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THE
CAMBRIDGE NATURAL HISTORY
EDITED BY
3. F. HARMER, Sc.D., F.R.S., Fellow of King’s College, Cambridge ; Superintendent of the University Museum of Zoolog
AND
A. E. SHIPLEY, M.A., Fellow of Christ’s College, Cambridge ; University Lecturer on the Morphology of Invertebrates
VOLUME IX
BIRDS
we wes = : By At: EVANS, M.A., Clare College, Cambridge
London MACMILLAN AND CO., LIMITED
NEW YORK: THE MACMILLAN COMPANY
1899
All rights reserved
In sicco ludunt fulicae.— VIRGIL.
‘* Loons disport themselves on dry matters.”
AY HSO he 4
AUG 6 1958 LIBRARY
Page
ERRATA
4, note 2, for Water-hens read Moor-hens. 10, line 19, after Owls vead and Pandion.
17, for Lord Howe’s read Lord Howe. 22, for Galapagos read Galapagos.
26, note 1, delete comma after Bronn’s. 30, line 2 from bottom, for Tarapaca read Tarapaca.
©» 6 a for Pelecanoidinae read Pelecanoidinae.
60, lines 14, 26, 34, for Pelecanoides read Pelecanoides.
67, line 6, for Thalassaeca read Thalassoeca.
16s. 43 aes 59, , TO? Oi s, ABN os, 122, ,, 138;. 3 160, ,, Oils 258, ,, Bolen BOUIN ss
10, for Phaenicopteridae read Phoenicopteridae. 12, for radloides read ralloides. 17, for dominicus read dominica. 2, after F. cristata, read the Tufted Duck. 5, after Wavy, read or Snow Goose. 8 from bottom, for cirrhatus read cirratus. 10, for praelatus read praclata. 15, for perhaps read probably not. 11, and page 357, line 6, for Phaenicophainae read Phoenicophainae. 11 from bottom, for Phaenicophaés read Phoenicophaés.
429, Fig. 89, for gugularus read jugularis.
550, line 20, for Seenopocetes read Scenopocetes.
568, ,,
9 from bottom, for a scale-insect vead an Aphid.
598.2
Ee a2
, (-
Birds
PREFACE
In this volume of the “ Cambridge Natural History” the author has attempted to meet a need which he beheves to be some- what widely felt. Recognising the fact that there is at the present time an abundance of popular, or only slightly scien- tific, works on Birds, some of which touch but superficially upon the individual species composing the various groups, as regards their plumage or habits, while others pay little or no attention to correctness of Classification, he has essayed the difficult and apparently unattempted task of including in some six hundred pages a short description of the majority of the forms in many of the Families, and of the most typical or important of the innumer- able species included in the large Passerine Order. Prefixed to each group is a brief summary of the Structure and Habits; a few further particulars of the same nature being subsequently added where necessary, with a statement of the main Fossil forms as yet recorded.
Thus it is hoped that the work may be of real use, not only to the tyro in Ornithology, but also to the traveller or resident in foreign parts interested in the subject, who, without time or opportunity for referring to the works of specialists, may yet need the aid of a concise account of the species likely to cross his path.
An introductory chapter has been written, to meet the claims of the present day, on the external and to a limited extent on
the internal structure of Birds, with short paragraphs on Classi-
&
vl PREFACE
fication, Geographical Distribution, and Migration, and a “ Ter- minology ” of the subject.
In accordance with the scheme of the Series generally, the order followed runs from the lowest forms and the Ratite Birds upwards; the Carinate Birds being divided, after Dr. Gadow’s plan, into two Brigades or main sections, and these again into Legions, Orders, and so forth. It should, however, be under- stood that the Species of each Genus are often merely placed in the most convenient order; and that, where a geographical range is given, it does not follow that it is unbroken from end to end.
In descriptions of colour, the names used for tints in the British Museum Catalogue of Birds have been commonly adopted, or for British species those in Mr. Howard Saunders’ Manual of British Birds.
Various subjects of a highly technical, or at least of a special character, have purposely been avoided in the main, as unfitted to the scope of the work; such are, Variation and Hybrids, with their accompaniments of Dimorphism, Dichromatism, and the like; Myology; Mechanism of Flight and the supposed Lines of Flight on Migration; the Classifications of Linnzus and the older writers; and the Strickland Code of Ornithological Nomen- elature. For these Professor Newton’s Dictionary of Birds, and especially the Introduction to it, may be consulted; besides a multitude of other works.
The woodcuts have been chiefly supphed by Mr. G. E. Lodge; but a few illustrations have been utilized from other sources.
The author does not hold himself responsible for the fact of the Family names being in Roman in place of Italic type, nor for the dissociation of the vowels in the diphthongs; in these minor points he personally differs from the writers of the former volumes, though he agrees with the wish of his Editors for
uniformity.
PREFACE vii
In conclusion, he must take the opportunity of acknow- ledging the invaluable assistance afforded by Mr. Howard Saunders, who carefully went over the whole of the proofs, while Dr. R. B. Sharpe was kind enough to do the same; nor must he fail to record his indebtedness to Professor Newton, Mr. Sclater, Dr. Gadow, Mr. Ogilvie Grant, and many others, not to mention the innumerable authors without whose previous labours to write a book of this description would be a well- nigh impossible task. Dr. Stejneger’s Volume on Birds in the
Standard Natural History should be mentioned in particular. A. H. Evans.
CAMBRIDGE, November 17, 1898.
ADDENDUM
Since the text has been printed off, several new species have been described, and of these it 1s necessary to mention at least the following ;—
Archaeopteryx siemensi, from Solenhofen, where the original form was obtained.—( Dames. )
Euryapteryx exilis (Dinornithidae) ; a new genus, Anomalornis, is also proposed for Anomalopteryx (preoccupied ).— (Hutton. )
Ammoperdix cholmleyi (Phasianidae), from Suakin.—(Ogilvie- Grant.)
Cepphus snowi (Alcidae), from the Kurile Is.—(Stejneger.) The range of C. columba will now be “ Bering Sea to California ; ” and of C. carbo “ North-East Asia and Japanese Seas.”
Podoces pleskii (Corvidae), from East Persia.—(Zarudny.)
Some new fossil forms from Patagonia.—(Mercerat. )
Mr. F. E. Blaauw has published a Monograph of the Cranes, and Mr. C. W. de Vis has described the eggs and young of Salvadorina (Anatidae).
In all these cases the Zoological Record for 1897 may be consulted.
CONTENTS
PAGE PREFACE : ; ; : ; : q , . : : . Vv ADDENDUM . s : : ; ; : : ; 5 2 villi SCHEME OF THE CLASSIFICATION ADOPTED IN THIS Book 2 ; g : xi CEPA Mh INTRODUCTION ; : : A 3 ; < , : 3 , al @UEPAREZ IME ie sul ARCHAEORNITHES—NEORNITHES RATITAE—NEORNITHES ODONTOLCAE. 5 eB CHEPATE ER lll NEORNITHES CARINATAE BricapE I—Lecion I (CoLyMBOMORPHAE). OrDERS: ICHTHYORNITHES— CoLYMBIFORMES—SPHENISCIFORMES—PROCELLARIIFORMES . : : 48 CEUAIR VERVE NEORNITHES CARINATAE CONTINUED BricADE I—Lercion II (PELARGOMORPHAE). ORDERS: CICONITFORMES— ANSERIFORMES—FALCONIFORMES . : Q : ; 5 : : 70
CHAPTER V
NEORNITHES CARINATAE CONTINUED
BricapE II]—Lxrcrion I (ALECTOROMORPHAE). OrbERsS: TINAMIFORMES— GALLIFORMES—GRUIFORMES—CHARADRIIFORMES . : , ; =~ 1182
x CONTENTS
CHAPTER VI
NEORNITHES CARINATAE CONTINUED
BricgapE II—-Lecron II (CorAcIOMORPHAE). ORDERS: COoRACIIFORMES
CHAPTER VII
CucULIFORMES—
NEORNITHES CARINATAE CONCLUDED
BricapE I]—Lraton II (CorAcIOMORPHAE CONCLUDED). FORMES
INDEX
ORDER : PASSERI-
PAGE
351
466
2
‘6 “JOA Atoisty TeameN aspieques
Ipy MemopouyTE TOP
i
(aang mmpog yaiony go @pors yp FT) “000 '099'SG OF T FAMIMN Jo PTHOG
D UVTOd HLA
[Par wage ier |
cae wma J? FULT
Mecha cg
Me By ae nye PAE IES
SSeS } Prebr Repe 7
Lath ee
SIOPE AY (ee ———gawotod 7
, | Py aprsOnl \ Wee: 3 on (eo) prumegrs ex |
Bie? ee )
le $ — &
&
a : } soul 7,
ftss
‘[ amuai? TS cee /
i] on @ Al ceeg p18 pe SAVED avid BIOs ~
SCHEME OF THE CLASSIFICATION ADOPTED IN THIS BOOK
CLASS AVES (p. 23.)
SUB-CLASS I. ARCHAEORNITHES (p. 23).
3)
Archaeopteryx (pp. vill, 23).
SUB-CLASS II. NEORNITHES (}). 25). Division A. NEORNITHES RATITAE (p. 25). I. Struthiones (p. 27): Fam. Struthionidae (p. 27). | II. Rheae (p. 30): Fam. Rheidae (p. 30). Nira cued e9,, fam. I. Casuariidae (p. 33). RATITAE (p. 26) < Gs lee Stns Aub 27) \ Fam. 11. Dromaeidae (p. 36). IV. Apteryges (p. 388): Fam. Apterygidae (p. 38). V. Dinornithes (p. 41): Fam. Dinornithidae (p. 41). VI. Aepyornithes (p. 43): Fam. Aepyornithidae (p. 43). Mesembriornis, etc. (p. 44).
; Diatryma (p. 45). a erenzonntraes| 2 [srr © 45).
eS 2 KBR) Eee (p. 45).
Gastornis (j). 45).
Division B, NEORNITHES ODONTOLCAE (p. 45).
HESPERORNITHES (p. 46). ? ENALIORNITHES (p. 46).
? Baptornis (p. 46). Division C. NEORNITHES CARINATAE (p. 48). ICHTHYORNITHES f Fam. IcHTHYORNITHIDAE (p. 48): Jchthyornis (p. 48).
(p- 48) l ’ Apatornis (p. 49). Order. Sub-Order. Family. Sub-family. Colymbi (p. 49) CoLYMBIDAE (p. 50). Te barca i Podicipedes PoDICIPEDIDAE Ms (p. 49) (p. 52), SPHENISCIFORMES Sphenisci SPHENISCIDAE (p. 54) (p. 54) (p. 54).
( Diomedeinae (p. 63). PROCELLARIIFORMES Tubinares PROCELLARIIDAE | Oceanitinae (p. 65). ' (p. 59) (p. 59) (p. 59) ) Procellariinae (p. 65). | Pelecanoidinae (p. 68).
xl SCHEME OF CLASSIFICATION
Order. Sub-Order. Family. Sub-family. ‘ PHAETHONTIDAE (p. 72). [scrips (p. 73). Steganopodes PHALACROCORACIDAE (p. 75). (p. 70) [ee ATIDAE (p. 81). PELECANIDAE ( (P- 83). j ARDE IDAE (p. 87).
: SCOPIDAE (p. 95 CICONTFORMES arden (p80) } Srcovitpal in 3) en: { Ibidinae (p. 100). Ciconiae (p. 95) || Totpipat ( 1b oe) \ Plataleinae (. 103). pe OPTERIDAE Phoenicopteri | (p. 105). (p. 105) ac euGe att | (pp. 105, 108). Palamedeae PALAMEDEIDAE (p. 108). (p. 108) Merginae (p. 115). Merganettinae (p. 116). Erismaturinae (p. 117). Fuligulinae (p. 118). ANSERIFORMES Anatinae (p. 123). (p. 108) Anseres (p. 110) ANATIDAE(p.111)¢ Chenonettinae (p. 130). Anserinae (p. 131). Cereopsinae (p. 133). Plectropterinae (p. 133). Anseranatinae ue 135). Cygninae (p. 135) Cathartae CATHARTIDAE (p.137) (p. 137). ( SERPENTARIIDAE (p. 141). VULTURIDAE (p. 143). FALCONIFORMES Gypaétinae (p. 150). (p. 137) Accipitres } Polyborinae (p. 151). (pp. 137, 141) FALCONIDAE Accipitrinae (p. 153). (p. 146) Aquilinae (p. 159). Buteoninae (p. 164). Falconinae (p. 178). PANDIONIDAE | (p. 180). TINAMIFORMES Tinami (p. 182) TINAMIDAE (CRYPTURIDAE) (p. 182) (p. 182). Mesitae (p. 186) MESITIDAE (p. 186). ‘TURNICIDAE Turnices (p. 187) | (pa Teo: SNe: a PE eno p- 189). ten A PoaIae (p: 90): ae (p. 196). racinae (p. EE TN es) ) Galli (p. 190) CRACIDAE (p. 194) ) Penelopinae (p. 197). (p. 186) Oreopl e (p. 198). phasinae (p. z ( Numidinae (p. 204). I EES SHADIN Meleagrinae (p. 206), (p. 198) >, Phasianinae (p. 206). [ Odontophorinae (p. 230). Tetraoninae (p. 233).
em OPISTHOCOMIDAE (p. 241) (p. 241).
SCHEME OF
CLASSIFICATION
xii
Sub-Order. Family.
RALLIDAE (p. 2
Order.
GRUIFORMES (p. 243)
OTIDIDAE (p.
RUINOCHETIDAE (p. 263).
EURYPYGIDAE (p. 265).
(p. 267).
( OHARADRIIDAE 272)
| ie ray | . 295 2) ise GL AREOLA © J vi wT Limicolae (p. 268) 5 p. 293) | ee aoe THIDAE (p. | OEDICNEMIDAE [ (p. 297).
PARRIDAE(p. 297)
CHARADRIIFORMES | ,
WH? Pe) 7 Lari (pp. 268, 300) LaAnripAk (p. 300) Alcae (p. 315) Pterocles (p. 321)
PTEROCLIDAER
(p. 321). DIDIDAE (p. DIDUNCULIDAE
(p. 331).
| CoLUMBIDAE (p. 333)
328
Columbae (p. all
CUCULIDAE
Cuculi (p. 351) (p. 351)
- CUCULIFORMES MvsoOPHAGIDAE (p. 351) \ (p. 359). PSITTACI YATE (p. 366)
Psittaci (p. 361) | TRICHOGLOSSIDAE
(p. 373)
. CORACIIDALE (p. 376) on
(p. 379)
CORACIIFORMES
(p. 376) J Coraciae (p. 376) \
(Continued on the neat page.)
243). GRUIDAE (p. 281). ARAMIDAE (p. 256). PsorHIIDAE (p. 257). CARTAMIDAE(]). 25 260).
HELIORNITHIDAE
996)
ALCIDAE (p. 315).
| Columbinae ().
Sub-family.
8).
Tringinae (p. 278
hee varadriinae (p. 272). ). | Scolopacinae (p. 289).
{ Glareolinae (p. 29 | Dromadinae (p.
( Stercorariinae (p. 504 Larinae (p. 305).
| Rhynchopinae (p. 310).
Sterninae (p. 310).
freee p. 334). Peristerinae (p. 334). 342).
Treroninae (p. 34 4).
Centropodinae (p. 356). Phaenicophainae (p. 357).
| Cuculinae (p. 352).
Diplopterinae (p. 359).
_Crotophaginae (p. 3859).
| Neomorphinae (p. 357).
if Stringopinae (p. 366). Psittacinae {p. 367).
\e Cacatuinae (p. 372).
frelon cree (p. 373): Loriinae (p.
Q79) Ov). \ Nestorinae (p.
374).
{ Coraciinae (p. 376).
| Leptosomatinae (p. 378). Momotinae (p. 380). Todinae (p. 381).
XIV
SCHEME OF CLASSIFICATION
Order.
CORACIIFORMES (continued)
Sub-Order.
Coraciae (contin ued )
Striges (p. 397) |
Cypseli (p. 419) |
Caprimulgi (p. 415)
Colii (p. 439)
Trogones (p. 441)
Family. ALCEDINIDAE (p. 382) MEROPIDAE
(p. 387). BuCcEROTIDAL
(p. 390). UpuPIDAE
(p. 895) STRIGIDAE
(p- 398) CAPRIMULGIDAE
(p. 417) PODARGIDAE
(p. 419). STEATORNITHI-
DAE (p. 419). CYPSELIDAE
(p. 420)
TROCHILIDAE (p. 426), CoLIIDAE (p. 439). TROGONIDAE (p. 441).
, GALBULIDAE
Pici (p. 445)
(p. 445) CAPITONIDAE (p. 448)
|) RHAMPHASTIDAE
(p. 458)
PICIDAE (p. 457)
Family.
Sub-family. { Haleyoninae (p. 385). | Alcedininae (p.386).
{ Upupinae (p. 395). | Irrisorinae (p. 397). { Striginae (p. 403), z_\ Buboninae (p. 404). | f Caprimulginae (p. 418). | Nyctibiinae (p. 418).
( Macropteryginae (p. 422). Chaeturinae (p. 422). ( Cypselinae (p. 424).
Galbulinae (p. 445). Bucconinae (p. 446).
Capitoninae (p. 448). Indicatorinae (p. 451).
{ Picinae (p. 457). | lynginae (p. 464).
Sub-family.
( SUBCLAMATORES EURYLAEMIDAE
Order. Group. Division. (p. 467) PASSERI- Passeres FORMES - aniso- } CLAMATORES (p. 466) | myodae (p. 469) ; (p. 467)
(Continued on the nest
( page.)
(p. 467). ( PITTIpDAE (p. 469).
PHILEPITTIDAE
(p. 471). XENICIDAE (p. 472).
TYRANNIDAE (p. 473)
a
(p. 477). PIPRIDAE (p. 477).
CorTINGIDAE (p. 479)
Taeniopterinae
J} Platyrhynchinae Elaineinae Tyranninae
.
| ! | 473. J
OXYRHAMPHIDAE
Tityrinae
| Lipanginae Attilinae
‘| Rupicolinae Cotinginae Gymnoderinae
| (p- ls
PHYTOTOMIDAE
(p. 483).
SCHEME OF CLASSIFICATION XV Order. Group. Division. Family. Sub-family. ( Furnariinae Dr eae OLAPTI- [syntax | (p. Daw (p- 488) | Dendkocolap- (a tinae Passeres aniso- J CLAMATORES =} eee | myodae (continued) ORMICARIIDAE nae K (p. pane pega ». 488) Formicariinae { 488). Keontmatied) eae | CONOPOPHAGIDAE (p. 489). PrEROPTOCHIDAE \ \ (p. 490). MENURIDAE ( SUBOSCINES | (p. 491). (p. 491) Nae RICHORNITHI- DAE (p. 493). ALAUDIDAE (p. 496). MOTACILLIDAE — § Motacillinae | (p. 498 (p. 498) YAnthinae f\P ) HENICURIDAE (p. 501). TIMELIIDAE (p. 501). PYCNONOTIDAE (p. 504). >ASSERI- MUSCICAPIDAE FORMES 4 (p. 506). (continued) Turdinae (p. 509), ite Myiodectinae . 513) (p. 509) > Sylviinae (p. 513). [ Retioptitinae (p. 514). Miminae (p. 514), CINCLIDAE Ds, O9)). Roeecres i OS Tt ane jacro- | OSCINES J (p. 521). ea kpa22s) | CHAMAEIDAE (p. 522). HIRUNDINIDAE
: |
.
Continued on the next page. )
(p. 522). CAMPEPHAGIDAE (p. 525). DICRURIDAE (p. 527). AMPELIDAE (p. 529). ARTAMIDAE (p. 530). Gymnorhininae (p. 532). Malaconotinae (p. 533). Pachycephalinae (p 533). Laniinae (p. 534). Prionopinae (p. 535).
LANIIDAE
(p. 531)
ed
XV1
SCHEME OF CLASSIFICATION
Order. Group.
Passeres diacro- myodae
(continwed)
PASSERI- FORMES < (continued )
| 1
Division.
OScCINES
(continued)
Family. Sub-family.
, VIREONIDAE
(p. 536). SITTIDAE (p. 586). PARIDAE (p. 538). PANURIDAE (p. 541). ORIOLIDAE (p. 542). PARADISEIDAE (p. 543). Corvinae Garrulinae | Fregilinae
CorRVIDAE (p. 552)
(p. 552).
STURNIDAE (os G5): DREPANIDIDAE (p. 562). MELIPHAGIDAE (p. 564) ZOSTEROPIDAE (p. 568). NECTARINIIDAE (p. 568). DICAEIDAE p. 570). CERTHIIDAE (p. 571). CoOEREBIDAE p- 572). SIOTILTIDAE Dod: TANAGRIDAE Dao). PLOCEIDAE (p. 576)
{ Myzomelinae \ (p. \ Meliphaginae J 564).
M
4,
Viduinae (p. 576). Ploceinae (p. 577).
{ i} Cassicinae Agelaeinae ape
ICTERIDAE ). 579) J Sturnellinae - We Icterinae |
(Juiscalinae
FRINGILLIDAE
ea eee | 582).
CHAP THRs INTRODUCTION
Definition.—“ A Bird is a feathered biped.” This popular saying undoubtedly furnishes a definition in the world of to-day, since no other existing creature has a clothing of feathers, and even the word “ biped” is thus superfluous.
The above should, however, be somewhat expanded, in order to shew in greater detail the differences between Birds and other Vertebrata. Care must nevertheless be taken to avoid the fault common to many modern definitions, of giving an abstract of the main characteristics of the object, rather than a clear guide to distinction.
Dr. Gadow ' defines Birds as “ oviparous, warm-blooded, amniotic Vertebrates, which have their anterior extremities transformed into wings. Metacarpus and fingers carrying feathers or quills. With an intertarsal joint. Not more than four toes, of which the first is the hallux.” .
Much of this the beginner might well postpone, his attention being solely drawn to the external characters; though of course those that are internal are by no means to be subsequently neglected. Indeed no satisfactory progress can be made in the serious study of Ornithology, or the Science of Birds, without a competent knowledge of their Anatomy and Development ; while, though at present comparatively few fossil remains of Birds have been found, some of them are of the highest importance, and there is every probability of future discoveries throwing much light not only on the mutual relationships of Birds among themselves, but also on their connexion with the Reptilia. Birds are, in fact, only extremely modified Reptiles, the two Classes forming the Sawr- opsida of Huxley, one of his three primary divisions of Vertebrata.
1 P.Z.8. 1892, p. 236. VOL. IX =z B
2 FEATHERS CHAP.
The aid of the Palaeontologist and Geologist must thus be called in to clear up many problems which present themselves to the Ornithologist who does not content himself with examining exist- ing forms of life alone. Archaeopteryx (p. 23) from the Jurassic System is the oldest Bird known, nor are any other pre-Tertiary forms recorded, save a small number from the rocks of the Creta- ceous Epoch, the chief of which are the so-called Odontornithes, or toothed species of America (p. 49).
The following paragraphs on the structure of Birds will help to explain the systematic account in the later chapters.
Feathers.—Returning to the outward character denoted by the popular saying with which we began, the Feathers’ con- stituting the plumage may not inconveniently be first considered. The general belief that they grow from almost every part of a Bird’s body, as do hairs in most Mammals, is erroneous; for, almost without exception, they grow in certain definite tracts called pterylae, the intervening spaces, whether they be wholly bare or covered with down, being termed apteria. The arrange- ment of these patches is at times of considerable assistance in determining a Bird’s affinities ; and the subject may be studied in Nitzsch’s Pterylographie* or in a shorter form in Dr. Gadow’s article “ Pterylosis” in Professor Newton’s Dictionary of Birds.
A feather originates thus. A conical papilla arises in the derma and pushes up the epidermis, a depression forming mean- while around the base; subsequently the derma supphes a nutritive pulp, while part of the epidermal layer is converted into a tuft of stiff rays, meeting and forming a short tube below ; these thereafter burst their covering and protrude as the rami or barbs, on which, apparently by secondary splitting, are commonly produced radii or barbules. In this state we have a “plumule” or “down-feather” ; but in the case of the feathers that have “webs” or “vanes” (veailla) often called contour feathers (pennae or plumae), afresh papilla forms at a deeper level, so that the earlier structure is thrust forward and eventually drops off from the apex of the later. Meanwhile the “ dorsal” portions of
1 The integument of a Bird consists of Skin and Feathers, the former being composed of a superficial epidermis and an underlying derma or cutis, which is rich in sensory organs but poor in blood-vessels. The epidermis itself has a horny outer layer and a softer (Malpighian) substratum. Feathers, hairs, bristles, scales, claws and bill-sheaths are epidermal structures.
* A translation was edited for the Ray Society by Mr. Sclater in 1867.
I COLOUR
Os
the barrel or quill (calamus or seapus) at the base of the tuft of rays have elongated into a principal shaft (rhachis) ; this is generally accompanied by a secondary “ aftershatt ° (hyporhachis), originating from the “ ventral” side, which in the Emeu and Cassowary rivals the shaft itself in size. On the rhachis a double series of lamellae or barbs are developed, carrying a similar double series of barbules, much as in the down-feather, but the barbules again give rise to barbicels (cia), which in the distal rows usually terminate in hooklets (damuli). These catch in the folded margins of the next proximal row, and a firm surface is thus secured, An after-shaft never, and a down-feather rarely, possesses barbicels; while in some cases by the absence of these and part of the barbules a “ disconnected ” web and a “ decomposed ” feather are formed, as in the decorative tufts of many species. The barbs may even be absent, as in the wing-quills of Cassowaries, the “wires” of Birds-of-Paradise, the “ bristle-feathers” at the gape of Night-jars or the eyelashes of Hornbills. In the hackles of Gallus (Fowl), and the secondaries or even the tail-feathers of Ampelis (Waxwing), the tip of the rhachis is flattened and wax- like; and similar structures are observable elsewhere. In the newly-hatched young the down is often partly or entirely sup- pressed, but in certain Birds this suppression is temporary, and a thick coat grows after a few days. “ Powder-down” feathers are those which never develop beyond the early stage, and continually cusintegrate at the tip into bluish- or greyish-white powder ; they occur in the Zinamidae, Ardeidae, Rhinochetidae, EBurypygidae, Mesitidae, Accipitres and Psittaci, in Podargus, Coracias, Lepto- soma, Gymnoderus and Artamus.
Colour.—The colour of Feathers is due to one of three causes. First, an actual pigment ' may be present in certain corpuscles, or in diffused solution, and the tint does not then vary according to the incidence of the hght. Secondly, it may arise from a pigment overlaid by colourless structures in the form of ridges or imbedded polygonal bodies; here, if the vanes are scraped or held up to the light, the pigmentary colour alone is visible? Thirdly, the colour may be iridescent or prismatic; that is, a blackish pig-
1 Of this nature are zoomelanin (black), zoonerythrin (red), zooxanthin (yellow), turacin (red—only known in the Musophagidae), and perhaps turacoverdin (green, from the same family). Brown is produced by a combination of red and black ; white is the appearance due to innumerable air-spaces.
* Such are many yellows, oranges, greens and blues.
4 MOULT CHAP.
ment may lie beneath a surface, which, whether polished, ridged, or pitted, acts as a series of prisms, causing the hue to vary according to the relative position of the spectator’s eye and the light. This is seen in a remarkable degree in Humming-birds.' Not uncommonly the vanes of feathers have an appearance like watered silk, due to very indistinct transverse striations. In regard to plumage generally, it may be noticed that the markings on a feather frequently indicate the age of a bird. In some the immature plumage is characterised by light-coloured tips to the feathers, which are lost as maturity is reached. _ In other groups, and especially in most of the Accipitres or Diurnal Birds of Prey, the markings of the immature bird are generally longitudinal, and in the adult transverse. In nearly all these cases the change is effected at the first moult. Females and young are usually duller than males, but in some cases, such as Phalaropus (Limicolae) and Eclectus (Psittaci), the hen-birds are the more brightly coloured. Moult.—Referring to p. 2, it should be remarked that, after the production of a feather, the formative substances become for a while dormant, but awake to renewed activity, if accidental or periodical loss needs to be made good; and so we naturally arrive at the phenomena of the annual Moult, which is often “double,” that is, occurring towards autumn, and again in spring. Though some Birds do not lose their quill-feathers the first year, they normally gain a winter plumage—differing in colour from the summer garb—by moulting or shedding their feathers. The wing-quills, and even those of the tail, are ordinarily discarded in pairs, though not quite simultaneously ; but most Anatidae (Swans, Geese and Ducks), and apparently the Phoenicopteridae (Flamingos), lose all the former at once,” and with them the power of flight; while in the first-named Famjly the males of many species assume for several weeks a dress resembling that of the female, and are said to undergo an “eclipse.” Young birds moult, as a rule, somewhat later than adults, but in the typicai Gallinae the original quills are shed before the possessors are fully grown, and are succeeded by others of proportionately in- creased size, the power of flight being attained very early.
1 Albinism is due to the absence of pigment; melanism, xanthochroism and erythrism are terms implying an abnormal proportion of black, yellow, or red in the plumage. They may be caused by food.
* Tn some cases at least Rails and Water-hens do the same.
1 SKELETON 5
The additional or spring moult affects the smaller feathers only, while it is still doubtful how far changes of colour are due to a mere dropping off of the fringe of barbicels. The decorative plumes of the males of many species are gained at the vernal moult. The double process is certainly not diagnostic of Families or even Genera, except in isolated cases; as an instance, however, the Larks have one moult, the Pipits and Wagtails two.
In such cases as Swallows and Diurnal Birds of Prey generally, the plumage is not changed till after the migration; in the Ptarmigan there is a triple moult, the breeding-swit being changed first to a greyish habit and then to a white; in Penguins the feathers of the wing come off in flakes.!
Skeleton, Digestive Organs, etc.—The plumage, however, though often striking, and of undoubted utility as a non-conductor of heat and a protection against wet, plays a subordinate part in determining the relationships of the larger groups of Birds. — For this we need the assistance of anatomy, if indeed we do not rely upon it almost entirely. It will be well before starting to state that structures which are morphologically similar, that is, which have a like origin in the embryo, are termed “homologous,” while those which perform the same physiological functions are “ analo- gous,” the word in its strictest sense implying initial diversity.
Any standard work on Vertebrate Anatomy ought to furnish & concise account of the bony framework or Skeleton of a Bird, but it will be convenient here to follow mainly the treatment of Dr. Gadow, in Prof. Newton’s Dictionary of Birds, pp. 848-8677.
According to this authority the Axial Skeleton consists of the Skull and Vertebral Column; the Appendicular Skeleton of the Ribs, the Sternum, the Limbs and their Arches, the Hyoid Apparatus or framework of the tongue, and the Jaws.
1. The Vertebral Column, which protects the Spinal Cord, is composed of a variable number of cervical, dorsal, sacral or. pelvic, and caudal vertebrae; that is, those of the neck, back, loins and tail respectively. The first cervical vertebra, which bears the head, articulating with it by a single condyle, is called the Atlas ; the second, on which it turns, the Axis; the succeeding cervicals
1 In certain of the Tetraonidae the claws are shed in spring ; in some A/eidae (Auks) the horny bill-sheath and the outgrowths over the eyes are lost after the breeding season ; the American White Pelican moults a horny projection on the culmen after nesting, while the beak of Redpolls is much elongated in summer.
6 SKELETON CHAP.
present a considerable number of processes or projections, which protect certain blood- vessels, and serve for the attachment of the muscles which turn the flexible neck.. The dorsal vertebrae follow, and some not unfrequently coalesce with each other, but this is always so with the sacrals, and in nearly all existing Birds with the terminal portion of the caudals, which are fused together to, form a“ pygostyle ” or upright triangular plate to carry the tail-feathers.| Archaeopteryx, so far as is known, stands alone in having all the caudal vertebrae free.
A typical vertebra consists of a centrum, and an arch, with articular surfaces for two ribs, and is called heterocoelous when the facets, or connecting surfaces, are saddle-shaped, a condition charac- teristic of, and restricted to, Birds. It is amphicoelous, or biconcave, when each end is hollowed, as in the dorsal region of Ichthyornis and probably in Archaeopteryx ; procoelous, when concave in front Fia. 1.—Third cervical vertebra of Wood- (as is common in Reptiles) ; opis-
pecker (Picus viridis). (Viewed thocoelous when concave behind anteriorly.) 7%, vertebrarterial fora- ( as in many Mamm als).
men; Ob, upper arch; Pa, articular process ; Psi, haemal spine ; Pt, Pt, 2. The Ribs are doubly attached the two bars of the transverse process, he aeaiecasya Iyer livennal Gan shewn on one side ancylosed with the to the vertebrae bY & Neac (capi- cervical rib (#2); Sa, articular surface ¢ulum) and a knob (tuberculum) : of centrum. (From Wiedersheim.) - and have a neck, a dorsal, and a ventral portion, each dorsal section (save on the last rib) possessing an “ uncinate process” or thin, bony posterior projec- tion, except in Archaeopteryx and the Palamedeidae. Should the ventral piece articulate with the sternum the rib is “ true,” otherwise it is called “false”; moreover the cervical and frequently the post-thoracic ribs are fused with the cervical vertebrae and the ilia respectively.
3. The Breast-bone (Sfernuwm) presents two different styles— according to whether it exhibits on its ventral surface a median ridge or keel (carina), or not. In the former ease, which is that of by far the greater number of existing Birds (hence termed Carinatae), the keel is of variable size, being correlated with the power of flight. It is exceedingly deep in the Swifts, Humming
1 The Ratitae, Crypturi and Hesperornis have no pygostyle.
I SKELETON
NI
Birds, and certain Petrels, but dwindles almost to disappearance in some flightless forms such as the Dodo, the Kakapo (Stringops), the extinct New Zealand Goose (Cnemiornis), and a good many Rails. The absence of a keel is characteristic of the other and smaller group of Birds, made up of the Ostrich, Rhea, Emeu and
Fic. 2.—-Skeleton of the trunk of a Falcon. Oa, coracoid, which articulates with the sternum (St) at +; C7, keel of sternum; Fw (CZ), furcula (clavicles) ; G, glenoid cavity for humerus; S, scapula; Un, uncinate process; V, vertebral, and Sp, sternal, portion of rib. (From Wiedershein.)
Cassowary, Moa and Kiwi, which from the resemblance the
sternum thus bears to a flat-bottomed boat (7atis) are known as
Ratitae. Whether keeled or not, the breast-bone affords a
surface of attachment to the principal muscles of the fore-limbs,
and its anterior end supports the coracoids, as in Fig. 2. Various processes are in most cases developed on the sides of the sternum itself, behind its junction with the ribs, especially towards the
8 SKELETON CHAP.
posterior portion, where they often take the form of prolonga- tions, the extremities of which occasionally meet and enclose what are called fenestrae; but these are un- important when compared with the features pre- Sch sented by the anterior part. Rs 4. The Pectoral Arch, or s,s. Shoulder-Girdle, consists of = three pairs of bones, the : .. Coracoids, the Scapulae Bie ( — = Ray s% “sx or Shoulder-blades,
’
Dar
“os and the Clavicles ae. or -@ Collare
Lp bones, the last two usu- \ ally coalescing in the median line into a V-shaped or U-shaped Furcula (the well-known “Merry-thought”); but V....yyp ™M some groups, as certain Parrots, the clavicles are practically me absent, while in others,
Fig. 3.—Skeleton of the Limbs and Tail of a Carinate @S several Owls, they do
Bird. (The skeleton of the body is indicated by not unite. The fureula dotted lines.) #, digits; Fi, fibula; HIV, carpus; : MF, tarsometatarsus ; MH, carpometacarpus ; OA, often — ossifies firmly
humerus; OS, femur; Py, pygostyle; R, coracoid ; with the anterior por- Rd, ulna; Sch, scapula; St, sternum, with its keel 3
(Cr) ; 7, tibiotarsus ; Ul, radius; Z, Z}, digits of tion of the keel, and in
foot. (From Wiedersheim.) Fregata, Didus and the fatitae, the coracoids and scapulae are fused together.
5. The Anterior Limbs, or Wings, are composed of the Humerus, or upper arm-bone, the Ulna and Radius (mak- ing the fore-arm), the Carpus or wrist, the Metacarpus and Digits, corresponding with the hand and fingers. The first of the three metacarpals bears the Pollex, or thumb, with one or two
1 SKELETON 9
phalanges (joints); the second the Index, representing man’s first finger, with two or three joints; the third a weak digit with only one phalanx, except in Archaeopteryx, where there are four. The Casuarii and Apteryges possess an index only, which in the Sphenisee fuses with the pollex. The basal joint of this is the normal place of attachment of the “bastard wing” (alula spuria). Archaeopteryx had claws on all its fingers, but in recent Birds they occur on the first two only, being functionless in the adult. Wing-spurs arise from the carpal and metacarpal bones,
6. The