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+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.
+
+
+
+
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+
+
+  _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. Dale Arvey
+
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+    .technt      {background:#d0d0d0; padding: 7px; border:solid black 1px;}
+
+    .vtop         {vertical-align: top;}
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+
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+  </head>
+<body>
+
+
+<pre>
+
+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
+
+
+
+
+
+
+</pre>
+
+
+
+<a name="typos"></a>
+<div class="technt">
+<div class="caption2">Transcriber's Notes</div>
+
+<p>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.</p>
+
+<p>One additional note, many of the figures list notation such as "&times; 1/2"
+to denote that the image is shown at half the actual size. The images
+reproduced herein most likely will not match the original printed scale due
+to the display resolution used by the viewer.</p>
+
+<div class="caption2">Typographical Corrections</div><br>
+
+<table summary="typo_list">
+<tr><td>Page 480 :</td><td><a href="#Luis_Potosi">Luis Potos&iacute; =&gt; Lu&iacute;s Potosi</a></td></tr>
+<tr><td>Page 481 :</td><td><a href="#Luis_Potosi2">Luis Potos&iacute; =&gt; Lu&iacute;s Potosi</a></td></tr>
+<tr><td>Page 481 :</td><td><a href="#Measurements">Measureemnts =&gt; Measurements</a></td></tr>
+<tr><td>Page 486 :</td><td><a href="#cedrorum">cedorum =&gt; cedrorum</a></td></tr>
+<tr><td>Page 496 :</td><td><a href="#Luis">Luis =&gt; Lu&iacute;s</a></td></tr>
+<tr><td>Page 516 :</td><td><a href="#Gastrocnemius">Gatrocnemius =&gt; Gastrocnemius</a></td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<hr style="width: 45%;">
+<div class="cover">
+<p><span class="pagenum"><a name="Cover" id="Cover">[Cover]</a></span></p>
+
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="caption2">Phylogeny of the Waxwings<br>
+and Allied Birds</div><br>
+<br>
+<div class="caption3">BY<br>
+M. DALE ARVEY</div><br>
+<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="caption3">University of Kansas Publications<br>
+Museum of Natural History</div><br>
+<br>
+<h4>Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables<br>
+October 10, 1951</h4><br>
+<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<h4>University of Kansas<br>
+LAWRENCE<br>
+1951</h4><br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_473" id="Page_473">[Pg&nbsp;473]</a></span></p>
+
+<div class="center">
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="caption2">Phylogeny of the Waxwings<br>
+and Allied Birds</div><br>
+<br>
+<div class="caption3">BY<br>
+M. DALE ARVEY</div><br>
+<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="caption3">University of Kansas Publications<br>
+Museum of Natural History</div><br>
+<br>
+<h4>Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables<br>
+October 10, 1951</h4><br>
+<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<h4>University of Kansas<br>
+LAWRENCE<br>
+1951</h4><br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<p><span class="pagenum"><a name="Page_474" id="Page_474">[Pg&nbsp;474]</a></span></p>
+<div class="center">
+<div class="caption3">
+<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br>
+<br>
+Editors: E. Raymond Hall, Chairman, Edward H. Taylor,<br>
+A. Byron Leonard, Robert W. Wilson<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables<br>
+<br>
+Published October 10, 1951<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<span class="smcap">University of Kansas</span><br>
+Lawrence, Kansas<br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="caption4">
+PRINTED BY<br>
+FERD VOILAND, JR., STATE PRINTER<br>
+TOPEKA, KANSAS<br>
+1950<br>
+<img src="images/union_label.png" width="71" height="26" border="0" alt="Look for the Union Label" title="Look for the Union Label"><br>
+23-1019<br>
+</div>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_475" id="Page_475">[Pg&nbsp;475]</a></span></p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="caption2">
+Phylogeny of the Waxwings<br>
+and Allied Birds<br>
+</div>
+
+<div class="caption3">
+by<br>
+M. DALE ARVEY<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="TOC"></a>
+<div class="caption2">CONTENTS</div>
+<table width="100%" summary="Table of Contents">
+<tr><td class="blank">&nbsp;</td><td class="text_rt blank">PAGE</td></tr>
+<tr><td><a href="#INTRODUCTION"><span class="smcap blank">Introduction</span></a></td><td class="text_rt blank">476</td></tr>
+<tr><td><a href="#ACKNOWLEDGMENTS"><span class="smcap blank">Acknowledgments</span></a></td><td class="text_rt blank">476</td></tr>
+<tr><td><a href="#NOMENCLATURAL_HISTORY"><span class="smcap blank">Nomenclatural History</span></a></td><td class="text_rt blank">477</td></tr>
+<tr><td><a href="#MATERIALS"><span class="smcap blank">Materials</span></a></td><td class="text_rt blank">478</td></tr>
+<tr><td><a href="#DIAGNOSES"><span class="smcap blank">Diagnoses</span></a></td><td class="text_rt blank">478</td></tr>
+<tr><td><a href="#COLORATION"><span class="smcap blank">Coloration</span></a></td><td class="text_rt blank">485</td></tr>
+<tr><td><a href="#COURTSHIP"><span class="smcap blank">Courtship</span></a></td><td class="text_rt blank">489</td></tr>
+<tr><td><a href="#NEST_BUILDING"><span class="smcap blank">Nest Building</span></a></td><td class="text_rt blank">491</td></tr>
+<tr><td><a href="#FOOD"><span class="smcap blank">Food</span></a></td><td class="text_rt blank">493</td></tr>
+<tr><td><a href="#SKELETON"><span class="smcap blank">Skeleton</span></a></td><td class="text_rt blank">494</td></tr>
+<tr><td><span style="margin-left: 1em;" class="smcap blank"><a href="#SKULL">Skull</a></span></td><td class="text_rt blank">494</td></tr>
+<tr><td><span style="margin-left: 1em;" class="smcap blank"><a href="#HUMERUS">Humerus</a></span></td><td class="text_rt blank">499</td></tr>
+<tr><td><a href="#PYGOSTYLE"><span style="margin-left: 1em;" class="smcap blank">Pygostyle</span></a></td><td class="text_rt blank">502</td></tr>
+<tr><td><a href="#STERNUM"><span style="margin-left: 1em;"><span class="smcap blank">Sternum</span></span></a></td><td class="text_rt blank">505</td></tr>
+<tr><td><a href="#RELATIVE_LENGTH"><span style="margin-left: 1em;" class="smcap blank">Relative Lengths of Bones</span></a></td><td class="text_rt blank">505</td></tr>
+<tr><td><span style="margin-left: 2em;" class="smcap blank"><a href="#LEG-TRUNK">Leg-trunk Percentages</a></span></td><td class="text_rt blank">509</td></tr>
+<tr><td><span style="margin-left: 2em;" class="smcap blank"><a href="#ARM-TRUNK">Arm-trunk Percentages</a></span></td><td class="text_rt blank">511</td></tr>
+<tr><td><a href="#MUSCULATURE"><span class="smcap blank">Musculature</span></a></td><td class="text_rt blank">514</td></tr>
+<tr><td><a href="#CAUDAL"><span style="margin-left: 1em;"><span class="smcap blank">Caudal Muscles</span></span></a></td><td class="text_rt blank">514</td></tr>
+<tr><td><a href="#PECTORAL"><span style="margin-left: 1em;"><span class="smcap blank">Pectoral Muscles</span></span></a></td><td class="text_rt blank">517</td></tr>
+<tr><td><a href="#HIND_LIMB"><span style="margin-left: 1em;"><span class="smcap blank">Hind Limb Musculature</span></span></a></td><td class="text_rt blank">517</td></tr>
+<tr><td><a href="#DIGESTIVE_TRACT"><span class="smcap blank">Digestive Tract</span></a></td><td class="text_rt blank">517</td></tr>
+<tr><td><a href="#ORIGIN_OF_THE_SPECIES"><span class="smcap blank">Origin of the Species</span></a></td><td class="text_rt blank">519</td></tr>
+<tr><td><a href="#CONCLUSIONS"><span class="smcap blank">Conclusions</span></a></td><td class="text_rt blank">521</td></tr>
+<tr><td><a href="#SUMMARY"><span class="smcap blank">Summary</span></a></td><td class="text_rt blank">524</td></tr>
+<tr><td><a href="#BIBLIOGRAPHY"><span class="smcap blank">Bibliography</span></a></td><td class="text_rt blank">525</td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="INTRODUCTION" id="INTRODUCTION"></a>
+<p><span class="pagenum"><a name="Page_476" id="Page_476">[Pg&nbsp;476]</a></span></p>
+<div class="caption2">INTRODUCTION</div>
+
+<p>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.</p>
+
+<p>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 (<a href="#ORIGIN_OF_THE_SPECIES">see p.
+519.</a>) 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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="ACKNOWLEDGMENTS" id="ACKNOWLEDGMENTS"></a>
+<div class="caption2">ACKNOWLEDGMENTS</div>
+
+<p>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,
+<span class="pagenum"><a name="Page_477" id="Page_477">[Pg&nbsp;477]</a></span>
+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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="NOMENCLATURAL_HISTORY" id="NOMENCLATURAL_HISTORY"></a>
+<div class="caption2">NOMENCLATURAL HISTORY</div>
+
+<p>The oldest name available for any species of the waxwings is
+<i>Lanius garrulus</i> Linnaeus (1758). <i>Lanius garrulus</i> and <i> Lanius garrulus</i>
+variety B <i>carolinensis</i> were described as conspecific. The description
+has been associated with the first of the two names. The
+latter name is a <i>nomen nudum</i> since it was not accompanied by a
+separate description. The generic name <i>Lanius</i> 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 <i>Bombycilla</i>, 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 <i>Ampelis</i> for the
+Bohemian Waxwing, and combined it with the specific name <i>garrulus</i>,
+the Cedar Waxwing being termed variety B. Vieillot (1807)
+proposed the generic name <i>Bombycilla</i> and combined it with a new
+specific name, <i>cedrorum</i>, for the Cedar Waxwing. Vieillot has been
+cited as the author of <i>Bombycilla</i> since that time, although Brisson
+used <i>Bombycilla</i> 33 years before. Oberholser (1917) did not cite
+Brisson's work in his discussion of the proper generic name for the
+waxwings, and <i>Bombycilla</i> 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 <i>Bombycilla cedrorum</i> Vieillot (1807).</p>
+
+<p>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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="MATERIALS" id="MATERIALS"></a>
+<p><span class="pagenum"><a name="Page_478" id="Page_478">[Pg&nbsp;478]</a></span></p>
+<div class="caption2">MATERIALS</div>
+
+<p>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.</p>
+
+<table class="center" width="100%" summary="Species Specimen Counts">
+<tr><td><span class="smcap">Species&nbsp;or&nbsp;Subspecies</span></td><td>Skin</td><td>Skeleton</td><td>Alcoholic</td></tr>
+<tr><td colspan=4><hr></td></tr>
+<tr><td class="text_lf"><i>Phainoptila melanoxantha melanoxantha</i></td><td>8</td><td>1</td><td>2</td></tr>
+<tr><td class="text_lf"><i>Phainoptila melanoxantha minor</i>&nbsp;</td><td>2</td><td>&nbsp;</td><td>&nbsp;</td></tr>
+<tr><td class="text_lf"><i>Ptilogonys cinereus cinereus</i>&nbsp; &nbsp;</td><td>13</td><td>3</td><td>4</td></tr>
+<tr><td class="text_lf"><i>Ptilogonys cinereus molybdophanes</i></td><td>6</td><td>&nbsp;</td><td>&nbsp;</td></tr>
+<tr><td class="text_lf"><i>Ptilogonys caudatus</i></td><td>16</td><td>3</td><td>4</td></tr>
+<tr><td class="text_lf"><i>Phainopepla nitens nitens</i></td><td>&nbsp;</td><td>1</td><td>5</td></tr>
+<tr><td class="text_lf"><i>Phainopepla nitens lepida</i></td><td>12</td><td>5</td><td>4</td></tr>
+<tr><td class="text_lf"><i>Bombycilla cedrorum</i></td><td>53</td><td>27</td><td>8</td></tr>
+<tr><td class="text_lf"><i>Bombycilla garrula garrula</i>&nbsp; &nbsp; &nbsp;</td><td>4</td><td>3</td><td>&nbsp;</td></tr>
+<tr><td class="text_lf"><i>Bombycilla garrula centralasiae</i></td><td>9</td><td>2</td><td>&nbsp;</td></tr>
+<tr><td class="text_lf"><i>Bombycilla garrula pallidiceps</i>&nbsp;</td><td>7</td><td>3</td><td>2</td></tr>
+<tr><td class="text_lf"><i>Bombycilla japonica</i></td><td>10</td><td>&nbsp;</td><td>&nbsp;<br>
+<tr><td class="text_lf"><i>Dulus dominicus dominicus</i></td><td>9</td><td>5</td><td>2</td></tr>
+<tr><td class="text_lf"><i>Dulus dominicus oviedo</i>&nbsp; &nbsp;</td><td>4</td><td>1</td><td>&nbsp;</td></tr>
+<tr><td colspan=4><hr></td></tr>
+<tr><td class="text_lf">Totals&nbsp; &nbsp;</td><td>153</td><td>54</td><td>31</td></tr>
+<tr><td colspan=4><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="DIAGNOSES" id="DIAGNOSES"></a>
+<div class="caption2">DIAGNOSES</div><br>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Family Bombycillidae</div>
+
+<p><i>Diagnosis.</i>&mdash;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 <i>Dulus</i> and <i>Phainoptila</i>);
+tarsus generally shorter than middle toe and claw, distinctly
+scutellate with five or six divisions, the lateral plate subdivided
+(except in <i>Phainoptila</i>); 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 <i>Dulus</i>); eggs spotted; nest in trees; three
+subfamilies, five genera, eight species.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Subfamily Ptilogonatinae</div>
+
+<p><i>Diagnosis.</i>&mdash;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,
+<span class="pagenum"><a name="Page_479" id="Page_479">[Pg&nbsp;479]</a></span>
+with small external condyle; plumage soft and silky, less so in
+<i>Phainoptila</i>; sexes dissimilar, young like adult female; three
+genera, four species.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Genus <b>Phainoptila</b> Salvin</div>
+
+<div class="blockquot smaller"><p><i>Phainoptila</i> Salvin, Proc. Zoöl. Soc. London, 1877:367, April 17,
+1877. Type <i>Phainoptila melanoxantha</i> Salvin.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Range.</i>&mdash;Costa Rica and Panam&aacute;.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Phainoptila melanoxantha melanoxantha</b> Salvin</div>
+<br>
+<div class="center">Phainoptila</div>
+
+<div class="blockquot smaller"><p><i>Phainoptila melanoxantha melanoxantha</i> Salvin, Proc. Zoöl. Soc.
+London, 1877:367; April 17, 1877. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 99.0, tail 88.5, culmen 15.2, tarsus 28.4.</p>
+
+<p><i>Range.</i>&mdash;Highlands of Costa Rica and extreme western Panam&aacute; (Volc&aacute;n
+de Chiriqu&iacute;).</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Phainoptila melanoxantha minor</b> Griscom</div>
+<br>
+<div class="center">Phainoptila</div>
+
+<div class="blockquot smaller"><p><i>Phainoptila melanoxantha minor</i> Griscom, Amer. Mus. Novitates,
+141:7, 1924. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration as in <i>P. m. melanoxantha</i>, 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 <i>P. m. melanoxantha</i>.</p>
+
+<p><i>Range.</i>&mdash;Highlands of westeran Panam&aacute; (Cerro Flores and eastern Chiriqu&iacute;).</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Genus <b>Ptilogonys</b> Swainson</div>
+
+<div class="blockquot smaller"><p><i>Ptilogonys</i> Swainson, Cat. Bullock's Mex. Mus., App. 4, 1824. Type
+<i>Ptilogonys cinereus</i> Swainson. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Range.</i>&mdash;Southwestern United States to Costa Rica.</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_480" id="Page_480">[Pg&nbsp;480]</a></span></p>
+
+<div class="caption3nb"><b>Ptilogonys cinereus cinereus</b> Swainson</div>
+<br>
+<div class="center">Ashy Ptilogonys</div>
+
+<div class="blockquot smaller"><p><i>Ptilogonys cinereus cinereus</i> Swainson, Cat. Bullock's Mex. Mus.,
+App. 4, 1824. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Measurements.</i>&mdash;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.</p>
+
+<p><i>Range.</i>&mdash;Mountainous districts of central and southern Mexico, in
+states of Durango, Zacatecas, Hidalgo, M&eacute;xico, Oaxaca, Colima,
+Morelos, Veracruz, San <a name="Luis_Potosi"></a><a href="#typos">Lu&iacute;s Potosi</a>, Guerrero and Michoac&aacute;n.</p>
+<p><br></p>
+
+<div class="caption3nb"><b>Ptilogonys cinereus molybdophanes</b> Ridgway</div>
+<br>
+<div class="center">Ashy Ptilogonys</div>
+
+<div class="blockquot smaller"><p><i>Ptilogonys cinereus molybdophanes</i> Ridgway, Man. N. American
+Birds, 464 (footnote), 1887. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration of adult male: Upper parts darker bluish than
+in <i>P. c. cinereus</i>; venter paler; flanks Olive-Green rather than
+Olive as in <i>P. c. cinereus</i>. Coloration of adult female: Like female
+of <i>P. c. cinereus</i> but colors darker throughout; dorsum more
+olivaceous.</p>
+
+<p><i>Measurements.</i>&mdash;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.</p>
+
+<p><i>Range.</i>&mdash;Western Guatemala, in subtropical and temperate zones.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Ptilogonys caudatus</b> Cabanis</div>
+<br>
+<div class="center">Costa Rican Ptilogonys</div>
+
+<div class="blockquot smaller"><p><i>Ptilogonys caudatus</i> Cabanis, Jour. für Orn., 1866:402, Nov. 1866.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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)
+<span class="pagenum"><a name="Page_481" id="Page_481">[Pg&nbsp;481]</a></span>
+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.</p>
+
+<p><i><a name="Measurements"></a><a href="#typos">Measurements</a></i>&mdash;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.</p>
+
+<p><i>Range.</i>&mdash;Highlands of Costa Rica and extreme western Panam&aacute;.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Genus <b>Phainopepla</b> Sclater</div>
+
+<div class="blockquot smaller"><p><i>Phainopepla</i> Sclater, Proc. Zoöl. Soc. London, 26:543, 1858. Type
+<i>Phainopepla nitens</i> (Swainson). </p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Phainopepla nitens nitens</b> (Swainson)</div>
+<br>
+<div class="center">Phainopepla</div>
+
+<div class="blockquot smaller"><p><i>Phainopepla nitens nitens</i> (Swainson), Anim. in Menag., 1838:285,
+Dec. 31, 1837. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Measurements.</i>&mdash;No specimens examined; larger than <i>P. n. lepida</i> (Van
+Tyne, 1925).</p>
+
+<p><i>Range.</i>&mdash;Central and southern Mexico, in states of Coahuila, San <a name="Luis_Potosi2"></a><a href="#typos">Lu&iacute;s Potosi</a>,
+Durango, Guanajuato, M&eacute;xico, Puebla, and Veracruz.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Phainopepla nitens lepida</b> Van Tyne</div>
+<br>
+<div class="center">Phainopepla</div>
+
+<div class="blockquot smaller"><p><i>Phainopepla nitens lepida</i> Van Tyne, Occ. Pap. Bost. Soc. Nat.
+Hist., 5:149, 1925. </p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration same as <i>P. n. nitens</i>; separated by smaller
+size.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 91.0, tail 90.3, culmen 11.5, tarsus 17.6.</p>
+
+<p><span class="pagenum"><a name="Page_482" id="Page_482">[Pg&nbsp;482]</a></span>
+<i>Range.</i>&mdash;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).</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Subfamily <b>Bombycillinae</b></div>
+
+<p><i>Diagnosis.</i>&mdash;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 <i>B.
+japonica</i>); sexes alike; humerus short with large external condyle; caudal
+muscles and pygostyle not well developed; bill flared widely at base; one genus,
+three species.</p>
+
+<p><i>Range of subfamily.</i>&mdash;Holarctic breeding area; wanders nomadically south
+in winter to Central America and West Indies, southern Europe and Asia.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Genus <b>Bombycilla</b> Brisson</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla</i> Brisson, Orn. ii, 1760:337. Type <i>Bombycilla garrula</i> (Linnaeus).</p></div>
+
+<p><i>Diagnosis.</i>&mdash;As described for the subfamily.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla cedrorum</b> Vieillot</div>
+<br>
+<div class="center">Cedar Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla cedrorum</i> Vieillot, Hist. Nat. Amer., 1:88, Sept. 1, 1807</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 92.9, tail 55.5, culmen 10.9, tarsus 16.8.</p>
+
+<p><i>Range.</i>&mdash;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 Panam&aacute;.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla garrula</b> (Linnaeus)</div>
+<br>
+<div class="center">Bohemian Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla garrula</i> (Linnaeus), Syst. Nat., 10th Ed., 1758:55.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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
+<span class="pagenum"><a name="Page_483" id="Page_483">[Pg&nbsp;483]</a></span>
+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.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla garrula garrula</b> (Linnaeus)</div>
+<br>
+<div class="center">Bohemian Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla garrula garrula</i> (Linnaeus), Syst. Nat., 10th Ed., 1758:55.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration: As described for the species, but darkest of the
+three subspecies; tending to be more Vinaceous dorsally than either <i>pallidiceps</i>
+or <i>centralasiae</i>.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 113.5, tail 63.1, culmen 12.5, tarsus 20.7.</p>
+
+<p><i>Range.</i>&mdash;Europe; breeds north to northern Russia and Norway, south to
+about 65&deg; N latitude; winters south to England and Ireland, southern France,
+northern Italy, and Turkey.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla garrula centralasiae</b> Poljakov</div>
+<br>
+<div class="center">Bohemian Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla garrula centralasiae</i> Poljakov, Mess. Orn. vi:137, 1915.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration: As described for the subspecies <i>garrula</i>, but less
+Vinaceous dorsally, and more Cinnamon; venter lighter gray than <i>garrula</i>, and
+much paler than <i>pallidiceps</i>.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 114.7, tail 63.0, culmen 12.2, tarsus 21.0.</p>
+
+<p><i>Range.</i>&mdash;Asia; breeds northern Siberia south to Vladivostok; winters to
+Turkestan and central eastern China and Japan.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla garrula pallidiceps</b> Reichenow</div>
+<br>
+<div class="center">Bohemian Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla garrula pallidiceps</i> Reichenow, Orn. Monats. 16:191, 1908.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration: As described for the species, but more grayish
+above and below than <i>B. g. garrula</i>; darker gray than in <i>centralasiae</i>.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 115.1, tail 71.7, culmen 12.6, tarsus 21.1.</p>
+
+<p><i>Range.</i>&mdash;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.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Bombycilla japonica</b> (Siebold)</div>
+<br>
+<div class="center">Japanese Waxwing</div>
+
+<div class="blockquot smaller"><p><i>Bombycilla japonica</i> (Siebold), Nat. Hist. Jap., St. No. 2:87, 1824.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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;
+<span class="pagenum"><a name="Page_484" id="Page_484">[Pg&nbsp;484]</a></span>
+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.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 108.3, tail 53.6, culmen 11.2, tarsus 19.4.</p>
+
+<p><i>Range.</i>&mdash;Breeds eastern Siberia, northern China; winters south in China,
+and to Japan (Hokkaido, Kyushu), Taiwan, and Korea.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Subfamily <i>Dulinae</i></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Range of subfamily.</i>&mdash;Islands of Haiti and Gonave, Greater Antilles.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb">Genus <i>Dulus</i> Vieillot</div>
+
+<div class="blockquot smaller"><p><i>Dulus</i> Vieillot, Analyse, 1816:42.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;Like the subfamily.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Dulus dominicus dominicus</b> (Linnaeus)</div>
+<br>
+<div class="center">Palm-chat</div>
+
+<div class="blockquot smaller"><p><i>Dulus dominicus dominicus</i> (Linnaeus), Syst. Nat., 12th Ed., 1766:316.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;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.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 85.0, tail 68.8, culmen 15.0, tarsus 24.7.</p>
+
+<p><i>Range.</i>&mdash;Island of Haiti, Greater Antilles.</p>
+<p>&nbsp;</p>
+
+<div class="caption3nb"><b>Dulus dominicus oviedo</b> Wetmore</div>
+<br>
+<div class="center">Palm-chat</div>
+
+<div class="blockquot smaller"><p><i>Dulus dominicus oviedo</i> Wetmore, Proc. Biol. Soc. Wash., 42:117, 1929.</p></div>
+
+<p><i>Diagnosis.</i>&mdash;Coloration: Like <i>D. d. dominicus</i>, but averaging more Grayish
+Olive; rump and tail coverts with less greenish wash.</p>
+
+<p><i>Measurements.</i>&mdash;Wing 90.1, tail 71.3, culmen 16.2, tarsus 25.1.</p>
+
+<p><i>Range.</i>&mdash;Gonave Island, off Haiti, Greater Antilles.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="COLORATION" id="COLORATION"></a>
+<p><span class="pagenum"><a name="Page_485" id="Page_485">[Pg&nbsp;485]</a></span></p>
+<div class="caption2">COLORATION</div>
+
+<p>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 <i>B. garrula</i> and the
+yellow color on the distal ends of the rectrices (red in <i>B.
+japonica</i>) 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.</p>
+
+<p>The red wax tips on the secondaries of the flight feathers, and
+sometimes found on the ends of the rectrices in <i>Bombycilla</i>, 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 <i>B. cedrorum</i> (Farley, 1924). It is well known that many
+individuals, especially of <i>B. cedrorum</i>, do not possess these tips;
+they are absent in a smaller proportion of individuals of <i>B.
+garrula</i>. Of the 53 skins of <i>B. cedrorum</i> 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 <i>B.
+garrula pallidiceps</i> 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 <i>B.
+garrula</i>, 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 <i>B. garrula garrula</i> killed by storms in England in the winter of
+1866-67, 69 individuals had
+<span class="pagenum"><a name="Page_486" id="Page_486">[Pg&nbsp;486]</a></span>
+wax tips. Of these, 41 were males and 27 were females; the remaining
+one was of uncertain sex. Among 38 definitely sexed <i>B. garrula
+pallidiceps</i> 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 <i>B.
+garrula</i> than in <i>B. cedrorum</i>. The potentiality for wax tips is
+possibly inherited according to Mendelian ratio.</p>
+
+<p><i>Bombycilla japonica</i> 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
+<i>B. garrula</i> (and in <a name="cedrorum"></a><i><a href="#typos">B. cedrorum</a></i>): "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." <i>Bombycilla japonica</i>
+thus may be close to the ancestral condition in the waxwing stock in
+the development of the waxy appendage.</p>
+
+<p>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 <i>B. garrula</i>,
+and only occasionally in <i>B. cedrorum</i>. The pigment in the tip
+of the tail of <i>B. japonica</i> 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.</p>
+
+<p>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.</p>
+
+<p>The wax tips are not limited to the adult birds in the species <i>B.
+garrula</i>. Swarth (<i>op. cit.</i>) mentions the capture of several young
+<span class="pagenum"><a name="Page_487" id="Page_487">[Pg&nbsp;487]</a></span>
+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 <i>pallidiceps</i>, but at
+least for the European subspecies <i>garrula</i> as well. On the other
+hand, the young of <i>B. cedrorum</i> lack the wax tips, at least as far as
+available data show.</p>
+
+<p>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.</p>
+
+<p>The wax tips are ruptive in effect, since the birds, especially in
+winter, are habitu&eacute;s 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.</p>
+
+<p>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.</p>
+
+<p>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
+<span class="pagenum"><a name="Page_488" id="Page_488">[Pg&nbsp;488]</a></span>
+months, the wax tips that appear like berries, are especially valuable
+to the birds, and operate selectively.</p>
+
+<p>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. <i>B. japonica</i> is
+closer to the ancestral stock insofar as wax tips are concerned, and
+generally lacks the tips. <i>B. cedrorum</i> has the tips in approximately
+half of the adults, and not at all in the young. <i>B. garrula</i> 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.</p>
+
+<p>The streaked plumage of <i>Dulus</i> 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 <i>Dulus</i> 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, <i>Dulus</i> has evidently gained sufficient
+protection against predators; other birds solve this problem by
+modifying their coloration.</p>
+
+<p><i>Ptilogonys</i> is ruptively colored, but in a different fashion than
+<i>Bombycilla</i>. 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 <i>Ptilogonys</i>, 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.</p>
+
+<p>It is difficult to suggest reasons for the brilliant coloration of the
+male <i>Phainopepla</i>, unless it is for decoying predators away from
+the nest. Possibly some birds survive not because of, but in spite
+of, their coloration, and <i>Phainopepla</i> may be a case of this sort.
+Anyone who has observed <i>Phainopepla</i> in life will agree, certainly,
+that the male makes no attempt at concealment, and flaunts his
+color to all comers.</p>
+
+<p><span class="pagenum"><a name="Page_489" id="Page_489">[Pg&nbsp;489]</a></span>
+The coloration of <i>Phainoptila</i>, in contrast to <i>Phainopepla</i>, 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.</p>
+
+<p>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, <i>Dulus</i> has
+developed large feet, heavy decurved beak, and the large communal nest
+that affords protection from enemies; as a consequence, perhaps
+<i>Dulus</i> did not need a plumage different from the primitive and
+streaked one. The survival of <i>Dulus</i> 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.</p>
+
+<p>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&mdash;the plumage of the male is
+outstandingly conspicuous&mdash;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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="COURTSHIP" id="COURTSHIP"></a>
+<div class="caption2">COURTSHIP</div>
+
+<p>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
+<i>Dulus</i>, in which the territory is not limited even to
+<span class="pagenum"><a name="Page_490" id="Page_490">[Pg&nbsp;490]</a></span>
+the nest, but to the individual compartment of the community nest.
+<i>Phainopepla</i>, a less gregarious bird than <i>Dulus</i> and waxwings, has a
+much more definite territory, although individuals of <i>Phainopepla</i>
+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.</p>
+
+<p>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 (<i>op.
+cit.</i>:2) considered as possibly concerned with courtship.
+<i>Phainopepla</i> has various call notes, and in addition, a succession of
+notes which are run together. <i>Ptilogonys</i> 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." <i>Dulus</i> is described by Wetmore and Swales
+(1931:349) as having only a variety of rather harsh chattering notes
+in chorus.</p>
+
+<p>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
+<span class="pagenum"><a name="Page_491" id="Page_491">[Pg&nbsp;491]</a></span>
+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.</p>
+
+<p>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 <i>Phainopepla</i> has
+a nuptial flight in which the male chases the female, and that <i>Dulus</i>
+(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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="NEST_BUILDING" id="NEST_BUILDING"></a>
+<div class="caption2">NEST BUILDING</div>
+
+
+<p>There are numerous papers concerning the nesting of waxwings. <i>B.
+garrula</i>, owing to its nesting in the far north, where observers are
+few, has received less attention than <i>B. cedrorum</i>. There is, on the
+other hand, no literature that deals with the nesting habits of the
+majority of the Ptilogonatines, with the exception of <i>Phainopepla</i>,
+on which there is considerable literature (Merriam, 1896; Myers, 1907,
+1908). No detailed study of the nesting of <i>Dulus</i> has been reported,
+although Wetmore and Swales (1931) have described carefully the large
+communal nest of this genus.</p>
+
+<p>In <i>Bombycilla</i>, 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.</p>
+
+<p>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,
+<i>B. garrula</i> according to Swarth (<i>op. cit.</i>:271), and various plant
+fibers and string are used by <i>B. cedrorum</i>. 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
+<span class="pagenum"><a name="Page_492" id="Page_492">[Pg&nbsp;492]</a></span>
+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.</p>
+
+<p>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 (<i>op.
+cit.</i>:1) remarks that <i>B. cedrorum</i> 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.</p>
+
+<p>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.</p>
+
+<p>The very large and bulky communal nest of <i>Dulus</i> is not radically
+different from the nest of waxwings. In the absence of sufficient
+nesting sites, a pair of gregarious birds such as <i>Dulus</i> 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 <i>Dulus</i> 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.</p>
+
+<p>In <i>Phainopepla</i> 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 <i>Bombycilla</i>, but the materials used for a base
+are stems of annual plants, whereas <i>Bombycilla</i> uses more woody
+twigs. The finer materials used by <i>Phainopepla</i> are more readily
+obtainable in the ecological association inhabited by <i>Phainopepla</i>
+than would be heavier twigs such as <i>Bombycilla</i> uses.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="FOOD" id="FOOD"></a>
+<p><span class="pagenum"><a name="Page_493" id="Page_493">[Pg&nbsp;493]</a></span></p>
+<div class="caption2">FOOD</div>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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 <i>Ptilogonys cinereus</i> 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 <i>Eurya theoides</i> and <i>Monnina xalapensis</i>. It is well known that
+<i>Phainopepla</i> 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 (<i>Schinus molle</i>)
+and feeds on the small red berries. The preserved specimens of
+<i>Ptilogonys</i> and <i>Phainoptila</i> available for this study contain only
+berries in the digestive tract. <i>Dulus</i> feeds mostly, if not wholly, on
+plant food. According to Wetmore and Swales (1931:349), berries,
+fruits, and parts of flowers are eaten.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="SKELETON" id="SKELETON"></a>
+<p><span class="pagenum"><a name="Page_494" id="Page_494">[Pg&nbsp;494]</a></span></p>
+<div class="caption2">SKELETON</div>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p><a name="SKULL"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Skull.</i>&mdash;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.</p>
+
+<p>The most specialized bill is to be found in <i>Dulus</i>; 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 <i>Bombycilla</i>.
+Moreover, the "vomer" in <i>Dulus</i> and in <i>Phainoptila</i> is larger and
+heavier than in <i>Bombycilla</i>, and the quadrate and pterygoid bones
+are relatively large for support of the beak. The palatines, however,
+<span class="pagenum"><a name="Page_495" id="Page_495">[Pg&nbsp;495]</a></span>
+are weak in <i>Phainoptila</i>. In the Ptilogonatinae, with the exception
+of <i>Phainoptila</i>, the wings of the palatines flare more than in <i>Bombycilla</i>,
+but not to the extent that they do in <i>Dulus</i>, nor does the palatine
+bone present a concave appearance in the Ptilogonatinae. The
+premaxilla is a relatively weak bone in <i>Bombycilla</i> and <i>Phainopepla</i>,
+stronger in <i>Ptilogonys</i>, and is notably heavy in <i>Phainoptila</i> and
+<i>Dulus</i>, 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, <i>Phainopepla</i> and
+<i>Phainoptila</i> show the least flaring in this region.</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<table summary="bird skulls" class="center">
+    <tr>
+        <td colspan=2><a href="images/fig_1_lg.png"><img src="images/fig_1_sm.png" border=0 width="230" height="120" title="Fig.1" alt="Fig.1"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_2_lg.png"><img src="images/fig_2_sm.png" border=0 width="202" height="105" title="Fig.2" alt="Fig.2"></a></td>
+        <td><a href="images/fig_3_lg.png"><img src="images/fig_3_sm.png" border=0 width="196" height="115" title="Fig.3" alt="Fig.3"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_4_lg.png"><img src="images/fig_4_sm.png" border=0 width="190" height="102" title="Fig.4" alt="Fig.4"></a></td>
+        <td><a href="images/fig_5_lg.png"><img src="images/fig_5_sm.png" border=0 width="211" height="124" title="Fig.5" alt="Fig.5"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_6_lg.png"><img src="images/fig_6_sm.png" border=0 width="199" height="105" title="Fig.6" alt="Fig.6"></a></td>
+        <td><a href="images/fig_7_lg.png"><img src="images/fig_7_sm.png" border=0 width="221" height="110" title="Fig.7" alt="Fig.7"></a></td>
+    </tr>
+</table>
+<p>&nbsp;</p>
+
+<span class="smcap">Figs.</span> 1-7. Skulls in lateral view of five genera of Bombycillidae. Natural size.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">&nbsp;1.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">&nbsp;2.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">&nbsp;3.</td><td class="text_lf"><i>Phainopepla nitens</i>, male, MNH no. 24752, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">&nbsp;4.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no. 297,<br>Xilitla Region, San Lu&iacute;s Potosi, Mexico.</td></tr>
+<tr><td class="vtop">&nbsp;5.</td><td class="text_lf"><i>Dulus dominicus</i>, female, USNM no. 292652, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">&nbsp;6.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">&nbsp;7.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_496" id="Page_496">[Pg&nbsp;496]</a></span></p>
+
+<div class="center">
+<table width="75%" summary="bird skulls" class="center">
+    <tr>
+        <td colspan=3><a href="images/fig_8_lg.png"><img src="images/fig_8_sm.png" border=0 width="97" height="186" title="Fig.8" alt="Fig.8"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_9_lg.png"><img src="images/fig_9_sm.png" border=0 width="88" height="166" title="Fig.9" alt="Fig.9"></a></td>
+        <td><a href="images/fig_10_lg.png"><img src="images/fig_10_sm.png" border=0 width="90" height="165" title="Fig.10" alt="Fig.10"></a></td>
+        <td><a href="images/fig_11_lg.png"><img src="images/fig_11_sm.png" border=0 width="79" height="157" title="Fig.11" alt="Fig.11"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_12_lg.png"><img src="images/fig_12_sm.png" border=0 width="96" height="179" title="Fig.12" alt="Fig.12"></a></td>
+        <td><a href="images/fig_13_lg.png"><img src="images/fig_13_sm.png" border=0 width="96" height="164" title="Fig.13" alt="Fig.13"></a></td>
+        <td><a href="images/fig_14_lg.png"><img src="images/fig_14_sm.png" border=0 width="106" height="173" title="Fig.14" alt="Fig.14"></a></td>
+    </tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<span class="smcap">Figs.</span> 8-14. Skulls in ventral view of five genera of Bombycillidae. Natural size.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">&nbsp;8.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26492, 15 mi. SE Cartago,<br>
+Costa Rica.</td></tr>
+<tr><td class="vtop">&nbsp;9.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">10.</td><td class="text_lf"><i>Phainopepla nitens</i>, male, MNH no. 24754, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">11.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no 297, Xilitla<br>
+Region, San <a name="Luis"></a><a href="#typos">Lu&iacute;s</a> Potosi, Mexico.</td></tr>
+<tr><td class="vtop">12.</td><td class="text_lf"><i>Dulus dominicus</i>, female, USNM no. 292652, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">13.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">14.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_497" id="Page_497">[Pg&nbsp;497]</a></span></p>
+
+<div class="center">
+<table width="75%" summary="bird skulls dorsal view" class="center">
+    <tr>
+        <td colspan=3><a href="images/fig_15_lg.png"><img src="images/fig_15_sm.png" border=0 width="90" height="193" title="Fig.15" alt="Fig.15"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_16_lg.png"><img src="images/fig_16_sm.png" border=0 width="93" height="170" title="Fig.16" alt="Fig.16"></a></td>
+        <td><a href="images/fig_17_lg.png"><img src="images/fig_17_sm.png" border=0 width="85" height="170" title="Fig.10" alt="Fig.17"></a></td>
+        <td><a href="images/fig_18_lg.png"><img src="images/fig_18_sm.png" border=0 width="81" height="156" title="Fig.11" alt="Fig.18"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_19_lg.png"><img src="images/fig_19_sm.png" border=0 width="91" height="170" title="Fig.12" alt="Fig.19"></a></td>
+        <td><a href="images/fig_20_lg.png"><img src="images/fig_20_sm.png" border=0 width="99" height="161" title="Fig.13" alt="Fig.20"></a></td>
+        <td><a href="images/fig_21_lg.png"><img src="images/fig_21_sm.png" border=0 width="93" height="174" title="Fig.14" alt="Fig.21"></a></td>
+    </tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<span class="smcap">Figs.</span> 15-21. Skulls in dorsal view of five genera of Bombycillidae. Natural size.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">15.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">16.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">17.</td><td class="text_lf"><i>Phainopepla nitens</i>, male, MNH no. 24752, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">18.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no. 297, Xilitla Region, San Lu&iacute;s Potosi, Mexico.</td></tr>
+<tr><td class="vtop">19.</td><td class="text_lf"><i>Dulus dominions</i>, female, USNM no. 292642, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">20.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">21.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_498" id="Page_498">[Pg&nbsp;498]</a></span>
+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 <i>Phainopepla</i> there is much variability in this character;
+some specimens have a narrower antorbital bridge than others. Only
+one skeleton of <i>Phainopepla n. nitens</i> was available. The flaring in
+the skull of this specimen is identical with that in <i>Ptilogonys</i>.
+Among the skulls of <i>P. n. lepida</i> 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 <i>P. n. nitens</i>, 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 <i>lepida</i> 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 <i>Phainoptila</i>; the skull of No. 24754 is
+long and thin, and the base of the bill is only slightly swollen. The
+skull of <i>Phainopepla nitens lepida</i> is more generalized than that of
+<i>Phainopepla n. nitens</i>, having a longer and narrower bill like the
+generalized <i>Phainoptila</i>. In <i>Phainopepla n. nitens</i> and in members
+of the genus <i>Ptilogonys</i>, more flaring occurs in the antorbital region.</p>
+
+<p><i>Phainoptila</i>, as noted above, has no great amount of flaring in
+the antorbital region. When more specimens of <i>Phainoptila</i> 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 <i>Phainopepla</i>. The premaxilla and maxilla of
+<i>Phainoptila</i> are similar to the same bones in <i>Dulus</i>, and there is a
+well-marked ridge on the tomium (possibly for cutting flower
+parts). In <i>Phainoptila</i>, 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
+<i>Dulus</i>; the skull of <i>Phainoptila</i> seems to be modified for a frugivorous
+rather than an insectivorous diet. The skull of <i>Phainoptila</i>
+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 <i>Phainoptila</i> has progressed
+less in this direction than have other species in the family.</p>
+
+<p><span class="pagenum"><a name="Page_499" id="Page_499">[Pg&nbsp;499]</a></span>
+The mandibles vary somewhat in the shape and proportionate
+size of the bones. The mandible is proportionately, as well as actually,
+highest in <i>Dulus</i>. The medial condyle varies to some extent,
+being slightly flattened mediad in <i>Bombycilla</i>, and less so in the
+other genera. The mandible of <i>Bombycilla</i> narrows to the symphysis
+much more gradually than it does in the other genera.</p>
+
+<p>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.</p>
+
+<p>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 <i>Phainoptila</i>. In order of decreasing relative
+length of the cranium, <i>Phainoptila</i> is followed by <i>Ptilogonys caudatus</i>,
+<i>P. cinereus</i>, and <i>Phainopepla</i>. <i>Bombycilla garrula</i> has the
+deepest cranium in the family.</p>
+
+<p>The measurements of the lengths and widths of the skulls are given
+in <a href="#Table_9">Table 9</a>. The relative length of the bill and relative width of the
+skull are given in <a href="#Table_10">Table 10</a>. 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 <i>Phainoptila</i> has a slightly narrower cranium.</p>
+
+<p><a name="HUMERUS"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Humerus.</i>&mdash;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 <i>Dulus</i>. The deltoid tuberosity is variable, being
+shorter but more elevated in <i>Bombycilla</i> 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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_500" id="Page_500">[Pg&nbsp;500]</a></span></p>
+
+<div class="center">
+<table width="75%" summary="bird humeri" class="center">
+    <tr>
+        <td colspan=3><a href="images/fig_22_lg.png"><img src="images/fig_22_sm.png" border=0 width="64" height="117" title="Fig.22" alt="Fig.22"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_23_lg.png"><img src="images/fig_23_sm.png" border=0 width="66" height="118" title="Fig.16" alt="Fig.23"></a></td>
+        <td><a href="images/fig_24_lg.png"><img src="images/fig_24_sm.png" border=0 width="63" height="113" title="Fig.10" alt="Fig.24"></a></td>
+        <td><a href="images/fig_25_lg.png"><img src="images/fig_25_sm.png" border=0 width="67" height="111" title="Fig.11" alt="Fig.25"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_26_lg.png"><img src="images/fig_26_sm.png" border=0 width="68" height="106" title="Fig.12" alt="Fig.26"></a></td>
+        <td><a href="images/fig_27_lg.png"><img src="images/fig_27_sm.png" border=0 width="56" height="96" title="Fig.13" alt="Fig.27"></a></td>
+        <td><a href="images/fig_28_lg.png"><img src="images/fig_28_sm.png" border=0 width="61" height="108" title="Fig.14" alt="Fig.28"></a></td>
+    </tr>
+</table>
+
+<span class="smcap">Figs.</span> 22-28. Humeri of five genera of Bombycillidae. Natural size.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">22.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">23.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">24.</td><td class="text_lf"><i>Phainopepla nitens</i>, male, MNH no. 24754, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">25.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no. 297, Xilitla Region, San Lu&iacute;s Potosi, Mexico.</td></tr>
+<tr><td class="vtop">26.</td><td class="text_lf"><i>Dulus dominicus</i>, female, USNM no. 292652, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">27.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">28.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>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 <i>Tachycineta</i>. In the Bombycillidae the condyle is
+smallest in <i>Phainoptila</i>, 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 <i>Ptilogonys caudatus</i>, and a
+slightly pointed process in <i>P. cinereus</i>. The external condyle in
+<i>Dulus</i> is not specialized, being low and rounded, but in <i>Bombycilla</i>,
+it is noticeably elongated, indicating a better attachment distally for
+<span class="pagenum"><a name="Page_501" id="Page_501">[Pg&nbsp;501]</a></span>
+the deltoid muscle. (No measurements are tabulated for this
+condyle, as the percentage of error in measuring this small structure
+is great.) <a href="#Table_1">Table 1</a> gives lengths of humeri, and <a href="#Table_2">Table 2</a> gives
+lengths of the humeri expressed as percentages of the length of the
+trunk, a standard measurement.</p>
+
+<p>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.</p>
+
+<a name="Table_1"></a>
+<p class="center"><span class="smcap"><b>Table 1.</b></span> Lengths of Arm Bones in cm.</p>
+
+<table class="center" width="100%" summary="Lengths of Arm Bones in cm">
+<tr><td>Species</td><td>Humerus</td><td>Radius</td><td>Ulna</td><td>Manus</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>2.39</td><td>2.57</td><td>2.79</td><td>2.25</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>2.24</td><td>2.48</td><td>2.78</td><td>2.38</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>2.21</td><td>2.59</td><td>2.82</td><td>2.39</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>2.40</td><td>2.51</td><td>2.70</td><td>2.25</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>2.23</td><td>2.38</td><td>2.63</td><td>2.31</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>2.35</td><td>2.58</td><td>2.88</td><td>2.67</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>2.06</td><td>2.34</td><td>2.60</td><td>2.38</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_2"></a>
+<p class="center"><span class="smcap"><b>Table 2.</b></span> Arm-trunk Ratios (in percent)</p>
+
+<table class="center" width="100%" summary="Arm-trunk Ratios (in percent)">
+<tr><td>Species</td><td>Humerus</td><td>Radius</td><td>Ulna</td><td>Manus</td><td>Total</td></tr>
+<tr><td colspan=6><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>85</td><td>92</td><td>93</td><td>80</td><td>2.58</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>84</td><td>90</td><td>103</td><td>89</td><td>2.76</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>84</td><td>98</td><td>107</td><td>91</td><td>2.82</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>73</td><td>77</td><td>82</td><td>69</td><td>2.31</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>78</td><td>83</td><td>92</td><td>81</td><td>2.51</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>69</td><td>75</td><td>87</td><td>78</td><td>2.34</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>67</td><td>76</td><td>85</td><td>77</td><td>2.29</td></tr>
+<tr><td colspan=6><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_502" id="Page_502">[Pg&nbsp;502]</a></span></p>
+<a name="Table_3"></a>
+<p class="center"><span class="smcap"><b>Table 3.</b></span> Arm-trunk Ratios (in percent)</p>
+
+<table class="center" width="100%" summary="Arm-trunk Ratios (in percent)">
+<tr><td><span class="smcap">Species</span></td><td>Humerus</td><td>Radius</td><td>Ulna</td><td>Manus</td><td>Total</td></tr>
+<tr><td colspan=6><hr></td></tr>
+<tr><td class="text_lf">Corvus brachyrynchos</td><td>90</td><td>101</td><td>111</td><td>106</td><td>307</td></tr>
+<tr><td class="text_lf">Dendroica audubonii</td><td>68</td><td>82</td><td>90</td><td>77</td><td>237</td></tr>
+<tr><td class="text_lf">Setophaga ruticilla</td><td>69</td><td>82</td><td>91</td><td>75</td><td>235</td></tr>
+<tr><td class="text_lf">Myadestes townsendi</td><td>71</td><td>84</td><td>96</td><td>81</td><td>248</td></tr>
+<tr><td class="text_lf">Sialia sialis</td><td>72</td><td>84</td><td>98</td><td>86</td><td>256</td></tr>
+<tr><td class="text_lf">Hylocichla mustelina</td><td>75</td><td>81</td><td>92</td><td>80</td><td>247</td></tr>
+<tr><td class="text_lf">Parus atricapillus</td><td>85</td><td>90</td><td>106</td><td>81</td><td>272</td></tr>
+<tr><td class="text_lf">Tachycineta thalassina</td><td>71</td><td>95</td><td>107</td><td>128</td><td>306</td></tr>
+<tr><td class="text_lf">Myiarchus crinitus</td><td>83</td><td>105</td><td>115</td><td>92</td><td>290</td></tr>
+<tr><td class="text_lf">Dumetella carolinensis</td><td>76</td><td>75</td><td>89</td><td>78</td><td>243</td></tr>
+<tr><td class="text_lf">Polioptila caerulea</td><td>85</td><td>93</td><td>105</td><td>71</td><td>261</td></tr>
+<tr><td class="text_lf">Eremophila alpestris</td><td>91</td><td>99</td><td>110</td><td>95</td><td>296</td></tr>
+<tr><td class="text_lf">Muscivora forficata</td><td>85</td><td>111</td><td>120</td><td>108</td><td>313</td></tr>
+<tr><td colspan=6><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><a name="PYGOSTYLE"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Pygostyle.</i>&mdash;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.</p>
+
+<p>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 <i>Ptilogonys caudatus</i>, and narrowest in <i>Phainoptila</i>, in
+which genus, however, the entire bone is of small size. The centrum
+is widest in <i>Ptilogonys caudatus</i>, and is progressively narrower in
+<i>P. cinereus</i>, <i>Phainopepla</i>, and <i>Phainoptila</i>. 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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_503" id="Page_503">[Pg&nbsp;503]</a></span></p>
+<div class="center">
+<table width="75%" summary="bird pygostyles" class="center">
+    <tr>
+        <td colspan=2><a href="images/fig_29_lg.png"><img src="images/fig_29_sm.png" border=0 width="66" height="106" title="Fig.29" alt="Fig.29"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_30_lg.png"><img src="images/fig_30_sm.png" border=0 width="76" height="125" title="Fig.30" alt="Fig.30"></a></td>
+        <td><a href="images/fig_31_lg.png"><img src="images/fig_31_sm.png" border=0 width="69" height="114" title="Fig.31" alt="Fig.31"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_32_lg.png"><img src="images/fig_32_sm.png" border=0 width="75" height="106" title="Fig.32" alt="Fig.32"></a></td>
+        <td><a href="images/fig_33_lg.png"><img src="images/fig_33_sm.png" border=0 width="65" height="107" title="Fig.33" alt="Fig.33"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_34_lg.png"><img src="images/fig_34_sm.png" border=0 width="63" height="93" title="Fig.34" alt="Fig.34"></a></td>
+        <td><a href="images/fig_35_lg.png"><img src="images/fig_35_sm.png" border=0 width="75" height="96" title="Fig.35" alt="Fig.35"></a></td>
+    </tr>
+</table>
+<p>&nbsp;</p>
+
+<a name="Figs29-35"></a>
+<span class="smcap">Figs.</span> 29-35. Pygostyles in posterior view of five genera of Bombycillidae.
+&times; 2.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">29.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">30.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">31.</td><td class="text_lf"><i>Phainopepla nitens</i>, male, MNH no. 24754, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">32.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no. 297, Xilitla Region, San Lu&iacute;s Potosi, Mexico.</td></tr>
+<tr><td class="vtop">33.</td><td class="text_lf"><i>Dulus dominicus</i>, female, USNM no. 292652, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">34.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">35.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>In proportionate size (see <a href="#Table_7">Table 7</a>), the pygostyle of <i>Bombycilla</i>
+is the smallest in the family. The dorsal spinous portion is acutely
+pointed instead of flattened as in the Ptilogonatinae. In <i>Dulus</i>, 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 <i>Ptilogonys</i>.</p>
+
+<p>The centrum in cross section varies considerably. In <i>Bombycilla</i>
+the walls are indented, with definite terminal knobs; both knobs and
+indentations are more pronounced in <i>B. garrula</i> than in <i>cedrorum</i>,
+however. The spinous portion is enlarged in both species, and the
+rest of the neck region is constricted (<a href="#Figs29-35">Figs. 29-35</a>).</p>
+
+<p>The centrum of <i>Dulus</i> in posterior aspect presents the appearance
+of a simple shield; little of the indentation seen in <i>Bombycilla</i> is
+<span class="pagenum"><a name="Page_504" id="Page_504">[Pg&nbsp;504]</a></span>
+present. The spinous portion is plain, with no constriction nor
+terminal enlargement in the neck. The centrum in <i>Phainopepla</i> is
+similar to that in <i>Dulus</i>, 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 <i>Ptilogonys</i> shows great width, and the spine is in a
+large expanded tip as in <i>Bombycilla</i>. The lateral edges of the centrum
+in <i>P. cinereus</i> are "winged" and in two separate halves; whereas
+the centrum of <i>P. caudatus</i> is fairly plain, its specialization being
+reflected primarily in breadth and flatness. In cross section of the
+centrum, <i>Phainoptila</i> is similar to <i>Phainopepla</i>, 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 <i>Phainopepla</i>.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<table width="75%" summary="bird pygostyles" class="center">
+    <tr>
+        <td colspan=2><a href="images/fig_36_lg.png"><img src="images/fig_36_sm.png" border=0 width="70" height="102" title="Fig.36" alt="Fig.36"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_37_lg.png"><img src="images/fig_37_sm.png" border=0 width="74" height="118" title="Fig.37" alt="Fig.37"></a></td>
+        <td><a href="images/fig_38_lg.png"><img src="images/fig_38_sm.png" border=0 width="82" height="108" title="Fig.38" alt="Fig.38"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_39_lg.png"><img src="images/fig_39_sm.png" border=0 width="79" height="104" title="Fig.39" alt="Fig.39"></a></td>
+        <td><a href="images/fig_40_lg.png"><img src="images/fig_40_sm.png" border=0 width="72" height="106" title="Fig.40" alt="Fig.40"></a></td>
+    </tr>
+    <tr>
+        <td><a href="images/fig_41_lg.png"><img src="images/fig_41_sm.png" border=0 width="60" height="91" title="Fig.41" alt="Fig.41"></a></td>
+        <td><a href="images/fig_42_lg.png"><img src="images/fig_42_sm.png" border=0 width="75" height="93" title="Fig.42" alt="Fig.42"></a></td>
+    </tr>
+</table>
+<p>&nbsp;</p>
+
+<span class="smcap">Figs.</span> 36-42. Pygostyles in lateral view of five genera of Bombycillidae. &times; 2.<br>
+
+<table summary="bird catalog info">
+<tr><td class="vtop">36.</td><td class="text_lf"><i>Phainoptila m. melanoxantha</i>, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">37.</td><td class="text_lf"><i>Ptilogonys caudatus</i>, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.</td></tr>
+<tr><td class="vtop">38.</td><td class="text_lf"><i>Phainoptila nitens</i>, male, MNH no. 24754, Pima Co., Arizona.</td></tr>
+<tr><td class="vtop">39.</td><td class="text_lf"><i>Ptilogonys cinereus</i>, female, Louisiana State University no. 297, Xilitla Region, San Lu&iacute;s Potosi, Mexico.</td></tr>
+<tr><td class="vtop">40.</td><td class="text_lf"><i>Dulus dominicus</i>, female, USNM no. 292652, Don Don, Haiti.</td></tr>
+<tr><td class="vtop">41.</td><td class="text_lf"><i>Bombycilla cedrorum</i>, male, MNH no. 15331, Bexar Co., Texas.</td></tr>
+<tr><td class="vtop">42.</td><td class="text_lf"><i>Bombycilla garrula</i>, sex?, USNM no. 223895, Bozeman, Montana.</td></tr>
+</table>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>In specialization for muscle attachment, the centra of the pygostyles
+of the Ptilogonatinae have more area for muscle attachment
+<span class="pagenum"><a name="Page_505" id="Page_505">[Pg&nbsp;505]</a></span>
+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 <i>Phainoptila</i>, and that
+of <i>Dulus</i> differs only slightly. In <i>Bombycilla</i> 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.</p>
+
+<p><a name="STERNUM"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Sternum.</i>&mdash;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.</p>
+
+<p>In lateral aspect, the carina is deeper in <i>Bombycilla</i> than in other
+genera of the family, and is deepest in <i>B. garrula</i>. 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 <i>B. cedrorum</i>. In <i>Dulus</i>, 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.</p>
+
+<p>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 <i>Bombycilla</i> and <i>Dulus</i> the same
+process is closer to the body of the sternum. In <i>Bombycilla</i> the
+xiphoid process is more dorsal in position than in other species in the
+family, and in <i>Dulus</i> 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
+<i>Ptilogonys</i>.</p>
+
+<p><a name="RELATIVE_LENGTH"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Relative Lengths of Bones.</i>&mdash;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
+<span class="pagenum"><a name="Page_506" id="Page_506">[Pg&nbsp;506]</a></span>
+be such a part. For example, the length of the humerus divided by
+the length of the thoracic region yields, in <i>Phainopepla</i> and <i>Ptilogonys</i>,
+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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_4"></a>
+<p class="center"><span class="smcap"><b>Table 4.</b></span> Lengths of Leg Bones in cm.</p>
+
+<table class="center" width="100%" summary="Lengths of Leg Bones in cm">
+<tr><td><span class="smcap">Species</span></td><td>Femur</td><td>Tibiotarsus</td><td>Tarsometatarsus</td></tr>
+<tr><td colspan=4><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>2.04</td><td>3.10</td><td>1.94</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>1.89</td><td>2.90</td><td>1.77</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>1.76</td><td>2.78</td><td>1.72</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>2.43</td><td>3.77</td><td>2.58</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>2.09</td><td>3.34</td><td>2.09</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>2.32</td><td>3.46</td><td>1.99</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>1.92</td><td>2.95</td><td>1.64</td></tr>
+<tr><td colspan=4><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_5"></a>
+<p class="center"><span class="smcap"><b>Table 5.</b></span> Leg-trunk Ratios (in percent)</p>
+
+<table class="center" width="100%" summary="Leg-trunk Ratios (in percent)">
+<tr><td><span class="smcap">Species</span></td><td>Femur</td><td>Tibiotarsus</td><td>Tarsometatarsus</td><td>Total</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>73</td><td>110</td><td>69</td><td>252</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>71</td><td>109</td><td>66</td><td>246</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>69</td><td>106</td><td>65</td><td>240</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>74</td><td>115</td><td>60</td><td>249</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>73</td><td>119</td><td>73</td><td>265</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>68</td><td>101</td><td>59</td><td>228</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>63</td><td>96</td><td>53</td><td>212</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_507" id="Page_507">[Pg&nbsp;507]</a></span></p>
+
+<a name="Table_6"></a>
+<p class="center"><span class="smcap"><b>Table 6.</b></span> Leg-trunk Ratios (in percent)</p>
+
+<table class="center" width="100%" summary="Leg-trunk Ratios (in percent)">
+<tr><td><span class="smcap">Species</span>&nbsp;</td><td>Femur</td><td>Tibiotarsus</td><td>Tarsometatarsus</td><td>Total</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Corvus brachyrynchos</td><td>71</td><td>120</td><td>77</td><td>268</td></tr>
+<tr><td class="text_lf">Corvus corax</td><td>73</td><td>139</td><td>78</td><td>290</td></tr>
+<tr><td class="text_lf">Dendroica audubonii</td><td>62</td><td>109</td><td>81</td><td>252</td></tr>
+<tr><td class="text_lf">Setophaga ruticilla</td><td>66</td><td>127</td><td>94</td><td>287</td></tr>
+<tr><td class="text_lf">Myadestes townsendi</td><td>61</td><td>99</td><td>60</td><td>220</td></tr>
+<tr><td class="text_lf">Sialia sialis</td><td>66</td><td>111</td><td>72</td><td>249</td></tr>
+<tr><td class="text_lf">Hylocichla mustelina</td><td>75</td><td>133</td><td>97</td><td>305</td></tr>
+<tr><td class="text_lf">Parus atricapillus</td><td>78</td><td>138</td><td>99</td><td>315</td></tr>
+<tr><td class="text_lf">Tachycineta thalassina</td><td>61</td><td>97</td><td>56</td><td>214</td></tr>
+<tr><td class="text_lf">Myiarchus crinitus</td><td>68</td><td>106</td><td>74</td><td>248</td></tr>
+<tr><td class="text_lf">Dumetella carolinensis</td><td>73</td><td>136</td><td>94</td><td>303</td></tr>
+<tr><td class="text_lf">Polioptila caerulea</td><td>75</td><td>144</td><td>113</td><td>332</td></tr>
+<tr><td class="text_lf">Eremophila alpestris</td><td>73</td><td>113</td><td>115</td><td>301</td></tr>
+<tr><td class="text_lf">Muscivora forficata</td><td>62</td><td>98</td><td>61</td><td>221</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_7"></a>
+<p class="center"><span class="smcap"><b>Table 7.</b></span> Actual Length and Width in mm. of Pygostyle and Proportionate
+Length and Width of Pygostyle in percent of Lacrimal Length</p>
+
+<table class="center" width="100%" summary="Comparitive Length and Width of Pygostyle">
+<tr><td class="vbot"><span class="smcap">Species</span></td><td class="vbot">Length</td><td class="vbot">Width</td><td>Length,<br>percent</td><td>Width,<br>percent</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>9.8</td><td>3.9</td><td>45</td><td>18</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>8.8</td><td>4.1</td><td>41</td><td>19</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>8.4</td><td>3.9</td><td>41</td><td>19</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>8.5</td><td>3.5</td><td>35</td><td>14</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>8.5</td><td>2.9</td><td>38</td><td>13</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>7.0</td><td>3.5</td><td>31</td><td>15</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>7.1</td><td>2.9</td><td>35</td><td>14</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_508" id="Page_508">[Pg&nbsp;508]</a></span></p>
+
+<a name="Table_8"></a>
+<p class="center"><span class="smcap"><b>Table 8.</b></span> Length of Sternum and Depth of Carina expressed as percentages
+of the Length of the Trunk</p>
+
+<table class="center" width="100%" summary="Comparison of Sternum and Carina to Trunk Length">
+<tr><td><span class="smcap">Species</span></td><td>Sternum</td><td>Carina</td></tr>
+<tr><td colspan=3><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>85</td><td>28</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>91</td><td>32</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>81</td><td>26</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>76</td><td>25</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>107</td><td>28</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>88</td><td>33</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>82</td><td>31</td></tr>
+<tr><td colspan=3><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_9"></a>
+<p class="center"><span class="smcap"><b>Table 9.</b></span> Skull and Sternum, Length and Width in mm.</p>
+
+<table class="center" width="100%" summary="Skull and Sternum Length and Width">
+<tr><td>Species</td><td>Length of Skull</td><td>Width of Skull</td><td>Length of Sternum</td><td>Width of Sternum</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>34.9</td><td>15.6</td><td>23.9</td><td>7.8</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>33.4</td><td>14.7</td><td>24.3</td><td>8.5</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>33.3</td><td>15.1</td><td>21.3</td><td>6.9</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>39.7</td><td>16.0</td><td>24.8</td><td>8.2</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>36.4</td><td>16.6</td><td>30.5</td><td>8.0</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>37.0</td><td>16.8</td><td>30.0</td><td>11.2</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>34.0</td><td>15.5</td><td>25.3</td><td>9.6</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_509" id="Page_509">[Pg&nbsp;509]</a></span></p>
+
+<a name="Table_10"></a>
+<p class="center"><span class="smcap"><b>Table 10.</b></span> Relative Length and Width of Skull (in percent)</p>
+
+<table class="center" width="100%" summary="Relative Length and Width of Skull">
+<tr><td>Species</td><td>Length of Skull</td><td>Width of Skull</td></tr>
+<tr><td colspan=5><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>160</td><td>72</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>158</td><td>69</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>162</td><td>73</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>161</td><td>65</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>164</td><td>75</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>164</td><td>74</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>162</td><td>74</td></tr>
+<tr><td colspan=5><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<img src="images/fig_43.png" width="600" height="404" title="skeleton of Bombycilla cedrorum" alt="skeleton of Bombycilla cedrorum"><br>
+<span class="smcap">Fig.</span> 43. Part of skeleton of <i>Bombycilla cedrorum</i> showing method of
+measuring the length of the trunk. Natural size.<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><a name="LEG-TRUNK"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Leg-trunk Percentages.</i>&mdash;<a href="#Table_4">Table 4</a> shows the relative lengths of the
+legs and of the separate bones in the legs of the different species of
+the Bombycillids. <a href="#Table_5">Table 5</a> 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
+<span class="pagenum"><a name="Page_510" id="Page_510">[Pg&nbsp;510]</a></span>
+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.</p>
+
+<p><a href="#Table_4">Table 4</a> shows that of the birds compared in this paper, <i>Dulus</i>
+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 <a href="#Table_4">Table
+5</a> that <i>Tachycineta</i> and <i>Myadestes</i> 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 <i>Hylocichla mustelina</i>, a typical ground dweller, and <i>Parus atricapillus</i>,
+which hops about in the trees and underbrush.</p>
+
+<p>Insofar as the lengths of the legs show, <i>Dulus</i> and <i>Phainoptila</i> 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, <i>Dulus</i> and <i>Phainoptila</i> have remained unspecialized,
+in contrast to the waxwings in which adaptive changes fitting them
+for a perching habit have taken place. <i>Ptilogonys</i> and <i>Phainopepla</i>
+are intermediate in length of leg between <i>Phainoptila</i> and <i>Bombycilla</i>,
+and <i>Ptilogonys</i> and <i>Phainopepla</i> have progressed from life on
+the ground toward the perching habit. <i>Bombycilla cedrorum</i> is
+more specialized than is <i>B. garrula</i> in shortness of leg, and the reduction
+is comparable, as is noted above, to that in the legs of
+<i>Tachycineta</i>.</p>
+
+<p>In birds which have the legs much modified for walking or for hopping
+in the brush, such as <i>Polioptila</i> and <i>Eremophila</i>, it is noteworthy
+that the distal segment, the tarsometatarsus, is the longest, whereas
+in birds such as <i>Myiarchus</i> and <i>Tachycineta</i>, 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
+<span class="pagenum"><a name="Page_511" id="Page_511">[Pg&nbsp;511]</a></span>
+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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<img src="images/fig_44.png" width="600" height="514" title="leg bone lenghts" alt="leg bone lenghts"><br>
+<span class="smcap">Fig.</span> 44. Graph showing relative lengths of bones of the leg. The percentage
+values are shown on the axis of the ordinates.<br>
+<br>
+A. <i>Bombycilla cedrorum</i>; B. <i>Bombycilla garrula</i>; C. <i>Dulus dominicus</i>; D. <i>Phainoptila melanoxantha</i>; E. <i>Phainopepla nitens</i>; F. <i>Ptilogonys cinereus</i>; G. <i>Ptilogonys caudatus</i>.<br>a. femur; b. tibiotarsus; c. tarsometatarsus; d. total.<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><a name="ARM-TRUNK"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Arm-trunk Percentages.</i>&mdash;<a href="#Table_1">Tables 1 and 2</a> show the total length
+of the arm, and lengths of the separate arm elements, relative to the
+trunk. <a href="#Table_3">Table 3</a> 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 <a href="#Table_4">tables 4 and 5</a>. The method of adding together
+the component parts does not give the entire length of the
+<span class="pagenum"><a name="Page_512" id="Page_512">[Pg&nbsp;512]</a></span>
+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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<img src="images/fig_45_46.png" width="472" height="600" title="wing bones" alt="wing bones"><br>
+<span class="smcap">Figs.</span> 45-46. Outlines of wings. × 1/2<br>
+<br>
+<p>45. <i>Ptilogonys caudatus</i>, showing relation of outline of wing to bones of arm.</p>
+<p>46. <i>Bombycilla cedrorum</i>, showing relation of outline of wing to bones of arm.</p>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>It may be noted that <i>Phainoptila</i> and <i>Bombycilla</i> 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 <i>Phainoptila</i> that affects
+the total. The manus in <i>Phainoptila</i> is comparatively smaller than
+in any other genus of the family Bombycillidae, and this indicates
+poor flight power. <i>Bombycilla</i> has a total length corresponding
+<span class="pagenum"><a name="Page_513" id="Page_513">[Pg&nbsp;513]</a></span>
+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 <i>Dulus</i>; measurements for it
+are roughly the same as those of <i>Myadestes</i>. The wing bones of the
+Ptilogonatinae, other than <i>Phainoptila</i>, are the longest in this series,
+and they most nearly resemble the same bones in flycatchers, Parids,
+and gnatcatchers.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<img src="images/fig_47.png" width="600" height="445" title="arm bone lenghts" alt="arm bone lenghts"><br>
+<span class="smcap">Fig.</span> 47. Graph showing relative lengths of bones of the arm. The percentage values are shown on the axis of the ordinates.<br>
+<br>
+A. <i>Bombycilla cedrorum</i>; B. <i>Bombycilla garrula</i>; C. <i>Dulus dominicus</i>; D. <i>Phainoptila melanoxantha</i>; E. <i>Phainopepla nitens</i>; F. <i>Ptilogonys cinereus</i>; G.<i> Ptilogonys caudatus</i>.<br>
+a. humerus; b. radius; c. ulna; d. manus; e. total.<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span><br>
+<span class="pagenum"><a name="Page_514" id="Page_514">[Pg&nbsp;514]</a></span></p>
+<a name="MUSCULATURE"></a>
+<div class="caption2">MUSCULATURE</div>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p><a name="CAUDAL"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Caudal Muscles.</i>&mdash;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 <a href="#Table_11">Table
+11</a>, 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
+<span class="pagenum"><a name="Page_515" id="Page_515">[Pg&nbsp;515]</a></span>
+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 <i>Dulus</i> and <i>Bombycilla</i> constitute a smaller
+percentage of the total weight of the bird than in any of the genera
+in the subfamily Ptilogonatinae.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<a href="images/fig_48_lg.png"><img src="images/fig_48_sm.png" width="149" height="215" border=0 title="caudal musculature" alt="caudal musculature"></a><br>
+<span class="smcap">Fig.</span> 48. Caudal musculature, of <i>Phainopepla nitens lepida</i>, in dorsal view. × 2.<br>
+a. Levator coccygis; b. Levator caudae; c. Lateralis caudae;<br>
+d. Lateralis coccygis; e. oil gland; f. dorsal tip of pygostyle.<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<a name="Table_11"></a>
+<p class="center"><span class="smcap"><b>Table 11.</b></span> Caudal Muscles (Actual and Relative Weights)</p>
+
+<table width="100%" summary="Caudal Muscle Data">
+<tr><td><span class="smcap">Species</span></td><td>Levator</td><td>Lateralis</td></tr>
+<tr><td colspan=3><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>.145g.</td><td>.022g.</td></tr>
+<tr><td>&nbsp;</td><td>.092%</td><td>.045%</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>.030g.</td><td>.010g.</td></tr>
+<tr><td>&nbsp;</td><td>.076%</td><td>.026%</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>.025g.</td><td>.008g.</td></tr>
+<tr><td>&nbsp;</td><td>.096%</td><td>.029%</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>.040g.</td><td>.015g.</td></tr>
+<tr><td>&nbsp;</td><td>.063%</td><td>.014%</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>.028g.</td><td>.006g.</td></tr>
+<tr><td>&nbsp;</td><td>.063%</td><td>.014%</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>.034g.</td><td>.010g.</td></tr>
+<tr><td>&nbsp;</td><td>.048%</td><td>.014%</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>.026g.</td><td>.008g.</td></tr>
+<tr><td>&nbsp;</td><td>.050%</td><td>.014%</td></tr>
+<tr><td colspan=3><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_516" id="Page_516">[Pg&nbsp;516]</a></span></p>
+<a name="Table_12"></a>
+<p class="center"><span class="smcap"><b>Table 12.</b></span> Weights of Muscles (These percentages expressed in terms of
+weights of the body)</p>
+
+<table class="center" width="100%" summary="Weights of Muscle">
+<tr><td><span class="smcap">Species</span></td><td>P. major</td><td>P. minor</td><td>Deltoid</td><td>Thigh</td><td>Peroneus</td><td><a name="Gastrocnemius"></a><a href="#typos">Gastrocnemius</a></td></tr>
+<tr><td colspan=7><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>2.42g.</td><td>.29g.</td><td>.55g.</td><td>.43g.</td><td>.15g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>4.94%</td><td>.59%</td><td>1.12%</td><td>.88%</td><td>.31%</td><td>.96%</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>2.19g.</td><td>.28g.</td><td>.53g.</td><td>.30g.</td><td>.08g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>5.57%</td><td>.71%</td><td>1.35%</td><td>.71%</td><td>.21%</td><td>1.02%</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>1.30g.</td><td>.20g.</td><td>.30g.</td><td>.28g.</td><td>.10g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>4.99%</td><td>.77%</td><td>1.15%</td><td>1.12%</td><td>.40%</td><td>1.42%</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>3.93g.</td><td>.44g.</td><td>.92g.</td><td>1.09g.</td><td>.48g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>6.18%</td><td>.69%</td><td>1.45%</td><td>1.61%</td><td>.75%</td><td>2.97%</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>2.09g.</td><td>.22g.</td><td>.50g.</td><td>.73g.</td><td>.18g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>4.81%</td><td>.50%</td><td>1.15%</td><td>1.68%</td><td>.41%</td><td>1.01%</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>3.85g.</td><td>.45g.</td><td>.55g.</td><td>.50g.</td><td>.15g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>5.31%</td><td>.62%</td><td>.76%</td><td>.69%</td><td>.18%</td><td>.59%</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>2.58g.</td><td>.35g.</td><td>.50g.</td><td>.37g.</td><td>.10g.</td><td>&nbsp;</td></tr>
+<tr><td>&nbsp;</td><td>5.00%</td><td>.68%</td><td>.97%</td><td>.73%</td><td>.19%</td><td>.83%</td></tr>
+<tr><td colspan=7><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><a name="PECTORAL"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span><br>
+<span class="pagenum"><a name="Page_517" id="Page_517">[Pg&nbsp;517]</a></span></p>
+
+<p><i>Pectoral Muscles.</i>&mdash;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 <i>Bombycilla</i> than in members
+of the other two subfamilies. The humerus in <i>Bombycilla</i> 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.</p>
+
+<p><a name="HIND_LIMB"></a><span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<i>Leg Musculature.</i>&mdash;The muscles of the thigh are noticeably larger
+in birds that have long leg bones. (See <a href="#Table_12">Table 12</a> 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 <i>Phainoptila</i>, in which
+genus all the leg muscles are well developed, but the gastrocnemius
+is larger in all the Ptilogonatinae and in <i>Dulus</i> than it is in the
+specialized <i>Bombycilla</i>, in which it has probably been reduced as
+the leg bones and other muscles have been reduced.</p>
+
+<p>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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="DIGESTIVE_TRACT" id="DIGESTIVE_TRACT"></a>
+<span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<div class="caption2">DIGESTIVE TRACT</div>
+
+<p>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, <i>Phainoptila</i> and
+<i>Ptilogonys caudatus</i> having the folds of the digestive tract loosely
+arranged, whereas <i>Ptilogonys cinereus</i> and <i>Phainopepla</i> have folds
+which adhere more tightly to the ventriculus and liver. In <i>Dulus</i>
+and <i>Bombycilla</i>, 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.</p>
+
+<p>The coiling of the intestine, if its degree of compactness is disregarded,
+is nearly identical in the birds of the family; there are
+<span class="pagenum"><a name="Page_518" id="Page_518">[Pg&nbsp;518]</a></span>
+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 <a href="#Table_13">Table 13</a>, in which the actual and relative
+lengths of the intestine are given. It may be seen that in <i>Bombycilla</i>
+and in <i>Phainopepla</i>, 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 <i>Bombycilla</i> and <i>Phainopepla</i>, thereby permitting
+the necessary nutriment to be extracted by a short digestive
+tract.</p>
+
+<p>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.</p>
+
+<a name="Table_13"></a>
+<p class="center"><span class="smcap"><b>Table 13.</b></span> Digestive Tract: Actual Length, and Length Relative to
+Thoracic Length</p>
+
+<table class="center" width="100%" summary="Digestive Tract Length Data">
+<tr><td><span class="smcap">Species</span></td><td>Length<br>in mm.</td><td>Relative length<br>(in percent)</td></tr>
+<tr><td colspan=3><hr></td></tr>
+<tr><td class="text_lf">Ptilogonys caudatus</td><td>134</td><td>476.9</td></tr>
+<tr><td class="text_lf">Ptilogonys cinereus</td><td>111</td><td>415.6</td></tr>
+<tr><td class="text_lf">Phainopepla nitens</td><td>94</td><td>357.5</td></tr>
+<tr><td class="text_lf">Phainoptila melanoxantha</td><td>150</td><td>457.1</td></tr>
+<tr><td class="text_lf">Dulus dominicus</td><td>130</td><td>451.0</td></tr>
+<tr><td class="text_lf">Bombycilla garrula</td><td>102</td><td>298.2</td></tr>
+<tr><td class="text_lf">Bombycilla cedrorum</td><td>95</td><td>309.5</td></tr>
+<tr><td colspan=3><hr></td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>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
+<span class="pagenum"><a name="Page_519" id="Page_519">[Pg&nbsp;519]</a></span>
+correlation between large caecae and more insectivorous diet and
+small caecae and frugivorous diet; however, the data are not conclusive
+in this regard.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<a name="ORIGIN_OF_THE_SPECIES" id="ORIGIN_OF_THE_SPECIES"></a>
+<div class="caption2">ORIGIN OF THE SPECIES</div>
+
+<p>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 (<i>Bombycilla</i> alone
+is reported, in essentially its modern form, from the late Pleistocene&mdash;Wetmore,
+1940a), the place and time of origin cannot certainly be
+determined.</p>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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
+<span class="pagenum"><a name="Page_520" id="Page_520">[Pg&nbsp;520]</a></span>
+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.</p>
+
+<p>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.</p>
+
+<p>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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<div class="center">
+<img src="images/fig_49.png" width="600" height="316" title="family tree" alt="family tree"><br><br>
+<span class="smcap">Fig. 49.</span> Hypothetical family tree of the Bombycillidae.<br>
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p>The present day <i>Phainoptila</i> is most nearly like the ancestral
+group, and the remainder of the Ptilogonatinae have diverged to
+<span class="pagenum"><a name="Page_521" id="Page_521">[Pg&nbsp;521]</a></span>
+fit conditions similar to those to which the Tyrannid flycatchers,
+which parallel them, are also fitted.</p>
+
+<p>In comparatively recent geological time, two basic lines developed
+from the Bombycilline stock, the future <i>B. garrula</i> and <i>B. cedrorum</i>.
+Possibly <i>garrula</i> originally was isolated in Europe and Asia, and
+later came into contact with <i>B. cedrorum</i>, following the time at
+which the two species were genetically well differentiated. It
+appears certain that <i>B. japonica</i> 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.</p>
+
+<p>Structural affinities of <i>Dulus</i> and <i>Bombycilla</i> are more pronounced
+than are those of <i>Dulus</i> and <i>Ptilogonys</i>, for example. Many of the
+structural features of <i>Dulus</i> parallel those of <i>Phainoptila</i>, 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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span>
+<a name="CONCLUSIONS" id="CONCLUSIONS"></a>
+<div class="caption2">CONCLUSIONS</div>
+
+<p>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
+(<a href="#DIAGNOSES">page 478</a>) 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.</p>
+
+<p>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.</p>
+
+<p>Consideration of the geographic distribution of the species of the
+family indicates that the center of origin of the family Bombycillidae
+<span class="pagenum"><a name="Page_522" id="Page_522">[Pg&nbsp;522]</a></span>
+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.</p>
+
+<p>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
+<i>Phainoptila</i> 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.</p>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p><span class="pagenum"><a name="Page_523" id="Page_523">[Pg&nbsp;523]</a></span>
+The shape of the pygostyle varies in the Bombycillidae, but the
+simple shieldlike bone of <i>Phainoptila</i> is judged to resemble closely
+the ancestral type. In <i>Ptilogonys</i> there is a tall dorsal spine,
+coupled with a wide and heavy centrum and flattened lateral areas,
+for support of the long rectrices. In <i>Bombycilla</i> the bone is small
+with knobs on the centrum that have been developed for muscle
+attachment.</p>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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 <i>Bombycilla</i> and
+in <i>Phainopepla</i>. 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, <i>Bombycilla</i> 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.</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="SUMMARY"></a>
+<span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span><br>
+<p><span class="pagenum"><a name="Page_524" id="Page_524">[Pg&nbsp;524]</a></span></p>
+<div class="caption2">SUMMARY</div>
+
+<table class="text_lf" summary="Conclusions" width="100%">
+<tr><td class="vtop">&nbsp;1.</td><td>The silky flycatchers, waxwings, and palm-chats are included
+in the family Bombycillidae; the Ptilogonatidae and Dulidae
+are reduced to subfamily rank.</td></tr>
+
+<tr><td class="vtop">&nbsp;2.</td><td>The coloration of the birds of each subfamily is different because
+the ecological needs are different.</td></tr>
+
+<tr><td class="vtop">&nbsp;3.</td><td>Waxwings were at one time regularly migratory, but are now
+nomadic, since they are adapted to live in northern latitudes for
+the entire year.</td></tr>
+
+<tr><td class="vtop">&nbsp;4.</td><td>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.</td></tr>
+
+<tr><td class="vtop">&nbsp;5.</td><td>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.</td></tr>
+
+<tr><td class="vtop">&nbsp;6.</td><td>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.</td></tr>
+
+<tr><td class="vtop">&nbsp;7.</td><td>The pygostyle varies in shape and size between genera and
+even between some species.</td></tr>
+
+<tr><td class="vtop">&nbsp;8.</td><td>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.</td></tr>
+
+<tr><td class="vtop">&nbsp;9.</td><td>The muscles of the hind limb vary in mass, but not in kind,
+in the members of the family Bombycillidae.</td></tr>
+
+<tr><td class="vtop">10.</td><td>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.</td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+<a name="BIBLIOGRAPHY"></a>
+<span class="pagenum"><a href="#TOC">[&uarr;&nbsp;TOC]</a></span><br>
+<p><span class="pagenum"><a name="Page_525" id="Page_525">[Pg&nbsp;525]</a></span></p>
+<div class="caption2">BIBLIOGRAPHY</div>
+
+<span class="smcap">Anderson, E. M.</span><br>
+<div class="references"><p>1915. Nesting of the Bohemian Waxwing in northern British Columbia. Condor, 17(4):145-148, 1915.</p></div>
+
+<span class="smcap">Anderson, M. P.</span><br>
+<div class="references"><p>1907. A collecting trip in Korea. Condor, 9(5):146-147, 1907.</p></div>
+
+<span class="smcap">Anderson, R. M.</span><br>
+<div class="references"><p>1909. Nesting of the Bohemian Waxwing (<i>Bombycilla garrulus</i>). Auk, 26(1):10-12, 1909.</p></div>
+
+<span class="smcap">Armstrong, E. A.</span><br>
+<div class="references"><p>1942. Bird display. Cambridge Univ. Press, xvi + 381 pp., 22 plates, 1942.</p></div>
+
+<span class="smcap">Baird, S. F.</span><br>
+<div class="references"><p>1860. The birds of North America. J. B. Lippincott Co., lvi + 1003 pp., 1860.</p></div>
+
+<span class="smcap">Beddard, F. E.</span><br>
+<div class="references"><p>1898. The structure and classification of birds. Longmans, Green &amp; Co., xx + 548 pp., 252 figs., 1898.</p></div>
+
+<span class="smcap">Bergtold, W. H.</span><br>
+<div class="references"><p>1917a. A study of the incubation period of birds. Kendrick-Bellamy Co., 109 pp., 1917.</p></div>
+<div class="references"><p>1917b. Regurgitation in the Bohemian Waxwing. Auk, 34(3):341-342, 1917.</p></div>
+<div class="references"><p>1924. A summer occurrence of the Bohemian Waxwing in Colorado. Auk, 41(4):614, 1924.</p></div>
+
+<span class="smcap">Boulton, R.</span><br>
+<div class="references"><p>1926. Remarks on the origin and distribution of the Zonotrichiae. Auk,
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+
+<span class="smcap">Burleigh, T. D.</span><br>
+<div class="references"><p>1921. Breeding birds of Warland, Lincoln County, Montana. Auk, 38(4):552-565, 1921.</p></div>
+
+<span class="smcap">Burt, W. H.</span><br>
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+Zoöl., 32(8):455-524, 29 figs. in text, 1930.</p></div>
+
+<span class="smcap">Carriker, M. A., Jr.</span><br>
+<div class="references"><p>1909-1912. An annotated list of the birds of Costa Rica including Cocos
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+
+<span class="smcap">Cory, C. B.</span><br>
+<div class="references"><p>1886. The birds of the West Indies, etc. Auk, 3(2):187-245, 1886.</p></div>
+
+<span class="smcap">Crouch, J. E.</span><br>
+<div class="references"><p>1936. Nesting habits of the Cedar Waxwing. Auk, 53(1):1-8, 1936.</p></div>
+<div class="references"><p>1943. Distribution and habitat relationships of the Phainopepla. Auk,
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+
+<span class="smcap">Engels, W. L.</span><br>
+<div class="references"><p>1938. Cursorial adaptations in birds&mdash;limb proportions in the skeleton of
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+Zoöl., 42(7):341-400, 24 figs. in text, 1940.</p></div>
+
+<p><span class="pagenum"><a name="Page_526" id="Page_526">[Pg&nbsp;526]</a></span></p>
+<span class="smcap">Farley, J. A.</span><br>
+<div class="references"><p>1924. Abnormal Cedar Waxwing. Auk, 41(1):160, 1924.</p></div>
+
+<span class="smcap">Fisher, H. I.</span><br>
+<div class="references"><p>1946. Adaptations and comparative anatomy of the locomotor apparatus of New World Vultures. Amer. Midl. Nat., 35:545-727, 14 plates, 42 tables, 28 figs. in text, 1946.</p></div>
+
+<span class="smcap">Frank, F.</span><br>
+<div class="references"><p>1939. Die Färbung der Vogelfeder durch Pigment und Struktur. Jour. für Orn., 87:426-523, 1939.</p></div>
+
+<span class="smcap">Garrod, A. H.</span><br>
+<div class="references"><p>1876. On some anatomical peculiarities which bear upon the major divisions of the passerine birds, Pt. I. Proc. Zoöl. Soc. London, 626-647, 1876.</p></div>
+
+<span class="smcap">Gerondet, P.</span><br>
+<div class="references"><p>1948. Le jaseur boreal en Suisse pendant l'hiver 1946-1947. Der Orn. Beob., 45(1):1-5, 1948.</p></div>
+
+<span class="smcap">Gould, J.</span><br>
+<div class="references"><p>1862. The birds of Great Britain. London, published by the author, 5 vols., text unpaged, 367 plates, 1862.</p></div>
+
+<span class="smcap">Grinnel, J.</span><br>
+<div class="references"><p>1901. The status of the Cedar Waxwing in California. Condor, 3(6):146-147, 1901.</p></div>
+
+<div class="references"><p>1909. A new cowbird of the genus <i>Molothrus</i>. Univ. California Publ. Zoöl., 5:275-281, 6 figs. in text, 1909.</p></div>
+
+<span class="smcap">Griscom, L.</span><br>
+<div class="references"><p>1934. The ornithology of Guerrero, Mexico. Mus. Comp. Zoöl. Bull. 75:367-422, 1934.</p></div>
+
+<span class="smcap">Hamilton, W. J., Jr.</span><br>
+<div class="references"><p>1933. A late nesting waxwing in central New York. Auk, 50(2):114-115, 1933.</p></div>
+
+<span class="smcap">Hanna, W. C.</span><br>
+<div class="references"><p>1931. Nesting of the Bohemian Waxwing in British Columbia. Condor, 33(6):253-254, 1 fig., 1931.</p></div>
+
+<span class="smcap">Heinroth, O.</span><br>
+<div class="references"><p>1924. Die Vögel Mitteleuropas. Berlin, Huge Bermühler, 1:51-58, 1924.</p></div>
+
+<span class="smcap">Hellmayr, C. E.</span><br>
+<div class="references"><p>1935. Catalogue of the birds of the Americas. Field Mus. Nat. Hist. Mus. Publ. 347, 8(pt. 8):vi + 541 pp., 1935.</p></div>
+
+<span class="smcap">Howell, A. B.</span><br>
+<div class="references"><p>1938. Muscles of the avian hip and thigh. Auk, 55(1):71-81, 2 figs. in text, 1938.</p></div>
+<div class="references"><p>1944. Speed in animals, their specialization for running and leaping. Univ. Chicago Press, xi + 270 pp., 55 figs. 1944.</p></div>
+
+<span class="smcap">Hudson, G. E.</span><br>
+<div class="references"><p>1937. Studies on the muscles of the pelvic appendage in birds. Amer. Midl. Nat., 18:1-108, 26 plates, 1937.</p></div>
+<div class="references"><p>1948. Studies on the muscles of the pelvic appendages in birds II, the heterogeneous order Falconiformes. Amer. Midl. Nat., 39(1):102-127, 1948.</p></div>
+
+<span class="pagenum"><a name="Page_527" id="Page_527">[Pg&nbsp;527]</a></span>
+<span class="smcap">Knowlton, F. H.</span>
+<div class="references"><p>1909. Birds of the world. Henry Holt &amp; Co., Ltd., xi + 873 pp., 15 plates, 233 figs. in text, 1909.</p></div>
+
+<span class="smcap">Konoda, N.</span><br>
+<div class="references"><p>1943. A dictionary of animals. Tokyo, 3 + 767 + 50 pp., profusely illustrated, 1943.</p></div>
+
+<span class="smcap">Koshantschikov, I.</span><br>
+<div class="references"><p>1930. Ein Beitrag zur Kenntnis der Okologie, Biologie, und Geographie des Zobels (<i>Martes zibellina</i> L.). Zeits. für Okol. der Tierre, 19(2):291-320, 2 maps, 1930.</p></div>
+
+<span class="smcap">Linsdale, J. M.</span><br>
+<div class="references"><p>1928. Variations in the Fox Sparrow (<i>Passerella iliaca</i>) with reference to natural history and osteology. Univ. California Publ. Zoöl., 30(12):251-392, 4 plates, 38 figs. in text, 1928.</p></div>
+
+<span class="smcap">Littlefield, M. J.</span>, and <span class="smcap">Lemkan, F.</span><br>
+<div class="references"><p>1928. History of a Cedar Waxwing family. Bull. NE Bird-Band. Assoc., 4:85-89, 1928.</p></div>
+
+<span class="smcap">Lucas, F. A.</span><br>
+<div class="references"><p>1897. The tongues of birds. U. S. Nat. Mus. Report for 1895, 1001-1019 pp., 2 plates, 1897.</p></div>
+
+<span class="smcap">McGregor, R. C.</span><br>
+<div class="references"><p>1906. Notes on birds observed while traveling from Yokohama to Manila. Condor, 8(4):98-100, 1906.</p></div>
+
+<span class="smcap">Matthew, W. D.</span><br>
+<div class="references"><p>1939. Climate and evolution. N. Y. Acad. Sci., Spec. Publ., 1:xi + 223 pp., 1939.</p></div>
+
+<span class="smcap">Mayr, E.</span><br>
+<div class="references"><p>1942. Systematics and the origin of species. Columbia Univ. Press, xiv + 334 pp., 29 figs. in text, 1942.</p></div>
+<div class="references"><p>1947. Ecological factors in speciation. Evolution, 1(4):263-288, 1947.</p></div>
+
+<span class="smcap">Merriam, F. A.</span><br>
+<div class="references"><p>1896. Nesting habits of <i>Phainopepla nitens</i> in California. Auk, 8(1):38-43, 1896.</p></div>
+
+<span class="smcap">Miller, A. H.</span><br>
+<div class="references"><p>1933. Postjuvenal molt and the appearance of sexual characters of plumage in <i>Phainopepla nitens</i>. Univ. California Publ. Zoöl., 38(13):425-444 pp., 8 pls., 1 fig. in text, 1933.</p></div>
+<div class="references"><p>1937. Structural modifications in the Hawaiian Goose (<i>Nesochen sandvicensis</i>). A study in adaptive evolution. Univ. California Publ. Zoöl., 42(1):1-80, 6 plates, 12 figs. in text, 1937.</p></div>
+<div class="references"><p>1941. Speciation in the avian genus Junco. Univ. California Publ. Zoöl., 44(3):173-434, 33 figs. in text, 1941.</p></div>
+
+<span class="smcap">Muller, C. S.</span><br>
+<div class="references"><p>1915. A northern winter record of the Phainopepla. Condor, 17(3):129, 1915.</p></div>
+
+<span class="smcap">Myers, H. W.</span><br>
+<div class="references"><p>1907. Nesting habits of <i>Phainopepla nitens</i>. Condor, 9(4):101-103, 1907.</p></div>
+<div class="references"><p>1908. Observations on the nesting habits of <i>Phainopepla</i>. Condor, 10(2):72-75, 1908.</p></div>
+<p><span class="pagenum"><a name="Page_528" id="Page_528">[Pg&nbsp;528]</a></span></p>
+<div class="references"><p>1909. Notes on the habits of <i>Phainopepla nitens</i>. Condor, 11(1):22-23, 1909.</p></div>
+
+<span class="smcap">Newton, A.</span>, and <span class="smcap">Gadow, H.</span><br>
+<div class="references"><p>1893-1896. A dictionary of birds. Adams and Charles Black, xii + 1086 pp., 1893-1896.</p></div>
+
+<span class="smcap">Nice, M. M.</span><br>
+<div class="references"><p>1940. Observations on the behavior of a young Cedar Waxwing. Condor, 43(1):58-64, 1940.</p></div>
+
+<span class="smcap">Oberholser, H. C.</span><br>
+<div class="references"><p>1917. A synopsis of the races of <i>Bombycilla garrula</i> (Linnaeus). Auk, 34(3):330-333, 1917.</p></div>
+
+<span class="smcap">Pemberton, J. R.</span><br>
+<div class="references"><p>1908. Northern range of the <i>Phainopepla</i>. Condor, 10(6):238, 1908.</p></div>
+
+<span class="smcap">Plath, K.</span><br>
+<div class="references"><p>1933. Molt of the Nonpareil. Auk, 50(2):121, 1933.</p></div>
+
+<span class="smcap">Post, K. C.</span><br>
+<div class="references"><p>1916. The Cedar Waxwing (<i>Bombycilla cedrorum</i>) during July and August, 1916. Wilson Bull., 28:175-193, 1916.</p></div>
+
+<span class="smcap">Rand, A. L.</span>, and <span class="smcap">Rand, R. M.</span><br>
+<div class="references"><p>1943. Breeding notes on <i>Phainopepla</i>. Auk, 60(3):333-341, 1943.</p></div>
+
+<span class="smcap">Richardson, F.</span><br>
+<div class="references"><p>1942. Adaptive modifications for trunk foraging in birds. Univ. California Pub. Zoöl., 46(4):317-368, 2 plates, 16 figs. in text, 1942.</p></div>
+
+<span class="smcap">Ridgway, R.</span><br>
+<div class="references"><p>1904. The birds of North and Middle America, Part III. U. S. Nat. Mus. Bull. 50:xx + 801 pp., 19 plates, 1904.</p></div>
+
+<span class="smcap">Saunders, A. A.</span><br>
+<div class="references"><p>1911. A study of the nesting of the Cedar Waxwing. Auk, 28(3):323-329, 1911.</p></div>
+
+<div class="references"><p>1912. The probable breeding of the Bohemian Waxwing in Montana. Condor, 14(6):224, 1912.</p></div>
+
+<span class="smcap">Sharpe, R. B.</span><br>
+<div class="references"><p>1885. Catalogue of the birds in the British Museum, Vol. 10, British Mus., xiii + 682 pp., 12 plates, 1885.</p></div>
+
+<span class="smcap">Shaw, W. T.</span>, and <span class="smcap">Culbertson, A. E.</span><br>
+<div class="references"><p>1944. A flock of Cedar Waxwings meets tragedy. Condor, 46(4):205-206, 1944.</p></div>
+
+<span class="smcap">Shufeldt, R. W.</span><br>
+<div class="references"><p>1887. A review of the muscles used in the classification of birds. Jour. Comp. Med. and Sur., 8(4):321-344, 1887.</p></div>
+<div class="references"><p>1889a. Comparative osteology of the families of North American birds. Jour. Morph., 3(1):81-114, 6 plates, 1889.</p></div>
+<div class="references"><p>1889b. Studies on the Macrochires, morphological and otherwise, with the view of indicating their relationships and defining their several positions in the system. Linn. Soc. London, Jour., 20(122):299-394, 1889.</p></div>
+<div class="references"><p>1890. The myology of the Raven. Macmillan &amp; Co., x + 344 pp., 76 figs., 1890.</p></div>
+<div class="references"><p>1909. Osteology of birds. New York State Mus. Bull., 130:381 pp., 1909.</p></div>
+
+<p><span class="pagenum"><a name="Page_529" id="Page_529">[Pg&nbsp;529]</a></span></p>
+<span class="smcap">Skutch, A.</span><br>
+<div class="references"><p>Manuscript&mdash;unpublished notes and personal correspondence.</p></div>
+
+<span class="smcap">Stevenson, H.</span><br>
+<div class="references"><p>1882. On the plumage of the waxwing, <i>Ampelis garrulus</i>, Linnaeus, from the examination and comparison of a large series of specimens killed, in Norfolk, in the winter of 1866-'67. Trans. Norfolk and Norwick Naturalists' Soc., 3:326-344, 2 figs. in text, 1882.</p></div>
+
+<span class="smcap">Sutton, G. M.</span>, and <span class="smcap">Burleigh, T. D.</span><br>
+<div class="references"><p>1940. Birds of Las Vigas, Veracruz. Auk, 57(2):234-243, 1940.</p></div>
+<div class="references"><p>1942. Birds recorded in the Federal District and States of Puebla and Mexico by the 1939 Semple Expedition. Auk, 59(3):418-423, 1942.</p></div>
+
+<span class="smcap">Swarth, H. S.</span><br>
+<div class="references"><p>1922. Birds and mammals of the Stikine River region of northern British Columbia and southeastern Alaska. Univ. California Publ. Zoöl., 24(2):125-314, 8 plates, 34 figs. in text, 1922.</p></div>
+
+<span class="smcap">Taylor, W. P.</span><br>
+<div class="references"><p>1918. Bohemian Waxwing (<i>Bombycilla garrulus</i>) breeding within the United States. Auk, 35(2):226-227, 1918.</p></div>
+
+<span class="smcap">Taverner, P. A.</span><br>
+<div class="references"><p>1934. Birds of Canada. Nat. Mus. Canada Bull., 72, series 19, 445 pp., 77 plates, 488 figs. in text, 1934.</p></div>
+
+<span class="smcap">Wayne, A. T.</span><br>
+<div class="references"><p>1924. A remarkable Cedar Waxwing. Auk, 41(3):485, 1924.</p></div>
+
+<span class="smcap">Wetmore, A.</span><br>
+<div class="references"><p>1926. The migrations of birds. Cambridge, Harvard Univ. Press, vii + 217 pp., 1926.</p></div>
+<div class="references"><p>1932. Notes from Dr. R. Ciferri on the birds of Hispaniola. Auk, 49(1):101-108, 1931.</p></div>
+<div class="references"><p>1940a. A check-list of the fossil birds of North America. Smithson. Misc. Coll., 99(4):1-88 pp., 1940.</p></div>
+<div class="references"><p>1940b. A systematic classification of the birds of the world. Smithson. Misc. Coll., 99(7):1-11 pp., 1940.</p></div>
+
+<span class="smcap">Wetmore, A.</span>, and <span class="smcap">Swales, B. H.</span><br>
+<div class="references"><p>1931. The birds of Haiti and the Dominican Republic. U. S. Nat. Mus. Bull. 155:iv + 482 pp., 26 plates, 1931.</p></div>
+
+<span class="smcap">Wheelock, I. G.</span><br>
+<div class="references"><p>1905. Regurgitation feeding of nestlings. Auk, 22(1):54-71, 1905.</p></div>
+
+<span class="smcap">Whittle, H. G.</span><br>
+<div class="references"><p>1928. The biography of a Cedar Waxwing. Bull. NE Bird-Band. Assoc., 4:77-85, 1928.</p></div>
+
+<span class="smcap">Wolfson, A.</span><br>
+<div class="references"><p>1945. The role of the pituitary, fat deposition, and body weight in bird migration. Condor, 47(3):95-127, 1945.</p></div>
+
+<span class="smcap">Wolley, J. J.</span><br>
+<div class="references"><p>1857. On the nest and eggs of the Waxwing (<i>Bombycilla garrula</i> Tamm.). Proc. Zoöl. Soc. London, 25:55-56, 1857.</p></div>
+
+<p>&nbsp;</p>
+<i>Transmitted July 29, 1949.</i><br>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<p><span class="pagenum"><a name="Page_530" id="Page_530">[Pg&nbsp;530]</a></span>
+Mention should be made here of an important paper by Jean Delacour and
+Dean Amadon (1949). The Relationships of <i>Hypocolius</i> (Ibis, 91:427-429,
+plates 19 and 20) which appeared after the present paper by Arvey was
+written. Delacour and Amadon stated that <i>Hypocolius</i>, a monotypic Persian
+genus, should be assigned to the Bombycillidae. Their conclusions (<i>op. cit.</i>:429)
+were as follows: "It might be advisable to set up three subfamilies in the
+Bombycillidae, one for <i>Bombycilla</i>, one for <i>Hypocolius</i>, and a third for the
+silky flycatchers, <i>Ptilogonys</i>, <i>Phainopepla</i> and <i>Phainoptila</i>. Further study may
+show that <i>Dulus</i> can be added as a fourth subfamily.</p>
+
+<p>"Previously the Bombycillidae appeared to be an American group of which
+one genus (<i>Bombycilla</i>) had reached the Old World. Inclusion of <i>Hypocolius</i>
+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."</p>
+
+<div class="text_rt">&mdash;Eds.</div>
+
+
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<div class="center">
+<img src="images/square.png" width="16" height="17" border="0" alt="square" title="square"><br>
+<br>
+23-1019
+</div>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+<hr style="width:65%">
+<p><span class="pagenum"><a name="Pub_i" id="Pub_i">[Pub&nbsp;i]</a></span></p>
+<div class="caption2">UNIVERSITY OF KANSAS PUBLICATIONS</div>
+
+<p>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 <span class="smcap">Exchange Desk, University of Kansas Library,
+Lawrence, Kansas, U. S. A.</span></p>
+
+<div class="blockquot">
+<p><span class="smcap">Museum of Natural History.</span>&mdash;E. Raymond Hall, Chairman, Editorial Committee.</p>
+
+<p>This series contains contributions from the Museum of Natural History.
+Cited as Univ. Kans. Publ., Mus. Nat. Hist.</p>
+</div>
+
+<table width="100%" class="text_lf" summary="UK Publication List">
+<tr><td class="vtop">Vol.&nbsp;1.</td><td class="vtop">1.</td><td>The pocket gophers (genus Thomomys) of Utah. By Stephen D. Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;2.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;3.</td><td>The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp. 93-96, 1 figure in text. August 15, 1946.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;4.</td><td>Hybridization between two species of garter snakes. By Hobart M. Smith. Pp. 97-100. August 15, 1946.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;5.</td><td>Selected records of reptiles and amphibians from Kansas. By John Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;6.</td><td>Kyphosis and other variations in soft-shelled turtles. By Hobart M. Smith. Pp. 117-124. July 7, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;7.</td><td>Natural history of the prairie vole (Mammalian genus Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;8.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;9.</td><td>Additions to the list of the birds of Louisiana. By George H. Lowery, Jr. Pp. 177-192. November 7, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;10.</td><td>A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-216. November 29, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;11.</td><td>Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R. and E. Raymond Hall. Pp. 217-236, 2 figures in text. November 29, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;12.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;13.</td><td>Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure in text. December 10, 1947.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;14.</td><td>A new pocket gopher (Thomomys) and a new spiny pocket mouse (Liomys) from Michoac&aacute;n, Mexico. By E. Raymond Hall and Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.</td></tr>
+
+<tr><td>&nbsp;<span class="pagenum"><a name="Pub_ii" id="Pub_ii">[Pub&nbsp;ii]</a></span></td><td class="vtop">&nbsp;15.</td><td>A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-264, 1 figure in text. August 16, 1948.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;16.</td><td>A new extinct emydid turtle from the Lower Pliocene of Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;17.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;18.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;19.</td><td>Speciation in the Brazilian spiny rats (genus Proechimys, Family Echimyidae). By João Moojen. Pp. 301-406, 140 figures in text. December 10, 1948.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;20.</td><td>Three new beavers from Utah. By Stephen D. Durrant and Harold S. Crane. Pp. 407-417, 7 figures in text. December 24, 1948.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;21.</td><td>Two new meadow mice from Michoac&aacute;n, M&eacute;xico. By E. Raymond Hall. Pp. 423-427, 6 figures in text. December 24, 1948.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;22.</td><td>An annotated check list of the mammals of Michoac&aacute;n, M&eacute;xico. By E. Raymond Hall and Bernardo Villa R. Pp. 431-472, 5 figures in text. December 27, 1949.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;23.</td><td>Subspeciation in the kangaroo rat, Dipodomys ordii. By Henry W. Setzer. Pp. 473-573, 27 figures in text, 7 tables. December 27, 1949.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;24.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;25.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;26.</td><td>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.</td></tr>
+
+<tr><td>&nbsp;</td><td colspan=2>Index. Pp. 605-638.</td></tr>
+
+<tr><td class="vtop">Vol.&nbsp;2.</td><td colspan=2>(Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948.</td></tr>
+
+<tr><td class="vtop">Vol.&nbsp;3.</td><td class="vtop">&nbsp;1.</td><td>The Avifauna of Micronesia, its origin, evolution, and distribution. By Rollin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;2.</td><td>A Quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 46 figures in text. June 29, 1951.</td></tr>
+
+<tr><td>&nbsp;</td><td class="vtop">&nbsp;3.</td><td>Phylogeny of the waxwings and allied species. By M. Dale Arvey. Pp. 473-530, 49 figures in text, 13 tables. October 10, 1951.</td></tr>
+</table>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Phylogeny of the Waxwings and Allied
+Birds, by M. Dale Arvey
+
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+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: ASCII
+
+*** 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 Zoological Committee on
+Nomenclature, states that generic names used by Brisson (1760) are
+valid under the Code. In consequence, the specific name available for
+the Cedar Waxwing, since Brisson is ruled not to be a binomialist, is
+_Bombycilla cedrorum_ Vieillot (1807).
+
+Most workers prior to 1900 utilized the family name Ampelidae to
+include waxwings, silky flycatchers, and palm-chats. Ridgway
+(1904:113) elevated the silky flycatchers to family rank under the
+name Ptilogonatidae, and assigned the palm-chats to a separate family,
+the Dulidae.
+
+
+
+
+MATERIALS
+
+
+The following specimens, numbering 238, and representing each
+currently recognized species and subspecies, were used in the study,
+and were supplemented by observation in 1947 on specimens in the
+United States National Museum.
+
+
+  ====================================================================
+          Species or Subspecies           | Skin | Skeleton| Alcoholic
+  ----------------------------------------+------+---------+----------
+  _Phainoptila melanoxantha melanoxantha_ |    8 |    1    |    2
+  _Phainoptila melanoxantha minor_        |    2 |         |
+  _Ptilogonys cinereus cinereus_          |   13 |    3    |    4
+  _Ptilogonys cinereus molybdophanes_     |    6 |         |
+  _Ptilogonys caudatus_                   |   16 |    3    |    4
+  _Phainopepla nitens nitens_             |      |    1    |    5
+  _Phainopepla nitens lepida_             |   12 |    5    |    4
+  _Bombycilla cedrorum_                   |   53 |   27    |    8
+  _Bombycilla garrula garrula_            |    4 |    3    |
+  _Bombycilla garrula centralasiae_       |    9 |    2    |
+  _Bombycilla garrula pallidiceps_        |    7 |    3    |    2
+  _Bombycilla japonica_                   |   10 |         |
+  _Dulus dominicus dominicus_             |    9 |    5    |    2
+  _Dulus dominicus oviedo_                |    4 |    1    |
+                                          |---------------------------
+      Totals                              |  153 |   54    |   31
+  --------------------------------------------------------------------
+
+
+
+
+DIAGNOSES
+
+
+Family Bombycillidae
+
+_Diagnosis._--Bill short, flat, somewhat obtuse, minutely notched near
+tip of each maxilla, flared at base; gape wide and deeply cleft;
+culmen convex; nasal fossa broad, exposed, or filled with short, erect
+or antrorse, close-set velvety feathers; nostril narrowly elliptical;
+rictal vibrissae long, short, or absent; lacrimal bone free,
+articulating at two points; wings long and pointed, or short and
+rounded; primaries ten, tenth reduced in some species; tail short,
+narrow, even, two thirds or less length of wing, or much longer and
+forked or rounded; feet weak (except in _Dulus_ and _Phainoptila_);
+tarsus generally shorter than middle toe and claw, distinctly
+scutellate with five or six divisions, the lateral plate subdivided
+(except in _Phainoptila_); lateral toes of nearly equal length; hallux
+approximately as long as inner lateral toe, or shorter; basal phalanx
+of middle toe more or less united to that of outer and inner toes;
+body stout; head generally conspicuously crested; plumage soft, smooth
+and silky (except in _Dulus_); eggs spotted; nest in trees; three
+subfamilies, five genera, eight species.
+
+
+Subfamily Ptilogonatinae
+
+_Diagnosis._--Rictus with conspicuous bristles; nasal fossa almost
+entirely exposed; tail long and rounded, graduated, or square; caudal
+muscles and pygostyle well developed; wings rounded and short, first
+primary a half to a third as long as second; second primary shorter
+than third; humerus long, with small external condyle; plumage soft
+and silky, less so in _Phainoptila_; sexes dissimilar, young like
+adult female; three genera, four species.
+
+
+Genus =Phainoptila= Salvin
+
+  _Phainoptila_ Salvin, Proc. Zool. Soc. London, 1877:367, April 17,
+    1877. Type _Phainoptila melanoxantha_ Salvin.
+
+_Diagnosis._--Without crest; tarsus longer than middle toe and claw,
+and booted or very slightly reticulate; tail shorter than wing,
+rounded; nostril exposed, ovate; rictal bristles distinct; first
+primary well developed; plumage normal, bill flared slightly at base.
+
+_Range._--Costa Rica and Panama.
+
+
+=Phainoptila melanoxantha melanoxantha= Salvin
+
+Phainoptila
+
+  _Phainoptila melanoxantha melanoxantha_ Salvin, Proc. Zool. Soc.
+    London, 1877:367; April 17, 1877.
+
+_Diagnosis._--Coloration of adult males: Pileum, hindneck, back,
+scapulars, and upper tail coverts Black (capitalized color terms after
+Ridgway, Color Standards and Color Nomenclature, Washington, D. C.,
+1912), with Bluish Gray-Green gloss; rump Lemon Yellow tinged with
+Olive; lower breast and abdomen Gull Gray or Slate Gray; sides and
+flanks clear Lemon Yellow; lower chest, upper breast, and under tail
+coverts Yellowish Olive-Green, extending to patch on sides and flanks
+of same color; bill and feet Black or Blackish Brown. Coloration of
+adult females: Most of upper parts Olive-Green, with Yellowish Olive
+on rump; thighs Olive-Gray, as are sides of head; rest of coloration
+as in male. Coloration of young: As in adult female, but duller
+throughout.
+
+_Measurements._--Wing 99.0, tail 88.5, culmen 15.2, tarsus 28.4.
+
+_Range._--Highlands of Costa Rica and extreme western Panama (Volcan
+de Chiriqui).
+
+
+=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 Panama (Cerro Flores and eastern
+Chiriqui).
+
+
+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, Mexico, Oaxaca, Colima,
+Morelos, Veracruz, San Luis Potosi, Guerrero and Michoacan.
+
+
+=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. fuer 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 Panama.
+
+
+Genus =Phainopepla= Sclater
+
+  _Phainopepla_ Sclater, Proc. Zool. 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 Luis
+Potosi, Durango, Guanajuato, Mexico, 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
+Panama.
+
+
+=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 deg. 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 Zoology, 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 habitues 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 Luis 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 Luis 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 Luis 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 Luis 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. x 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 Luis 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. x 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 Luis 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. x 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. x 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.
+
+
+
+
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+
+    1928.  Variations in the Fox Sparrow (_Passerella iliaca_) with
+           reference to natural history and osteology. Univ. California
+           Publ. Zool., 30(12):251-392, 4 plates, 38 figs. in text,
+           1928.
+
+LITTLEFIELD, M. J., and LEMKAN, F.
+
+    1928.  History of a Cedar Waxwing family. Bull. NE Bird-Band.
+           Assoc., 4:85-89, 1928.
+
+LUCAS, F. A.
+
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+           1001-1019 pp., 2 plates, 1897.
+
+MCGREGOR, R. C.
+
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+           Manila. Condor, 8(4):98-100, 1906.
+
+MATTHEW, W. D.
+
+    1939.  Climate and evolution. N. Y. Acad. Sci., Spec. Publ.,
+           1:xi + 223 pp., 1939.
+
+MAYR, E.
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+           xiv + 334 pp., 29 figs. in text, 1942.
+
+    1947.  Ecological factors in speciation. Evolution, 1(4):263-288,
+           1947.
+
+MERRIAM, F. A.
+
+    1896.  Nesting habits of _Phainopepla nitens_ in California. Auk,
+           8(1):38-43, 1896.
+
+MILLER, A. H.
+
+    1933.  Postjuvenal molt and the appearance of sexual characters of
+           plumage in _Phainopepla nitens_. Univ. California Publ.
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+
+    1937.  Structural modifications in the Hawaiian Goose (_Nesochen
+           sandvicensis_). A study in adaptive evolution. Univ.
+           California Publ. Zool., 42(1):1-80, 6 plates, 12 figs. in
+           text, 1937.
+
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+
+MULLER, C. S.
+
+    1915.  A northern winter record of the Phainopepla. Condor,
+           17(3):129, 1915.
+
+MYERS, H. W.
+
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+
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+           Condor, 10(2):72-75, 1908.
+
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+           11(1):22-23, 1909.
+
+NEWTON, A., and GADOW, H.
+
+    1893-1896. A dictionary of birds. Adams and Charles Black,
+           xii + 1086 pp., 1893-1896.
+
+NICE, M. M.
+
+    1940.  Observations on the behavior of a young Cedar Waxwing.
+           Condor, 43(1):58-64, 1940.
+
+OBERHOLSER, H. C.
+
+    1917.  A synopsis of the races of _Bombycilla garrula_ (Linnaeus).
+           Auk, 34(3):330-333, 1917.
+
+PEMBERTON, J. R.
+
+    1908.  Northern range of the _Phainopepla_. Condor, 10(6):238, 1908.
+
+PLATH, K.
+
+    1933.  Molt of the Nonpareil. Auk, 50(2):121, 1933.
+
+POST, K. C.
+
+    1916.  The Cedar Waxwing (_Bombycilla cedrorum_) during July and
+           August, 1916. Wilson Bull., 28:175-193, 1916.
+
+RAND, A. L., and RAND, R. M.
+
+    1943.  Breeding notes on _Phainopepla_. Auk, 60(3):333-341, 1943.
+
+RICHARDSON, F.
+
+    1942.  Adaptive modifications for trunk foraging in birds. Univ.
+           California Pub. Zool., 46(4):317-368, 2 plates, 16 figs.
+           in text, 1942.
+
+RIDGWAY, R.
+
+    1904.  The birds of North and Middle America, Part III. U. S. Nat.
+           Mus. Bull. 50:xx + 801 pp., 19 plates, 1904.
+
+SAUNDERS, A. A.
+
+    1911.  A study of the nesting of the Cedar Waxwing. Auk,
+           28(3):323-329, 1911.
+
+    1912.  The probable breeding of the Bohemian Waxwing in Montana.
+           Condor, 14(6):224, 1912.
+
+SHARPE, R. B.
+
+    1885.  Catalogue of the birds in the British Museum, Vol. 10,
+           British Mus., xiii + 682 pp., 12 plates, 1885.
+
+SHAW, W. T., and CULBERTSON, A. E.
+
+    1944.  A flock of Cedar Waxwings meets tragedy. Condor,
+           46(4):205-206, 1944.
+
+SHUFELDT, R. W.
+
+    1887.  A review of the muscles used in the classification of birds.
+           Jour. Comp. Med. and Sur., 8(4):321-344, 1887.
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+           birds. Jour. Morph., 3(1):81-114, 6 plates, 1889.
+
+    1889b. Studies on the Macrochires, morphological and otherwise,
+           with the view of indicating their relationships and
+           defining their several positions in the system. Linn. Soc.
+           London, Jour., 20(122):299-394, 1889.
+
+    1890.  The myology of the Raven. Macmillan & Co., x + 344 pp.,
+           76 figs., 1890.
+
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+           1909.
+
+SKUTCH, A.
+
+    Manuscript--unpublished notes and personal correspondence.
+
+STEVENSON, H.
+
+    1882.  On the plumage of the waxwing, _Ampelis garrulus_, Linnaeus,
+           from the examination and comparison of a large series of
+           specimens killed, in Norfolk, in the winter of 1866-'67.
+           Trans. Norfolk and Norwick Naturalists' Soc., 3:326-344,
+           2 figs. in text, 1882.
+
+SUTTON, G. M., and BURLEIGH, T. D.
+
+    1940.  Birds of Las Vigas, Veracruz. Auk, 57(2):234-243, 1940.
+
+    1942.  Birds recorded in the Federal District and States of Puebla
+           and Mexico by the 1939 Semple Expedition. Auk,
+           59(3):418-423, 1942.
+
+SWARTH, H. S.
+
+    1922.  Birds and mammals of the Stikine River region of northern
+           British Columbia and southeastern Alaska. Univ. California
+           Publ. Zool., 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. Zool. 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 Michoacan, 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 Joao 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 Michoacan, Mexico. By E. Raymond
+            Hall. Pp. 423-427, 6 figures in text. December 24, 1948.
+
+        22. An annotated check list of the mammals of Michoacan,
+            Mexico. 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         => Luis
+
+    Page 480, 481         : Luis Potosi  => Luis 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. Dale Arvey
+
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