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THE

CAMBRIDGE NATURAL HISTORY

EDITED BY

F. HARMER, Sc.D., F.R.S., Fellow of King's College, Cambridge; Superintendent of the University Museum of Zoology

A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge ; University Lecturer on the Morphology of Invertebrates

VOLUME IX

BIRDS

By A^.'^H^^EvANs, M.A., Clare College, Cambridge

Hontion MACMILLAN AND CO., Limited

NEW YORK: THE MACMILLAN COMPANY 1899

A /I ri^lits rese>-ied

In sicco ludunt fulicae. Virgil. " Loons disport themselves on dry matters."

AUG 61958

ERRATA

Page 4, note 2, for Water-hens rr,ml Moor-hens.

10, line 19, after Owls read and Fandion. 16, 17, fo7- Lord Howe's read Lord Howe. 16, 22, for Galapagos read, Galapagos.

26, note 1, delete comma after Bronn's.

30, line 2 from bottom, for Tarapaca read Tarajjaca.

59, 6 ,, ,, for Pelecanoidinac read Pelecano'idinae.

60, lines 14, 26, 34, for Felecanoides read Pelecanoides. 67, line 6, for Tlialassaeca read Thalassoeca. 70, ,, 10, for Phaenicopteridae read Phoenicopteridae. 91, 12, for ralloides read ralloides.

118, ,, 17, for dominicus read dominica.

122, 2, after F. cristata, read the Tufted Duck.

133. 5, after Wavy, read or Snow Goose.

160, ,, 8 from bottom, for drrhattis read rirrafus.

215, ., Id, iov iwaelcUus vea,^ praclata.

258, 15, for perhaps read probably not.

351, ,, 11, and page 357, line 6, for Fhacnicojjhainae read Fhocnicoplminae.

357, 11 from bottom, for Fhaenicojihaes read Fhocnicopkaes.

429, Fig. 89, for jugiikwus Tea,d jiigularis.

550, line 20, for Seenopoeetes read Scenopocetes.

568, 9 from bottom, /or a scale-insect j^ead an Aphid.

5"^^^-^ If 6- 0 6^^

3 fV/^

PREFACE

In this volume of the " Cambridge Natural History " the author has attempted to meet a need which he believes to be some- what widely felt. Eecognisiug 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-

c

vi PREFACE

fication, Geographical Distribution, and Migration, and a " Ter- minology " of the subject.

In accordance with the scheme of the Series generally, tlie order followed runs from the lowest forms and the Eatite 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 siven, it does not follow that it is unl)roken 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 tlie like ; Myology ; Mechanism of Flight and the supposed Lines of Flight on Migration; the Classiiications of Linnaeus and the older writers ; and the Strickland Code of Ornithological Nomen- clature. For these Professor Newton's Dictionai'y of Birds, and especially the Introduction to it, may be consulted; l)esides a multitude of other w^orks.

The woodcuts have been chiefly supplied 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 Eoman in place of Italic type, nor for tlie dissociation of the vowels in the diphthongs ; in these minor points he personally differs from the writers of the former volumes, though he agree>s with the wish of his Editors for uniformity.

PREFACE vii

In conclusion, he must take the opportunity of acknow- ledging the invaluable assistance afforded hy Mr. Howard Saunders, who carefully went over the whole of the proofs, while Dr. E. B. Sharpo was kind enough to do the same ; nor must he fail to record his indebtedness to Professor Xewton, Air. Sclater, Dr. Gadow, Mr. Ogilvie Grant, and many others, not to mention the innumerable authors without whose previous lal)ours to write a book of this description would be a well- nigh impossible task. Dr. Stejneger's Volume on Birds in the Standard Xatural History should be mentioned in particular.

A. H. Evans. Cambridge, Novemher 17, 1898.

ADDENDUM

Since the text has been printed off, several new species have been described, and of these it is necessary to mention at least the following;

Archaeopteryx siemensi, from Solenhofeu, where the original form was obtained. (Dames.)

Fjuryapteryx exilis (Dinornithidae) ; a new genus, Anomalornis, is also proposed for Anomalopteryx (preoccupied). (Hutton.)

Ammoperclix cholmlcyi (Phasianidae), from Suakin. (Ogilvie- Grant.)

Cepphus snowi (Alcidae), from the Kurile Is. (Stejneger.) The range of C. columha will now be " Bering Sea to California ; " and of C. carlo " North-East Asia and Japanese Seas."

Fodoces 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 Yis has described the eggs and young of Salvador in a ( Anatidae).

In all these cases the Zoological Record for 1897 may be consulted.

CONTENTS

PllEFACF. . V

Addendum ............ viii

Scheme of the Classification adopted in this Book . . . . xi

CHAPTER I

Introduction . 1

CHAPTER II

Archaeornithes Neornithes Ratitae Neornithes Odontolcae . . 2-3

CHAPTER in

neornithes carinatae Brigade I Legion I (Colymbomorphae). Orders : Ichthyornithes

COLYMBIFORMES SPHENISCIFORMES PrOCELLARIIFORMES ... 48

CHAPTER IV

NEORNITHES CARINATAE CONTIXUED

Brigade I Legion II (Pelargomorphae). Orders: Ciconiiformes

Anseriformes Falconiformes ........ 70

CHAPTER V

neornithes carinatae continued

Brig.\de II Legion I (Alectoromorphae). Orders : Tin,\miforme.s

Galliformes^Gruiformes Charadriiformes ..... 182

CONTENTS

CHAPTER VI

NEORNITHES {'AllINATAE CONTINUED

PAGE

Brigade II Legion II (Coraciomorphae). Orders : CrrrLiFORMES

CoRACIirORMES ........... 351

CHAPTER YII

NEORNITHES CARINATAE CONCLUDED

Brigade II Legion II (Coraciomorphae concluded). Order : Passeri-

roRMES 466

Index . . . 589

r:TK

\

'is- F

SCHEME OF THE CLASSIFICATION ADOPTED m THIS BOOK

CLASS AVES (i.. 23.)

RATITAE (p. 26)

SUBCLASS I. ARCHAEORNITHES (p. 23). A rchaeojJtcryx (pp. viii, 23).

Sub-Class II. Neornithes (p. 2'.). Dirmox A. XEOllXITIIES llATITAE {i^. 25).

I. Struthioiies (p. 27) : Fani. Strutliionidae (p. 27). II. Rheae (p. 30) : Fam. Hheidae (ji. 30).

ITT Mpo-istanes fn 32^- Z^""^" ^- ^asuariidae (p. 33).

III. Megistanes Qi. 62.) . ^ j,^^^ jj Dromaeidae (p. 36).

IV. Apteryges (p. 38) : Fam. Apterygidae (p. 38). V. Diuoniitlies (p. 41) : Fain. Dinornithidae (]>. 41).

VI. Aepyornithes (p. 43) : Fam. Aeiiyoniithidae (p. 43).

1 ? STEREORNITHES

(p. 43)

(Mesemhriornis, etc. (p. 44).

Dicdnjw.a (p. 45). - Dasornis (p. 45).

Jiemioruis (p. 45). \Gastornis {[>. 45).

iJivisjoxB. KEORXITHES ODOXTOLCAE {i<. 45).

HESPERORNITHES (p. 46). ENALIORNITHES (p. 46). i. liiijjioniis (p. 46).

ICHTHYORNITHES

(p. 48) Order.

COLYMBIFORMES

;p. 49)

SPHENISCIFORMES

Divmox a. XEORXITHES CAEIXA TAE (i). iS).

j Fam. IcnTiiYORNiTiiin.\E (p. 48) : Ichthyornis (p. 48). (^ ? A-patornis (p. 49).

Sub-Order. Family. Sub-family.

CColymbit'p. 49) Colymbidaj: (p. 50)

Podicipedes

vl'- J'')

Sphenisci

(p. 54)

PROCELLARIIFORMES Tubinares

(p. 59)

(p. 59)

PuDICIPKDIDAE

(p. 52),

Sl'HKNI.'^CIDAE

(p. 54).

I Diomedeinae (p. 63). Pr.ocELLARiiDAE | Oceanitiiiae (p. 65). 9) 1 Procellaviinae (p. 65).

(^ Pelecanoidiuae (p. 68).

(p.

SCHEME OF CLASSIFICATION

CICONIIFORMES

(p. 70,

ANSERIFORMES

(p. lOS)

FALCONIFORMES

TINAMIFORMES

(p. 182)

GALLIFORMES

(p. 186)

75).

Family. Sub-family.

f Phaethontidae (p. 72).

SULIDAE (p. 73).

- Phalacrocoracidae (p. Fregatidae (p. 81).

1^ PELI'XANIDAE(p. 83).

J Ardbidae (p. 87).

\ SCOPIDAE (p. 95). ( CiCONIIDAE (p. 95).

1 IriDiDAE (n c)9^ f I^^idinae (p. 100). [ ^i^iD'DAC ip. yyj -^ Plataleinae (j). 103). I Phoesicopteriuae J (p. 105). 1 Palaelodidae l_ (pp. 105, 108). Palamedkidae (p. 108)

' Merginae (p. 115). Merganettiiiae (p. 116). Erismaturinae (p. 117). Fuligulinae (p. 118). Aiiatinae (p. 123). Clienonettiiiae (p. 130). Anseriiiae (p. 131). Cereopsinae (p. 133). Plectropterinae (p. 133). Anseranatiiiae (p. 135). - Cygniuae (p. 135). Cathartidae

(p. 137). Seri'entariidae

(p. 141). Vri/ruRiriAE (p. 143).

Sub-Order.

Steganopodes

(p. 70)

Ardeae (p. 86) Ciconiae (p. 95)

Phoenicoptcri

(p. 105)

Palamedeae

(p. 108).

Anseres (p. 110) An'atidae (p. Ill)

r Cathartae

(p. 137)

Accipitres

(pp. 137, 141)

Tinami (p. 182) Mesitae (p. 186) Turnices (p. 187)

Gain (p. 190)

Opisthocomi

(p. --^Jl)

Falcoxidae

(p. 146)

PAXniOXIDAE

. (p. ISO). Tinami DAE (Crypturidae) (p. 182).

MESITIDAE(p. 186). j'TuRNICIDAE

I (p. 187). i Pedionomidae I (p. 189). f Meoapodiidae (p. 190).

Gypaetinae (ji. 150). Polyboriuae (p. 151). Accipitrinae (]>. 153). Aquilinae (p. 159). Buteoninae (p. 164). Falcoiiinae (p. 173).

Cracidae (p. 194)

Phasiaxidaio

(p. 198)

Opisthocomidae (I'. 241).

rCraciiiae (p. 196). PeiiL'lopinae (p. 197). (^ Oreopliasiuae (p. 198).

f Numidiiiae (p. 204). Meleagrinae (p. 206). ~l Phasianinae (p. 206). I Odontophorinae (p. 230). iTetraoniiiae (p. 233). ,

SCHEME OF CLASSIFICATION

xni

Sub-Order.

GRUIFORMES

(!i. -243)

Family. Ralliuak (p. 243). Gruidae (p. 251).

AUAMIDAE(p. 25(5). PSOPHIIDAE (p. 257).

Caiiiamidae(i>. 258). Otididae (p. 260).

liHINOCHETIPAE

(p. 263).

EniYPYGIDAE

(p. 265). Hkliormithidae

(p. 267).

Limicolae (n. 268)

CHARADRIIFORMES

(p. 26S)

Charadriidae

(p. 272)

Ghionididae

(p. 292). Glareolidae

(p. 293) Thinocory-

THIDAE (p. 296). OEniCNEMIDAE

(p. 297).

.PARRIDAE(p. 297)

Lari (pp. 268, 300) Laridae (p. 300)

Alcae (p. 315) Pterocles (p. 321)

Columbae (p. 325)

Alcidae (p. 315) Pterocliuae (].. 321).

' UlDIDAE (p. 328). DiDUNCULIDAE

(p. 331).

COLUMBIDAE

(p. 333)

{ Charadriiiiae (p. 272). - Triiigiiiae (p. 278). 1^ Scolopacinae (p. 289).

( Glareoliiiae (p. 293). \ Drouiadiiiao ([i. 296).

Stercoraiiinae (p. 304). Larinae (p. 305). Ilhyneliopmae (]<. 310). Sterninae (p. 310).

Goiirinae (p. 334). Peristerinae (p. 334). Colunibinae ([>. 342). . Treroniuae (p. 344).

CUCULIFORMES

(p. 351)

CORACIIFORMES

(p. 376)

Cuculi (p. 351)

Psittaci (]'. 361;

Coraciae (p. 376)

{Cunlhuicd on the next ikkjc.)

CrCULlDAE

(p. 351)

MlTSOPHAGIDAE

(p. 359),

PSITTACIDAK

(p. 366)

Trichoglossidae (p. 373)

CORACIIDAK

(p. 376)

MoMOTIDAE

(p. 379)

Cuculinae (p. 352). Centropodiiiae (p. 356). Phaeuicophainae (p. 357). Neomorphinae (p. 357). Diplopteriiiae (p. 359). Crotophaginae (p. 359).

j' Striiigopinae (p. 366). - Psittaciiiae (p. 367). ( Cacatuinae (p. 372). I Cyclopsittaciiiae (p. 373)- I Loriinae (p. 373). I_ Nestoriiiau (p. 374).

f Coraciinae (p. 376). \ Lepto.somatinae (p. 378). ] ]\Iomotniae (p. 380). \Todmae (p. 381).

SCHEME OF CLASSIFICATION

CORACIIFORMES

{continued)

Sub-Order.

Coraciae

{continued)

Striges (p. 397)

Caprimulgi

(p. 415)

Sub-family, f Halcvoniiiae (p. 385). \ Alcedininae (p. 386).

Family. A LC ED IN I DAE (p. 382)

Mekopidae

(p. 387).

BUCEROTIDAE

(p. 390).

Ul'UPIDAE

^ (p. 395) f Stkiuidae

.. (p. 398) ___ _,. ,.

CA?iiiMri,iiiDAE j ( Capriniulginae (p. 418).

f Upupinae (p. 395). ( Irrisorinae (p. 397). / Striginae (p. 403), -jjj Buboninae (p. 404).

Cypseli ^p. 419)

Colii (p. 439) Trogones (p. 441)

Pici (p. 445)

I Trochi I (P- '

(p. 417)

PODARGIDAE

(p. 419). Steatoiinithi-

DAE (p. 419).

Cypsemdae (p. 420)

ilLIDAE

426).

COLIIDAE

(p. 439). Trogonidae

(p. 441). Galbulidae

(p. 445) Capitonidae

(p. 448) Khamphastidae

(p. 453)

PiCIDAE (p. 457)

\ Nyctibiiuae (p. 418).

i' Macropteryginae (p. 422). ^ Cliaeturinae (p. 422). \ Cyijselinae (p. 424).

/ Galbulinae (p. 445). \ BucL'ouinae (p. 446).

/ Caintoninae (p. 448). \ Indicatoi'iiiae (p. 451)

f Picinae (p. 457). [ lynginae (p. 464).

Order.

Group.

PASSERI FORMES

(p. 466)

Passeres aniso- myodae

(p. 467)

Division. SUBCLAMATORES (p. 467)

Clamatores (p. 469)

{Contimied on the next page.)

Family. EURYLAEMIDAE

(p. 467).

|- PiTTIDAE

(p. 469). Philepittidae

(p. 471).

Xenicidae

(p. 472).

Tyrannidae

(p. 473)

^ OXY'RHAMPHIDAE

(p. 477).

PiPRIDAE

(p. 477).

COTINGIDAE

(p. 479)

Phytotomidae

(p. 483).

Sub-family.

Taenioptei'inae ^ Platyrhynchinae |^ p.

\ Elaiiicinae I^Tyraniiinae

Titj'rinae

Lipaiiginae

Attilinae

Rupifolinae

Cotiiiginae

Gymnoderiuae

1473.

(P- 479).

SCHEME OF CLASSIFICATION

Order.

PASSERI- FORMES

[cuidiiLucd)

Gruup.

Passeres aniso myodac

{continued}

Clamatouks {cuntinucd)

Passeres diacro- myodae

(p. 491)

SUBOSCINES

(p. 491)

Continued o n the next iJa/je. )

OSCINES

(p. 494)

Siib-fainily. Furnariiiiae

J

j Formicariinae r488). I Grallariiiiae I

f Motacillinae

iA:uthi;;;ri(p-498).

Family.

r I'urnaninae ^ DKSl,r.c.roi,.SPTI- SynalUsii.ae ,

'■"■ (•'■ ^«»' Dtd"s;,. ' ^^^■>-

I tinae [ Thamnophili- FiuiMU'AUiiDAE I nae

ip. 4SS)

CONOPOPHAGIDAE (p. 489).

Pteroptochidae

(p. 490).

j Menfridae J (P- 491). j Atuichornithi-

l DAE (p. 493).

Alaudidae

(p. 496). Motacillidae

(p. 498) Henicukidae

(p. 601). Timeliidae

(p. 501). Pycnonotidae

(p. 504).

MUSCICAPIDAE

(p. 506).

TrUDIDAE

(p. 509)

ClXCLIDAE

(p. 519).

TUOGLODYTIDAE

(p. 521). Chamaeidae (p. 522).

HiRUNDIXIDAE

(p. 522). Campephagidae

(p. 525). Dicruridae

(p. 527). Ampelidae

(p. 529). Artamidae

(p. 530).

Turdinae (p. 509). Myiodectinae (p. 513) Sylviinae (p. 513). Polioptilinae (p. 514).

, Miminae (p. 514).

Laniidae (p. 531)

Gyiniiorliiiiiuae

(p. 532). Malacoiio tinae

(p. 533). Pachycephalinae

(p 533). Laniiiiae (p. 534). Prioiiopiuae (p. 535).

SCHEME OF CLASSIFICATION

Group.

Division.

PASSERI- FORMES

{co nil lined)

Passeres diacro- myodae

{cviiiinucd)

OSCINES

{continued)

Family. VlUEOXIDAE

(p. 536).

SlTTIDAE

(p. 536). Paridae

(p. 538). Panuridae

(p. 541). Orioi.idae

(p. 542). Paradiseidae

(p. 543).

Corvidae (p. 552)

Sturnidae

(p. 559). Drepanididae

(p. 562). Meliphagihae

(p. 564) Zosteropidae

(p. 568). Necta rixiid.ae

(p. 568).

DiCAEIDAE

(p. 570). Certhiidae coerebidae

(p. 572).

ilXKITILTIDAE

(p. 573. Tanagridae

(p. 575). Ploceidae

(p. 576)

Sub-faniilv.

[ Corvinae - Garruliuac \ Fi'egiliuae

(p. 552).

f Myzomelinae \ (p. [ Meliphagiuae j 564).

(p. 571 ;

f Viduinae (p. 576). \ Ploceinae ([>. 577).

Icteridae

(p. 579)

Fringillidae \ ,. + Emberizidae \ 582).

Cassiciiiae

Agelaeinae

Stunielliuae

Icterinae

Quiscaliiiae

P-

I (P- 9)

f

CHAPTER I

INTRODUCTION

Definition. "A Bird is a featlieivd 1)iped." This popiilar 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 supertiuous.

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, anniiotic 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 Ijeginner might well postpone, his attention being solely drawn to the external characters ; though of course those that are internal are by no means to l)e 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 l^iirds 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 Heptilia. Birds are, in fact, only extremely modified Reptiles, the tw^o Classes forming the Saur- 02)sida of Huxley, one of his three primary divisions of Vertebrata.

1 P.Z.S. 1892, p. 236. VOL. IX B

FEATHERS

The aid of the Palaeontologist and Geologist must thus be called in to clear up many problems which present themselves to the Ornithologist wlio does not content himself with examining exist- ing forms of life alone. Arehacoptcri/x (p. 23) from the Jurassic System is the oldest Bird known, nor are any other pre-Tertiary forms recorded, save a small nvunber from the rocks of the Creta- ceous Epoch, the chief of which are the so-called Odontornithes, or toothed species of America (p. 45).

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 IMammals, is erroneous ; for, almost without exception, they grow in certain definite tracts called lAerylae, 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 consideral^le 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 Xewton'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 supplies 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 " welis " or " vanes " (vexilla) often called contour feathers {pennae or plumae), a fresh 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

^ 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 3

the barrel or quill (calamus (jr sc(/pus) at the base of the tuft of rays have elongated into a principal shaft (rhachis) ; this is generally accompanied by a secondary " aftershaft " {hyjiorliaehu), originating from the " ventral " side, whicli in the Emeu and Cassowary rivals the shaft itself in size. On the rhachis a doulJe 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 {cilia), which in the distal rows usually terminate in booklets {hamuli). 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 disintegrate at the tip into bluish- or greyish-white powder ; they occur in the Tinamidae, Ardeidae, IlMnoclietidae, Eurypygidae, Mesitidae, Accipitres and Psittaci, in Podargus, Coracias, Lepto- soma, Gym,noderus 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 light. 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-

^ Of this nature are zoomelanin (black), zoonerytlirin (red), zooxantliin (yellow), turacin (red— only known in the Mtcsoijhncjiduc), and perhaps turacoverdin (green, from the same family). Brown is jDroduced 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 unconnnonly 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 innnature 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 inmiature 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 {TAmicolae) and Eclectus {Pslttaci), 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 Phocnico2'>teridae (Flamingos), lose all the former at once,'' and with them the power of flight ; while in the first-named Family 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 typical 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.

^ Albinism is due to the absence of pigment ; melanism, xantliocliroism and erythrism are terms implying an abnormal proportion of black, yellow, or red in the plumage. They may be caused bj' food.

~ In some cases at least Rails and Water-hens do the same.

I SKELETON 5

The additional or spring moult affects the smaller feathers only, while it is still doul)tfnl how far changes of colour are due to a mere dropping off of the fringe of bai'bicels. 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, tlie breeding-suit 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 w"et, plays a suljordinate part in determining the relationships of the larger groups of Birds. For tliis we need the assistance of anatomy, if indeed we do not rely upon it almost entirely. It will Ije 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 Yerteln-ate Anatomy ought to furnish a 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 Xewton's Dictionary of Birds, pp. 848-867.

According to this authority the Axial Skeleton consists of the Skull and Vertebral Column ; the Appendicular Skeleton of the Eibs, the Sternum, tlie Limbs and their Arches, the Hyoid Apjjaratus 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, 1 )ack, 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 Tctrctonidae the claws are shed in spring; in some A Icidae (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.

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r-7&

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 vmfrequently 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 Arcliaeoj)teryx ; procoelous, when concave in front (as is common in Eeptiles) ; opis- thocoelous when concave beliind (as in many Mammals).

2. The Eibs are doubly attached to the vertebrae by a head [ca/pi- t ilium) and a knob (tuherculum) ; and have a neck, a dorsal, and a ventral portion, each dorsal section (save on the last rili) possessing an " uncinate process " or thin, bony posterior projec- tion, except in Archaeopteryx and the Falaviedeidae. Should the ventral piece articulate with the sternum the rib is " true," otherwise it is called " false " ; moreover the cervical frequently the post -thoracic ribs are fused with the vertebrae and the ilia respectively.

3. The Breast-bone (Sternum) presents two different styles according to whether it exhibits on its ventral surface a median ridge or keel (carina), or not. In the former case, 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

^ The Ratitae, Crypturt and I£es2)crornis have no pygostyle.

Fig. 1. Third cervical vertebra ofWood- pecker {Picus viridis). (Viewed anteriorly.) Ft, vertebrarterial fora- men ; Ob, upper arch ; Pa, articular process ; Psi, haemal spine ; Pt, Pt, the two bars of the transverse process, shewn on one side aucylosed with the cervical rib [R) ; Sa, articular surface of centrum. (From Wiedersheim.)

ind cervical

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Birds, and certain Petrels, but dwindles almost to disappearance in some flightless forms such as the Dodo, the Kakapo {^String irps), the extinct New Zealand Goose {Cnemiornis), and a good many liails. The absence of a keel is characteristic of the other and smaller group of Birds, made up of the Ostrich, Ifhea, Emeu and

Fig. 2. Skeleton of the trunk of a Falcon. Ca, coracoiil, which articulates with the sterimm {St) at t ; t.'', keel of .steruuiu ; Fit {CI), furcula (clavicles) ; (!, glenoid cavity for humerus ; ^, scapula; t'/(, uncinate process ; T', vertebral, and ^^^a stei-nal, portion of rib. (lYoin Wiedersheim.)

Cassowary, Moa and Kiwi, which from the resemblance the sternum tlius bears to a flat-bottomed boat gratis) are known as Jlatltae. 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

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posterior portion, where they often take the form of prolonga- tions, the extremities of which occasionally meet and enclose what are called fcnestrae ; but these are un- important when compared with the features pre- sented hy the anterior part.

4. The Pectoral Arch, or Shoulder -Girdle, consists of •:-. three pairs of bones, the :%'•■::., Coracoids, the Scapulae VvX. ■••;■>. or Shoulder-blades, ■•■v^ and the Clavicles or Collar-

bones, the last two usu- ally coalescino; in the median line into a V-shaped or U-shaped Furcula (the well-known " Merry-thought ") ; but in some groups, as certain Parrots, the clavicles are practically absent, wdiile in others.

Fig. 3.— Skeleton of tlie Limhs and Tail of n, Cariiiate '^^ Several Owls, tllCy do Bird. (The skeleton of the body is indicated by ^lOt Unite. The furcula dotted lines.) /; digits ; Fi, fibula; HW, carpus; " .

.l//', tarsometatarsus ; J/S", carpometacarpus ; 0^, 01 ten OSSlfieS firmly Immerus; femur ; Pypygostyle ; if .coracoid ; ^^ith the anterior por- R<l, ulna ; Sch, scapula ; St, sternum, with its keel . ^

(Cr) ; T, tibiotarsus ; Ul, radius; Z, Z^, digits of tion of the keel, and in

foot. (From Wiedersheim. ) Frcgata, Bidus and the

Ratitae, 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 Eadius (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 thuml), with one or two

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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 Archacoptrryx, where there are four. The Casuarii and Apteryges possess an index only, which in the Sploenisci fuses with the pollex. The basal joint of tliis is the normal place of attachment of the " bastard wing " {alula sjj'uria). ArcJiaeopteryx had claws on all its fingers, but in recent Birds they occur on the first two fmly, being functionless in the adult. Wing-spurs arise from the carpal and metacarpal bones.

6. The Pelvic Arch consists of the Ilium, Ischium, and Os ])ubis, these three paired bones meeting from each side at the cup {((cctcdnduin) that receives tlie head of the femur, and coalescing early in life ; while the incisura ischiadtca or notch between the

Fig. 4. Pelvis of Apteryx mistralis. Lateral view, a. Acetabulum ; {7, ilium : is, isclainm : p, pectineal process of pubis ; ^^ jiubis. (From Wiedersheim, after Marsli.)

ischium and the ilium becomes an inclosed space (jorffmen) in all Birds except the Batitae and Crypturi.

V. The Posterior Limbs, or Legs, are composed of the Femur or thigh, the Tibia and Pibula, making the shank or " drum- stick," and the bones of the Foot. The thigh, however, being hidden l)y the plumage, the sliank of a Bird might easily be taken for the thigh, and the metatarsus (the cannon-l)one of some) for the shank. The tibia and fibula commonly unite to some extent, and the former, as it now exists in adult Birds, is strictly a " tibio-tarsus," since with it is fused the proximal portion of the originally existing tarsal elements. Similarly the distal tarsal

I o SKELETON

elements unite with the metatarsus, which is therefore properly a " tarso-metatarsus," though often called merely "tarsus" by ornithologists. This arises from a fusion of the second, third, and fourth metatarsal bones, which in the adult (except among the Splicnisci and to some extent in Fsittaci) do not lie in the same plane ; the middle one having its upper end thrust back- ward and its lower end forward in the course of growth to maturity. The fifth metatarsal practically disappears, while the first remains more or less separate, and lies behind the distal portion of the other metatarsals.

Of the toes the fifth is not traceable in Birds ; the first is often aborted, but the second only in Struthio, and to a less extent in Cei/x and Alcyone, and the fourth (nearly) in Cholornis. The hallux, or hind toe, has two phalanges, the second digit three, the tlnrd four, and the fourth five ; Cypselus and Pamjptila (Swifts), however, are exceptions, and possess only three in each of the anterior toes, while the Ccqwimidginae (true Nightjars) and Fteroclidae (Sand-Grouse) have only four joints on the outer. In OwlS/^the fourth digit is reversible at will, the same being true to a less extent of the Musophagidae (Plantain-eaters) and Leptosoma (akin to the Eoller) ; when this condition is permanent, as in the Cuculidae, Fsittaci and Fid the foot is termed zygodactylous. In Trogones the second toe is reversed (heterodactylous). Coliiis can turn the first toe forward and the fourth backward, while certain Swifts, and to a less degree some Nightjars, have the whole number permanently pointing to the front (pamprodactylous). Membranes more or less connecting the anterior digits produce a webbed or swimming foot, even the hallux being united with the rest in the Steganopodes. The hind-toe is often elevated, or higher than its fellows, when it is conniionly reduced and some- times lacks a nail. The Ostrich has little or no claw on the outer toe, while that of the third toe is toothed or serrated in a considerable number of Birds, but this is a character of very slight importance.

The covering of the metatarsus is usually " scutellated," but when the scutellae, or scales, which may be oblong or polygonal, are smaller than usual and generally hexagonal it is called " reticulated." In some cases the surface becomes nearly or quite smooth (" ocreated " or " booted "), or more or less granulated.

8. The structure of the Skull is a study in itself and affords

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I I

considerable help in Taxonomy (Classification). It must suffice here to refer for the names of the parts to the suljjoined figure.

The Bill, or Beak, is composed of an upper jaw or maxilla, and an under jaw or mandiljle. From the figure it will Ije seen that "maxilla" is not strictly the whole upper portion, thougli the term is thus used for convenience, as is the plural "mandibles" for the two jaws when mentioned simultaneously. The " rham- photlieca," or horny sheath, may be simple (undivided), or com- pound, that is, made of several distinct pieces. In tlie Ansercs the covering is soft with a horny (corneous) tip or " nail " ; in

em ^"'n f

t.-p

ar

Fig. 5. Skull of a Wild Duck {Anas hoscas\ from the side. o,g. Angular; aU, alispheuoid ; «r, articular ; ht, basitemporal ; d, deutary ; en, external nostrils ; c.o, exoccipital ; eth, ethmoid ; fr, frontal ; j, jugal ; Ic, lacrymal ; mx, maxilla ; mx.p, niaxillopalatine process ; n, nasal ; f, parietal ; 'p(j, pterygoid ; jjl, palatine ; ps, presphenoid ; ^j.x-, premaxilla ; q, quadrate ; q.j, quadratojugal ; s.ojj, supra-angular ; s.n, supraoccipital ; sq, squamosal ; Ui, tympanic cavity ; i\ vomer ; //, foramen for optic nerve ; I', for trigeminal. (From Wiedersheim, after Parker.)

the Liiiiicolac it varies extremely, producing a hard pickaxe, as in