In a migratory bird, or one that depends on flight power to find food +and escape capture by predators, as in the case of the waxwings, the +compacted and shortened visceral mass would seem to be advantageous, +because of the consequent reduction in weight. I consider the longer +intestine to be the ancestral condition, and that the intestine has +become shorter to meet new environmental conditions. + + + Table 13. Digestive Tract: Actual Length, and Length Relative to + Thoracic Length + + =========================+========+============== + | | Relative + Species | Length | length + | in mm. | (in percent) + -------------------------+--------+-------------- + Ptilogonys caudatus | 134 | 476.9 + Ptilogonys cinereus | 111 | 415.6 + Phainopepla nitens | 94 | 357.5 + Phainoptila melanoxantha | 150 | 457.1 + Dulus dominicus | 130 | 451.0 + Bombycilla garrula | 102 | 298.2 + Bombycilla cedrorum | 95 | 309.5 + -------------------------+--------+-------------- + + +Beddard (1898:30) states that caecae in the tract may be highly +variable in a single family of birds. The Bombycillidae is no +exception in this regard. At the junction of the cloaca and the large +intestine, there are two small caecae, the function of which is +unknown to me. The caecae are largest in the Ptilogonatinae, smaller +in the Bombycillinae, and smallest in the Dulinae. There may be a +correlation between large caecae and more insectivorous diet and small +caecae and frugivorous diet; however, the data are not conclusive in +this regard. + + + + +ORIGIN OF THE SPECIES + + +It is here postulated that the center of origin for the ancestral +stock of the Bombycillidae was in a region of North America, which at +the time concerned was temperate or possibly even semi-tropical in +climate. Probably Northern Mexico was the place and probably the +climate was temperate. It is reasonably certain, because of the +distribution of the species of the family, that they originated in the +Americas. In the absence of paleontological data (_Bombycilla_ alone +is reported, in essentially its modern form, from the late +Pleistocene--Wetmore, 1940a), the place and time of origin cannot +certainly be determined. + +The distribution of the family is such that the more primitive groups +are in the south. These are the Ptilogonatinae in Central America and +Mexico, and the isolated Dulinae in Haiti and the Dominican Republic. +This distribution would support the view that the origin was in the +south. However, the Holarctic Bombycillinae are so typically birds of +northern latitudes that, were it not for such close relatives south of +their range, it would appear logical to infer a northerly origin with +a subsequent shifting of populations both southward and northward. The +phyletic age of the family is probably great, however, as evidenced by +the spotty distribution of the birds. + +In the evolution of this family, population pressure possibly played +the initial role in forcing members of the primitive, southern stock +to seek habitable areas on the periphery of the range. Some birds +also, being possessed of the "adventuresome spirit", aided the +northerly movement, thus effecting an extension of the breeding ranges +to the north. So far as is now known, this family did not seek living +space in South America. By extending its range, a species might find +more abundant food and nesting sites. This process of extending the +range probably would be costly to the species concerned, because only +those individuals best able to adapt themselves to the new +environmental conditions would be able to survive long enough to +reproduce their kind. + +The return flight to the south could, in time, be dispensed with, +except in the coldest weather or when the local berry- and fruit-crop +failed. Birds such as waxwings are, of course, able to subsist on +dried fruits and berries in the critical winter season when strictly +insectivorous birds, not so catholic in their food habits, must return +south. It appears that waxwings are descendants of migratory birds +that have adjusted themselves to a life in the north; and they are +judged not to have evolved from year-round residents of the north. + +Even a short migratory journey in spring by part of a population of +birds, while the other part remained in the original range, would +quickly isolate one breeding population from the other, resulting in +the formation of different genetic strains that lead to subspecies, +species, and finally to genera and families. Any variation away from +the ancestral, "sedentary" stock would become established more quickly +because of such isolation at the breeding period. By the same token, +the parental stock can, and no doubt does, become modified to suit its +environment more perfectly, thus accelerating the tempo of this type +of divergent evolution. + +The original "split" of the Bombycillines is thought then to have been +the result of migration on the part of some of the ancestral stock, +with subsequent loss of regular migration because the need to return +south was lost. Early in development, and before the migrational +tendency was entirely lost, an isolated population, which later became +sedentary, as it was an island population, diverged to give rise to +the Dulinae. The Dulinae are a homogeneous group since on the islands +now inhabited by the birds, they have not been isolated sufficiently +long to produce even well-marked subspecies. + + + [Illustration: Fig. 49. Hypothetical family tree of the + Bombycillidae.] + + +The present day _Phainoptila_ is most nearly like the ancestral group, +and the remainder of the Ptilogonatinae have diverged to fit +conditions similar to those to which the Tyrannid flycatchers, which +parallel them, are also fitted. + +In comparatively recent geological time, two basic lines developed +from the Bombycilline stock, the future _B. garrula_ and _B. +cedrorum_. Possibly _garrula_ originally was isolated in Europe and +Asia, and later came into contact with _B. cedrorum_, following the +time at which the two species were genetically well differentiated. It +appears certain that _B. japonica_ was an offshoot of the Bombycilline +stock at an early time, since it has characteristics that seem +relatively unspecialized. It possibly was isolated in the Orient. + +Structural affinities of _Dulus_ and _Bombycilla_ are more pronounced +than are those of _Dulus_ and _Ptilogonys_, for example. Many of the +structural features of _Dulus_ parallel those of _Phainoptila_, and it +seems likely that the Dulinae were separated early in the history of +the family, perhaps as an isolated offshoot of the early migratory +Bombycillinae. + + + + +CONCLUSIONS + + +Nomenclature, as used by a taxonomist, should of course indicate +affinities as well as apply a name, and the rank of the family should +be applied to a structural unit based on common anatomical characters +that are more fundamental than, in my opinion, are those used by +Ridgway (1904) in proposing family status for the silky flycatchers +and the palm-chats. The characters in the diagnosis (page 478) of the +family Bombycillidae are common features regarded as warranting a +single family unit for the waxwings, silky flycatchers, and +palm-chats. The differences in morphology used by previous workers to +characterize each of these groups: (1) the silky flycatchers; (2) +waxwings and; (3) palm-chats are regarded as more properly characters +of only subfamily rank. + +The existing coloration of the species of the Bombycillidae appears to +have been acquired relatively late, geologically speaking. The three +subfamilies responded to ecological stimuli in three different ways, +and the resulting color patterns are unlike in the three groups. +Dulinae to this day have a color pattern that is most like the +ancestral color pattern, and this is recapitulated in the juvenal +plumage of the Bombycillinae before they attain their adult plumage. + +Consideration of the geographic distribution of the species of the +family indicates that the center of origin of the family Bombycillidae +was south of the present range of the waxwings (subfamily +Bombycillinae). Waxwings probably are the descendants of a migratory +population that diverged from the primitive population at an early +time in the history of the family. Owing to their adaptations to +survive in the north, waxwings no longer return south in the autumn. +Palm-chats (subfamily Dulinae) are descendants of an isolated +population of the family stock that developed communal living habits +as one specialization. Silky Flycatchers (subfamily Ptilogonatinae) +became modified to catch insects, and have specializations that +roughly parallel those of the Tyrannid flycatchers. + +Osteologically, the various species of the Bombycillidae are +remarkably similar. Small variations do exist, but these are primarily +differences in relative size. The modifications of the beak enable +palm-chats to feed on parts of plants, and the beak of _Phainoptila_ +shows some similarity in this respect. Rounded wings, which cause a +bird to fly by means of short, relatively weak strokes, are correlated +with a comparatively long humerus, whereas long and pointed wings, +which enable a bird to fly with more powerful strokes of the wing, are +correlated with a relatively short humerus. There is a positive +correlation between a short humerus and a long external condyle, and +between a long humerus and the absence or smallness of the external +condyle. + +In the Bombycillidae short bones of the leg are adaptive, and long +bones of the leg are the generalized condition. Although all passerine +birds were differentiated relatively late in geologic time, long hind +limbs still could have been present in the immediate ancestors of +passerine birds. As adaptive radiation took place in the class Aves, +some birds, the Bombycillidae included, became more and more adapted +for an arboreal, and eventually an aerial habitat, with consequent +loss of saltatorial and running ability. + +Birds, like mammals, have a short femur, the most proximal element in +the leg, if the species is adapted to run fast. If the species is not +adapted to run fast, birds, unlike mammals, have the tibiotarsus +longer than any of the other elements; in mammals that are not adapted +to run fast, the femur and tibia are approximately the same length. In +non-running birds as compared with running birds, the leg element +distal to the tibiotarsus, and the one proximal to it, are +considerably shortened. In waxwings, all three elements of the hind +limb are shortened, indicating that the reduction in length has been, +evolutionarily speaking, a rapid process, in order to reduce the limbs +to a convenient size as soon as possible. + +The shape of the pygostyle varies in the Bombycillidae, but the simple +shieldlike bone of _Phainoptila_ is judged to resemble closely the +ancestral type. In _Ptilogonys_ there is a tall dorsal spine, coupled +with a wide and heavy centrum and flattened lateral areas, for support +of the long rectrices. In _Bombycilla_ the bone is small with knobs on +the centrum that have been developed for muscle attachment. + +The muscles were carefully dissected in each genus and in most of the +species. The same homologous muscles are present in all species. +Significant differences were found only in the relative size of +certain muscles. No satisfactorily accurate method of measuring these +differences was found. Consequently, less use was made of the results +of the dissections than was originally planned. + +The set of pectoral muscles varies but slightly in relative mass, and +the variation is not considered significant. The deltoid muscle was +selected for measurement since its point of insertion is unusually +variable, while the mass of the muscle varies little. We can conclude +that the extent of the area of insertion of the tendon of a muscle can +determine that muscle's relative efficiency, while the muscle itself +remains the same in bulk. + +The muscles of the hind limb are notably larger in species that have +long legs, and a good index of the hopping ability may be gained by +study of certain of these muscles. In the Bombycillidae, and in those +Ptilogonatinae that do not use the hind limbs for hopping, the bones +are shortened, and the associated muscles are correspondingly smaller. + +The gross anatomy of the digestive tract is practically identical in +the members of the family. The variability noted is mainly in the +degree of compactness of the visceral mass in _Bombycilla_ and in +_Phainopepla_. Also there is a tendency for the Bombycillinae and the +Dulinae to have the mass situated more posteriorly than it is in the +Ptilogonatinae. Moreover, _Bombycilla_ has a shorter intestine than do +the other genera. All of this indicates that the waxwings +(Bombycillinae) have the center of gravity situated more +advantageously for flight than do the birds of the two other +subfamilies. + + + + +SUMMARY + + +1. The silky flycatchers, waxwings, and palm-chats are included in the +family Bombycillidae; the Ptilogonatidae and Dulidae are reduced to +subfamily rank. + +2. The coloration of the birds of each subfamily is different because +the ecological needs are different. + +3. Waxwings were at one time regularly migratory, but are now nomadic, +since they are adapted to live in northern latitudes for the entire +year. + +4. The corresponding bones in different members of the family closely +resemble one another, and the differences which do exist are the results +of responses within relatively recent times to changes in habits. + +5. In the Bombycillidae a rounded wing is judged to be the primitive +condition. As the wing becomes more pointed, the humerus becomes shorter +and its external condyle longer. + +6. The hind limbs are short in birds that depend most on flight power, +but are longer and the distal elements are disproportionately longer in +birds that depend on saltation or on running. + +7. The pygostyle varies in shape and size between genera and even +between some species. + +8. The pectoral muscles differ in size only slightly in the different +members of the family, but the insertions are more extensive for these +muscles in birds that fly a great deal. + +9. The muscles of the hind limb vary in mass, but not in kind, in the +members of the family Bombycillidae. + +10. In the Bombycillidae that depend on flight power, rather than on +saltation or on running power, there is a tendency for the digestive +tract to become shorter and for the whole visceral mass to become more +compact. + + + + +BIBLIOGRAPHY + + +ANDERSON, E. M. + + 1915. Nesting of the Bohemian Waxwing in northern British + Columbia. Condor, 17(4):145-148, 1915. + +ANDERSON, M. P. + + 1907. A collecting trip in Korea. Condor, 9(5):146-147, 1907. + +ANDERSON, R. M. + + 1909. Nesting of the Bohemian Waxwing (_Bombycilla garrulus_). + Auk, 26(1):10-12, 1909. + +ARMSTRONG, E. A. + + 1942. Bird display. Cambridge Univ. Press, xvi + 381 pp., + 22 plates, 1942. + +BAIRD, S. F. + + 1860. The birds of North America. J. B. Lippincott Co., lvi + + 1003 pp., 1860. + +BEDDARD, F. E. + + 1898. The structure and classification of birds. Longmans, Green + & Co., xx + 548 pp., 252 figs., 1898. + +BERGTOLD, W. H. + + 1917a. A study of the incubation period of birds. 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Zoöl., 24(2):125-314, 8 plates, 34 figs. in text, 1922. + +TAYLOR, W. P. + + 1918. Bohemian Waxwing (_Bombycilla garrulus_) breeding within + the United States. Auk, 35(2):226-227, 1918. + +TAVERNER, P. A. + + 1934. Birds of Canada. Nat. Mus. Canada Bull., 72, series 19, + 445 pp., 77 plates, 488 figs. in text, 1934. + +WAYNE, A. T. + + 1924. A remarkable Cedar Waxwing. Auk, 41(3):485, 1924. + +WETMORE, A. + + 1926. The migrations of birds. Cambridge, Harvard Univ. Press, + vii + 217 pp., 1926. + + 1932. Notes from Dr. R. Ciferri on the birds of Hispaniola. Auk, + 49(1):101-108, 1931. + + 1940a. A check-list of the fossil birds of North America. Smithson. + Misc. Coll., 99(4):1-88 pp., 1940. + + 1940b. A systematic classification of the birds of the world. + Smithson. Misc. Coll., 99(7):1-11 pp., 1940. + +WETMORE, A., and SWALES, B. H. + + 1931. The birds of Haiti and the Dominican Republic. U. S. Nat. + Mus. Bull. 155:iv + 482 pp., 26 plates, 1931. + +WHEELOCK, I. G. + + 1905. Regurgitation feeding of nestlings. Auk, 22(1):54-71, 1905. + +WHITTLE, H. G. + + 1928. The biography of a Cedar Waxwing. Bull. NE Bird-Band. Assoc., + 4:77-85, 1928. + +WOLFSON, A. + + 1945. The role of the pituitary, fat deposition, and body weight + in bird migration. Condor, 47(3):95-127, 1945. + +WOLLEY, J. J. + + 1857. On the nest and eggs of the Waxwing (_Bombycilla garrula_ + Tamm.). Proc. Zoöl. Soc. London, 25:55-56, 1857. + + + _Transmitted July 29, 1949._ + + +Mention should be made here of an important paper by Jean Delacour and +Dean Amadon (1949). The Relationships of _Hypocolius_ (Ibis, +91:427-429, plates 19 and 20) which appeared after the present paper +by Arvey was written. Delacour and Amadon stated that _Hypocolius_, a +monotypic Persian genus, should be assigned to the Bombycillidae. +Their conclusions (_op. cit._:429) were as follows: "It might be +advisable to set up three subfamilies in the Bombycillidae, one for +_Bombycilla_, one for _Hypocolius_, and a third for the silky +flycatchers, _Ptilogonys_, _Phainopepla_ and _Phainoptila_. Further +study may show that _Dulus_ can be added as a fourth subfamily. + +"Previously the Bombycillidae appeared to be an American group of +which one genus (_Bombycilla_) had reached the Old World. Inclusion of +_Hypocolius_ in the family makes this theory uncertain. Without +obvious affinities to other families, and consisting of a small number +of scattered and rather divergent genera, the Bombycillidae would seem +to be a declining group whose origin cannot safely be deduced from the +distribution of the few existing species." + + --Eds. + + + 23-1019 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + + +The University of Kansas Publications, Museum of Natural History, are +offered in exchange for the publications of learned societies and +institutions, universities and libraries. For exchanges and information, +address the EXCHANGE DESK, UNIVERSITY OF KANSAS LIBRARY, LAWRENCE, +KANSAS, U. S. A. + +MUSEUM OF NATURAL HISTORY.--E. Raymond Hall, Chairman, Editorial +Committee. + +This series contains contributions from the Museum of Natural History. +Cited as Univ. Kans. Publ., Mus. Nat. Hist. + + + Vol. 1. 1. The pocket gophers (genus Thomomys) of Utah. By Stephen D. + Durrant. Pp. 1-82, 1 figure in text. August 15, 1946. + + 2. The systematic status of Eumeces pluvialis Cope, and + noteworthy records of other amphibians and reptiles from + Kansas and Oklahoma. By Hobart M. Smith. Pp. 85-89. + August 15, 1946. + + 3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. + Pp. 93-96, 1 figure in text. August 15, 1946. + + 4. Hybridization between two species of garter snakes. + By Hobart M. Smith. Pp. 97-100. August 15, 1946. + + 5. Selected records of reptiles and amphibians from Kansas. + By John Breukelman and Hobart M. Smith. Pp. 101-112. + August 15, 1946. + + 6. Kyphosis and other variations in soft-shelled turtles. + By Hobart M. Smith. Pp. 117-124. July 7, 1947. + + 7. Natural history of the prairie vole (Mammalian genus + Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures + in text. October 6, 1947. + + 8. The postnatal development of two broods of great horned + owls (Bubo virginianus). By Donald F. Hoffmeister and + Henry W. Setzer. Pp. 157-173, 5 figures in text. + October 6, 1947. + + 9. Additions to the list of the birds of Louisiana. + By George H. Lowery, Jr. Pp. 177-192. November 7, 1947. + + 10. A check-list of the birds of Idaho. By M. Dale Arvey. + Pp. 193-216. November 29, 1947. + + 11. Subspeciation in pocket gophers of Kansas. By Bernardo + Villa-R. and E. Raymond Hall. Pp. 217-236, 2 figures in + text. November 29, 1947. + + 12. A new bat (genus Myotis) from Mexico. By Walter W. Dalquest + and E. Raymond Hall. Pp. 237-244, 6 figures in text. + December 10, 1947. + + 13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. + By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, + 1 figure in text. December 10, 1947. + + 14. A new pocket gopher (Thomomys) and a new spiny pocket + mouse (Liomys) from Michoacán, Mexico. By E. Raymond Hall + and Bernardo Villa-R. Pp. 249-256, 6 figures in text. + July 26, 1948. + + 15. A new hylid frog from eastern Mexico. By Edward H. Taylor. + Pp. 257-264, 1 figure in text. August 16, 1948. + + 16. A new extinct emydid turtle from the Lower Pliocene of + Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. + August 16, 1948. + + 17. Pliocene and Pleistocene records of fossil turtles from + western Kansas and Oklahoma. By Edwin C. Galbreath. + Pp. 281-284, 1 figure in text. August 16, 1948. + + 18. A new species of heteromyid rodent from the Middle + Oligocene of northeastern Colorado with remarks on the + skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. + August 16, 1948. + + 19. Speciation in the Brazilian spiny rats (genus Proechimys, + Family Echimyidae). By João Moojen. Pp. 301-406, + 140 figures in text. December 10, 1948. + + 20. Three new beavers from Utah. By Stephen D. Durrant and + Harold S. Crane. Pp. 407-417, 7 figures in text. + December 24, 1948. + + 21. Two new meadow mice from Michoacán, México. By E. Raymond + Hall. Pp. 423-427, 6 figures in text. December 24, 1948. + + 22. An annotated check list of the mammals of Michoacán, + México. By E. Raymond Hall and Bernardo Villa R. + Pp. 431-472, 5 figures in text. December 27, 1949. + + 23. Subspeciation in the kangaroo rat, Dipodomys ordii. + By Henry W. Setzer. Pp. 473-573, 27 figures in text, + 7 tables. December 27, 1949. + + 24. Geographic range of the hooded skunk, Mephitis macroura, + with description of a new subspecies from Mexico. + By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, + 1 figure in text. January 20, 1950. + + 25. Pipistrellus cinnamomeus Miller 1902 referred to the genus + Myotis. By E. Raymond Hall and Walter W. Dalquest. + Pp. 581-590, 5 figures in text. January 20, 1950. + + 26. A synopsis of the American bats of the genus Pipistrellus. + By E. Raymond Hall and Walter W. Dalquest. Pp. 591-602, + 1 figure in text. January 20, 1950. + + Index. Pp. 605-638. + + + Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. + Pp. 1-444, 140 figures in text. April 9, 1948. + + + Vol. 3. 1. The Avifauna of Micronesia, its origin, evolution, and + distribution. By Rollin H. Baker. Pp. 1-359, 16 figures in + text. June 12, 1951. + + 2. A Quantitative study of the nocturnal migration of birds. + By George H. Lowery, Jr. Pp. 361-472, 46 figures in text. + June 29, 1951. + + 3. Phylogeny of the waxwings and allied species. By M. Dale + Arvey. Pp. 473-530, 49 figures in text, 13 tables. + October 10, 1951. + + * * * * * + + +Transcriber's Notes: + +The text herein presented was derived from scans of the original report +which were OCRed and proofread. Minor typographical errors (genus name +initial not italicized, missing parenthis, missing or superfluous +commas, etc.) were made but are not noted here. With the exception of +those corrections and those noted below, it is the same text. + + +Typographical Corrections + + Page 481 : Measureemnts => Measurements + + Page 486 : cedorum => cedrorum + + Page 496, Fig. 11 : Luis => Luís + + Page 480, 481 : Luis Potosí => Luís Potosi + + Page 516, Table 12 : Gatrocnemius => Gastrocnemius + + +Emphasis Notation: + + _text_ : italicized + + =text= : bold + + * * * * * + + + + + +End of the Project Gutenberg EBook of Phylogeny of the Waxwings and Allied +Birds, by M. 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