NEORNITHES CARINATAE _CONCLUDED_
BRIGADE II–LEGION II (CORACIOMORPHAE _CONCLUDED_). ORDER:
PASSERIFORMES 466
INDEX 589
{xi}SCHEME OF THE CLASSIFICATION ADOPTED IN THIS BOOK
CLASS AVES (p. 23.)
SUB-CLASS I. ARCHAEORNITHES (p. 23).
_Archaeopteryx_ (pp. viii, 23).
SUB-CLASS II. NEORNITHES (p. 25).
_DIVISION A. NEORNITHES RATITAE_ (p. 25).
RATITAE (p. 26) { I. Struthiones (p. 27): Fam. Struthionidae (p. 27).
{ II. Rheae (p. 30): Fam. Rheidae (p. 30).
{ III. Megistanes (p. 32): { Fam. I. Casuariidae (p. 33).
{ { Fam. II. Dromaeidae (p. 36).
{ IV. Apteryges (p. 38): Fam. Apterygidae (p. 38).
{ V. Dinornithes (p. 41): Fam. Dinornithidae (p. 41).
{ VI. Aepyornithes (p. 43): Fam. Aepyornithidae (p. 43).
?? STEREORNITHES { { _Mesembriornis_, etc. (p. 44).
(p. 43) { { _Diatryma_ (p. 45).
{ ? { _Dasornis_ (p. 45).
{ { _Remiornis_ (p. 45).
{ { _Gastornis_ (p. 45).
_DIVISION B. NEORNITHES ODONTOLCAE_ (p. 45).
HESPERORNITHES (p. 46).
? ENALIORNITHES (p. 46).
? _Baptornis_ (p. 46).
_DIVISION C. NEORNITHES CARINATAE_ (p. 48).
ICHTHYORNITHES { Fam. ICHTHYORNITHIDAE (p. 48): _Ichthyornis_ (p. 48).
(p. 48) { ? _Apatornis_ (p. 49).
Order. Sub-Order. Family. Sub-family.
COLYMBIFORMES { COLYMBI (p. 49) COLYMBIDAE (p. 50).
(p. 49) { PODICIPEDES PODICIPEDIDAE (p. 52).
{ (p. 49)
SPHENISCIFORMES SPHENISCI SPHENISCIDAE
(p. 54) (p. 54) (p. 54).
PROCELLARIIFORMES
(p. 59) TUBINARES PROCELLARIIDAE { Diomedeinae (p. 63).
(p. 59) (p. 59) { Oceanitinae (p. 65).
{ Procellariinae (p. 65).
{ Pelecanoïdinae (p. 68).
CICONIIFORMES{ STEGANOPODES { PHAËTHONTIDAE (p. 72).
(p. 70) { (p. 70) { SULIDAE (p. 73).
{ { PHALACROCORACIDAE (p. 75).
{ { FREGATIDAE (p. 81).
{ { PELECANIDAE (p. 83).
{
{ ARDEAE (p. 86) { ARDEIDAE (p. 87).
{ { SCOPIDAE (p. 95).
{
{ CICONIAE { CICONIIDAE (p. 95).
{ (p. 95) { IBIDIDAE { Ibidinae (p. 100).
{ { (p. 99) { Plataleinae (p. 103).
{
{ PHOENICOPTERI { PHOENICOPTERIDAE (p. 105).
{ (p. 105) { PALAELODIDAE (pp. 105, 108).
ANSERIFORMES { PALAMEDEAE { PALAMEDEIDAE
(p. 108) { (p. 108). { (p. 108)
{
{ ANSERES { ANATIDAE { Merginae (p. 115).
{ (p. 110) { (p. 111) { Merganettinae (p. 116).
{ { { Erismaturinae (p. 117).
{ { { Fuligulinae (p. 118).
{ { { Anatinae (p. 123).
{ { { Chenonettinae (p. 130).
{ { { Anserinae (p. 131).
{ { { Cereopsinae (p. 133).
{ { { Plectropterinae(p. 133).
{ { { Anseranatinae (p. 135).
{ { { Cygninae (p. 135).
FALCONIFORMES{ CATHARTAE { CATHARTIDAE
(p. 137) { (p. 137) { (p. 137).
{
{ ACCIPITRES { SERPENTARIIDAE (p. 141).
{ (pp. 137, 141) { VULTURIDAE (p. 143).
{ { FALCONIDAE { Gypaëtinae (p. 150).
{ { (p. 146) { Polyborinae (p. 151).
{ { { Accipitrinae (p. 153).
{ { { Aquilinae (p. 159).
{ { { Buteoninae (p. 164).
{ { { Falconinae (p. 173).
{ { PANDIONIDAE (p. 180).
TINAMIFORMES { TINAMI (p. 182) { TINAMIDAE (CRYPTURIDAE)
(p. 182) { { (p. 182).
GALLIFORMES { MESITAE (p. 186){ MESITIDAE (p. 186).
(p. 186) {
{ TURNICES { TURNICIDAE (p. 187).
{ (p. 187) { PEDIONOMIDAE (p. 189).
{
{ GALLI (p. 190) { MEGAPODIIDAE
{ { (p. 190).
{ { CRACIDAE { Cracinae (p. 196).
{ { (p. 194) { Penelopinae (p. 197).
{ { { Oreophasinae (p. 198).
{ { PHASIANIDAE { Numidinae (p. 204).
{ { (p. 198) { Meleagrinae (p. 206).
{ { { Phasianinae (p. 206).
{ { { Odontophorinae (p. 230).
{ { { Tetraoninae (p. 233).
{
{ OPISTHOCOMI { OPISTHOCOMIDAE
{ (p. 241) { (p. 241).
GRUIFORMES { { RALLIDAE (p. 243).
(p. 243) { { GRUIDAE (p. 251).
{ { ARAMIDAE (p. 256).
{ { PSOPHIIDAE (p. 257).
{ { CARIAMIDAE (p. 258).
{ { OTIDIDAE (p. 260).
{ { RHINOCHETIDAE (p. 263).
{ { EURYPYGIDAE (p. 265).
{ { HELIORNITHIDAE (p. 267).
CHARADRIIFORMES
(p. 268) { LIMICOLAE { CHARADRIIDAE { Charadriinae (p. 272).
{ (p. 268) { (p. 272) { Tringinae (p. 278).
{ { { Scolopacinae (p. 289).
{ { CHIONIDIDAE (p. 292).
{ { GLAREOLIDAE { Glareolinae (p. 293).
{ { (p. 293) { Dromadinae (p. 296).
{ { THINOCORYTHIDAE (p. 296).
{ { OEDICNEMIDAE (p. 297).
{ { PARRIDAE (p. 297)
{
{ LARI { LARIDAE { Stercorariinae (p. 304).
{ (pp. 268, 300) { (p. 300) { Larinae (p. 305).
{ { { Rhynchopinae (p. 310).
{ { { Sterninae (p. 310).
{
{ ALCAE (p. 315) { ALCIDAE (p. 315).
{
{ PTEROCLES { PTEROCLIDAE
{ (p. 321) { (p. 321).
{
{ COLUMBAE { DIDIDAE (p. 328).
{ (p. 325) { DIDUNCULIDAE (p. 331).
{ { COLUMBIDAE { Gourinae (p. 334).
{ { (p. 333) { Peristerinae (p. 334).
{ { { Columbinae (p. 342).
{ { { Treroninae (p. 344).
CUCULIFORMES { CUCULI (p. 351) { CUCULIDAE { Cuculinae (p. 352).
(p. 351) { { (p. 351) { Centropodinae (p. 356).
{ { { Phaenicophainae(p. 357).
{ { { Neomorphinae (p. 357).
{ { { Diplopterinae (p. 359).
{ { { Crotophaginae (p. 359).
{ { MUSOPHAGIDAE (p. 359).
{
{ PSITTACI { PSITTACIDAE { Stringopinae (p. 366).
{ (p. 361) { (p. 366) { Psittacinae (p. 367).
{ { { Cacatuinae (p. 372).
{ {TRICHOGLOSSIDAE{ Cyclopsittacinae(p. 373)
{ { (p. 373) { Loriinae (p. 373).
{ { { Nestorinae (p. 374).
CORACIIFORMES{ CORACIAE { CORACIIDAE { Coraciinae (p. 376).
(p. 376) { (p. 376) { (p. 376) { Leptosomatinae (p. 378).
{ { MOMOTIDAE { Momotinae (p. 380).
{ { (p. 379) { Todinae (p. 381).
{ { ALCEDINIDAE { Halcyoninae (p. 385).
{ { (p. 382) { Alcedininae (p. 386).
{ { MEROPIDAE (p. 387).
{ { BUCEROTIDAE (p. 390).
{ { UPUPIDAE { Upupinae (p. 395).
{ { (p. 395) { Irrisorinae (p. 397).
{
{ STRIGES { STRIGIDAE { Striginae (p. 403).
{ (p. 397) { (p. 398) { Buboninae (p. 404).
{
{ CAPRIMULGI { CAPRIMULGIDAE { Caprimulginae (p. 418).
{ (p. 415) { (p. 417) { Nyctibiinae (p. 418).
{ { PODARGIDAE (p. 419).
{ { STEATORNITHIDAE (p. 419).
{
{ CYPSELI { CYPSELIDAE { Macropteryginae(p. 422).
{ (p. 419) { (p. 420) { Chaeturinae (p. 422).
{ { { Cypselinae (p. 424).
{ { TROCHILIDAE (p. 426).
{
{ COLII (p. 439) { COLIIDAE (p. 439).
{
{ TROGONES { TROGONIDAE
{ (p. 441) { (p. 441).
{
{ PICI (p. 445) { GALBULIDAE { Galbulinae (p. 445).
{ { (p. 445) { Bucconinae (p. 446).
{ { CAPITONIDAE { Capitoninae (p. 448).
{ { (p. 448) { Indicatorinae (p. 451).
{ { RHAMPHASTIDAE (p. 453)
{ { PICIDAE { Picinae (p. 457).
{ { (p. 457) { Iynginae (p. 464).
Order PASSERIFORMES (p. 466)
Group. Division. Family. Sub-family.
PASSERES { SUBCLAMATORES { EURYLAEMIDAE
ANISOMYODAE{ (p. 467) { (p. 467).
(p. 467) {
{ CLAMATORES { PITTIDAE
{ (p. 469) { (p. 469).
{ { PHILEPITTIDAE
{ { (p. 471).
{ { XENICIDAE
{ { (p. 472).
{ { TYRANNIDAE { Taeniopterinae }
{ { (p. 473) { Platyrhynchinae } p.
{ { { Elaineinae } 473
{ { { Tyranninae }
{ { OXYRHAMPHIDAE
{ { (p. 477).
{ { PIPRIDAE
{ { (p. 477).
{ { COTINGIDAE { Tityrinae }
{ { (p. 479) { Lipauginae }
{ { { Attilinae } (p.
{ { { Rupicolinae } 479).
{ { { Cotinginae }
{ { { Gymnoderinae }
{ { PHYTOTOMIDAE
{ { (p. 483).
{ { DENDROCO- { Furnariinae }
{ { LAPTIDAE { Synallaxinae } (p.
{ { (p. 483) { Sclerurinae } 484).
{ { { Dendrocolaptinae }
{ { FORMICARIIDAE { Thamnophilinae } (p.
{ { (p. 488) { Formicariinae } 488).
{ { { Grallariinae }
{ { CONOPOPHAGIDAE
{ { (p. 489).
{ { PTEROPTOCHIDAE
{ { (p. 490).
PASSERES { SUBOSCINES { MENURIDAE
DIACROMYODAE{ (p. 491) { (p. 491).
(p. 491) { { ATRICHORNITHIDAE
{ { (p. 493).
{
{ OSCINES { ALAUDIDAE
{ (p. 494) { (p. 496).
{ { MOTACILLIDAE { Motacillinae } (p. 498).
{ { (p. 498) { Anthinae }
{ { HENICURIDAE
{ { (p. 501).
{ { TIMELIIDAE
{ { (p. 501).
{ { PYCNONOTIDAE
{ { (p. 504).
{ { MUSCICAPIDAE
{ { (p. 506).
{ { TURDIDAE { Turdinae (p. 509).
{ { (p. 509) { Myiodectinae (p. 513).
{ { { Sylviinae (p. 513).
{ { { Polioptilinae (p. 514).
{ { { Miminae (p. 514).
{ { CINCLIDAE
{ { (p. 519).
{ { TROGLODYTIDAE
{ { (p. 521).
{ { CHAMAEIDAE
{ { (p. 522).
{ { HIRUNDINIDAE
{ { (p. 522).
{ { CAMPEPHAGIDAE
{ { (p. 525).
{ { DICRURIDAE
{ { (p. 527).
{ { AMPELIDAE
{ { (p. 529).
{ { ARTAMIDAE
{ { (p. 530).
{ { LANIIDAE { Gymnorhininae (p. 532).
{ { (p. 531) { Malaconotinae (p. 533).
{ { { Pachycephalinae
{ { { (p 533).
{ { { Laniinae (p. 534).
{ { { Prionopinae (p. 535).
{ { VIREONIDAE
{ { (p. 536).
{ { SITTIDAE
{ { (p. 536).
{ { PARIDAE
{ { (p. 538).
{ { PANURIDAE
{ { (p. 541).
{ { ORIOLIDAE
{ { (p. 542).
{ { PARADISEIDAE
{ { (p. 543).
{ { CORVIDAE { Corvinae }
{ { (p. 552) { Garrulinae } (p. 552).
{ { { Fregilinae }
{ { STURNIDAE
{ { (p. 559).
{ { DREPANIDIDAE
{ { (p. 562).
{ { MELIPHAGIDAE { Myzomelinae } (p.
{ { (p. 564) { Meliphaginae } 564).
{ { ZOSTEROPIDAE
{ { (p. 568).
{ { NECTARINIIDAE
{ { (p. 568).
{ { DICAEIDAE
{ { (p. 570).
{ { CERTHIIDAE (p. 571).
{ { COEREBIDAE
{ { (p. 572).
{ { MNIOTILTIDAE
{ { (p. 573.
{ { TANAGRIDAE
{ { (p. 575).
{ { PLOCEIDAE { Viduinae (p. 576).
{ { (p. 576) { Ploceinae (p. 577).
{ { ICTERIDAE { Cassicinae }
{ { (p. 579) { Agelaeinae } (p.
{ { { Sturnellinae } 579).
{ { { Icterinae }
{ { { Quiscalinae }
{ { FRINGILLIDAE } (p. 582).
{ { + EMBERIZIDAE }
ERRATA
Page 4, note 2, _for_ Water-hens _read_ Moor-hens.
" 10, line 19, after Owls _read_ and _Pandion_.
" 16, " 17, _for_ Lord Howe's _read_ Lord Howe.
" 16, " 22, _for_ Galapagos _read_ Galápagos.
" 26, note 1, delete comma after Bronn's.
" 30, line 2 from bottom, _for_ Tarapaca _read_ Tarapacá.
" 59, " 6 " " for _Pelecanoidinae_ read _Pelecanoïdinae_.
" 60, lines 14, 26, 34, for _Pelecanoides_ read _Pelecanoïdes_.
" 67, line 6, for _Thalassaeca_ read _Thalassoeca_.
" 70, " 10, _for_ Phaenicopteridae _read_ Phoenicopteridae.
" 91, " 12, for _ralloides_ read _ralloïdes_.
" 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 _cirrhatus_ read _cirratus_.
" 215, " 10, for _praelatus_ read _praelata_.
" 258, " 15, _for_ perhaps _read_ probably not.
" 351, " 11, and page 357, line 6, for _Phaenicophainae_ read
_Phoenicophainae_.
" 357, " 11 from bottom, for _Phaenicophaës_ read _Phoenicophaës_.
" 429, Fig. 89, for _jugularus_ read _jugularis_.
" 550, line 20, for _Seenopoeetes_ read _Scenopoeetes_.
" 568, " 9 from bottom, _for_ a scale-insect _read_ an Aphid.
{1}CHAPTER I
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[1] 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 _Sauropsida_ of Huxley, one of his three
primary divisions of Vertebrata. {2}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
existing 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 Cretaceous 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[2] constituting 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 arrangement 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_[3]
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 meanwhile 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 "webs" 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 {3}the barrel or quill (_calamus_ or _scapus_) at the
base of the tuft of rays have elongated into a principal shaft (_rhachis_);
this is generally accompanied by a secondary "aftershaft" (_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
(_cilia_), which in the distal rows usually terminate in hooklets
(_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 suppressed, 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_, _Rhinochetidae_, _Eurypygidae_, _Mesitidae_,
_Accipitres_ and _Psittaci_, in _Podargus_, _Coracias_, _Leptosoma_,
_Gymnoderus_ and _Artamus_.
COLOUR.–The colour of Feathers is due to one of three causes. First, an
actual pigment[4] 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.[5] Thirdly, the colour may be iridescent or prismatic; that is, a
blackish {4}pigment 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.[6]
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,[7] 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 increased size, the power of flight being attained very
early.
{5}The additional or spring moult affects the smaller feathers only, while
it is still doubtful how far changes of colour are clue 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-suit being changed first to a greyish habit and then to
a white; in Penguins the feathers of the wing come off in flakes.[8]
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 "analogous," the word in its strictest sense implying initial
diversity.
Any standard work on Vertebrate 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. Newton'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 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 {6}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.[9]
_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 characteristic 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 (as is common in
Reptiles); opisthocoelous when concave behind (as in many Mammals).
2. The Ribs are doubly attached to the vertebrae by a head (_capitulum_)
and a knob (_tuberculum_); 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 projection, 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.
[Illustration: FIG. 1.–Third cervical vertebra of Woodpecker (_Picus
viridis_). (Viewed anteriorly.) _Ft_, vertebrarterial foramen; _Ob_, upper
arch; _Pa_, articular process; _Psi_, haemal spine; _Pt_, _Pt_, the two
bars of the transverse process, shewn on one side ancylosed with the
cervical rib (_R_); _Sa_, articular surface of centrum. (From
Wiedersheim.)]
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 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.
{7}[Illustration: FIG. 2.–Skeleton of the trunk of a Falcon. _Ca_,
coracoid, which articulates with the sternum (_St_) at †; _Cr_, keel of
sternum; _Fu_ (_Cl_), furcula (clavicles); _G_, glenoid cavity for humerus;
_S_, scapula; _Un_, uncinate process; _V_, vertebral, and _Sp_, sternal,
portion of rib. (From Wiedersheim.)]
The absence of a keel is characteristic of the other and smaller group of
Birds, made up of the Ostrich, Rhea, Emeu and Cassowary, Moa and Kiwi,
which from the resemblance the sternum thus bears to a flat-bottomed boat
(_ratis_) 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}posterior
portion, where they often take the form of prolongations, the extremities
of which occasionally meet and enclose what are called _fenestrae_; but
these are unimportant when compared with the features presented by the
anterior part.
4. The Pectoral Arch, or Shoulder-Girdle, consists of three pairs of bones,
the Coracoids, the Scapulae or Shoulder-blades, and the Clavicles or
Collar-bones, the last two usually coalescing 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, while in
others, as several Owls, they do not unite. The furcula often ossifies
firmly with the anterior portion of the keel, and in _Fregata_, _Didus_ and
the _Ratitae_, the coracoids and scapulae are fused together.
[Illustration: FIG. 3.–Skeleton of the Limbs and Tail of a Carinate Bird.
(The skeleton of the body is indicated by dotted lines.) _F_, digits; _Fi_,
fibula; _HW_, carpus; _MF_, tarsometatarsus; _MH_, carpometacarpus; _OA_,
humerus; _OS_, femur; _Py_, pygostyle; _R_, coracoid; _Rd_, ulna; _Sch_,
scapula; _St_, sternum, with its keel (_Cr_); _T_, tibiotarsus; _Ul_,
radius; _Z_, _Z^1_, digits of foot. (From Wiedersheim.)]
5. The Anterior Limbs, or Wings, are composed of the Humerus, or upper
arm-bone, the Ulna and Radius (making 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
{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 _Sphenisci_ 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 Pelvic Arch consists of the Ilium, Ischium, and Os pubis, these
three paired bones meeting from each side at the cup (_acetabulum_) that
receives the head of the femur, and coalescing early in life; while the
_incisura ischiadica_ or notch between the ischium and the ilium becomes an
inclosed space (_foramen_) in all Birds except the _Ratitae_ and
_Crypturi_.
[Illustration: FIG. 4.–Pelvis of _Apteryx australis_. Lateral view. _a_,
Acetabulum; _il_, ilium; _is_, ischium; _p_, pectineal process of pubis;
_p^1_, pubis. (From Wiedersheim, after Marsh.)]
7. The Posterior Limbs, or Legs, are composed of the Femur or thigh, the
Tibia and Fibula, making the shank or "drumstick," and the bones of the
Foot. The thigh, however, being hidden by the plumage, the shank of a Bird
might easily be taken for the thigh, and the metatarsus (the cannon-bone 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
{10}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 _Sphenisci_ and to some extent
in _Psittaci_) do not lie in the same plane; the middle one having its
upper end thrust backward 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 _Ceyx_
and _Alcyone_, and the fourth (nearly) in _Cholornis_. The hallux, or hind
toe, has two phalanges, the second digit three, the third four, and the
fourth five; _Cypselus_ and _Panyptila_ (Swifts), however, are exceptions,
and possess only three in each of the anterior toes, while the
_Caprimulginae_ (true Nightjars) and _Pteroclidae_ (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 Roller); when this condition
is permanent, as in the _Cuculidae_, _Psittaci_ and _Pici_ the foot is
termed zygodactylous. In _Trogones_ the second toe is reversed
(heterodactylous). _Colius_ 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 commonly reduced and sometimes 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
{11}considerable help in Taxonomy (Classification). It must suffice here to
refer for the names of the parts to the subjoined figure.
[Illustration: FIG. 5.–Skull of a Wild Duck (_Anas boscas_), from the side.
_ag_, Angular; _als_, alisphenoid; _ar_, articular; _bt_, basitemporal;
_d_, dentary; _en_, external nostrils; _e.o_, exoccipital; _eth_, ethmoid;
_fr_, frontal; _j_, jugal; _lc_, lacrymal; _mx_, maxilla; _mx.p_,
maxillopalatine process; _n_, nasal; _p_, parietal; _pg_, pterygoid; _pl_,
palatine; _ps_, presphenoid; _px_, premaxilla; _q_, quadrate; _q.j_,
quadratojugal; _s.ag_, supra-angular; _s.o_, supraoccipital; _sq_,
squamosal: _ty_, tympanic cavity; _v_, vomer; _II_, foramen for optic
nerve; _V_, for trigeminal. (From Wiedersheim, after Parker.)]
The Bill, or Beak, is composed of an upper jaw or maxilla, and an under jaw
or mandible. From the figure it will be seen that "maxilla" is not strictly
the whole upper portion, though the term is thus used for convenience, as
is the plural "mandibles" for the two jaws when mentioned simultaneously.
The "rhamphotheca," or horny sheath, may be simple (undivided), or
compound, that is, made of several distinct pieces. In the _Anseres_ the
covering is soft with a horny (corneous) tip or "nail"; in the _Limicolae_
it varies extremely, producing a hard pickaxe, as in the Oystercatcher, or
a delicate sensory organ as in the Snipe and Woodcock. The rhamphotheca at
times has extraordinary outgrowths, as in the Hornbills, Sheathbills, and
elsewhere. In the _Accipitres_, or Diurnal Birds of Prey, and most
_Psittaci_, the base is soft and becomes a "cere," while the similar
formation in the _Columbae_ is due to a swelling of the _operculum_ or
covering of the nostrils. This operculum, moreover, may be leathery
(coriaceous), as in the _Charadriidae_, _Trochilidae_ and so forth, or
rolled up, as in _Rhinochetus_; it may even result in a short soft tube, as
in _Caprimulgus_, or in the hard double tube which gives the name of
_Tubinares_ to the Petrels. "Impervious" nostrils are those with a septum,
or division, between the nasal cavities, "pervious" {12}those with none.
The narrow slit-like or entirely closed nostrils of the _Steganopodes_
should also be mentioned.
The form of the bill varies from the "spoon" of _Platalea_ and
_Eurynorhynchus_ (spatulate) to the "arch" of _Numenius_, the scissors of
_Rhynchops_, the "wedge" of _Picus_, the big rounded feature of the
_Psittaci_, and so forth; but for details the characters of the several
Families must be consulted, as also for helmets, shields, horns, knobs, and
peculiarities due to the elongation, distorting or crossing of the
mandibles. These, too, are often notched, serrated, lobed or "festooned,"
or emarginate (slightly indented); the curious transverse serrations or
_lamellae_ of the beak in _Anseres_, and the somewhat similar sifting
apparatus in _Phoenicopterus_, _Prion_ and _Anastomus_ being especially
remarkable. Teeth were probably lost by Birds before Tertiary times, but
were possessed at least by _Archaeopteryx_, _Hesperornis_ and
_Ichthyornis_. The so-called "egg-tooth" of embryos is merely a calcareous
protuberance on the upper surface of the bill, which is cast after being
used to crack the shell.
9. The organs of deglutition and digestion begin with the tongue, which is
subject to much variation of structure, according to the different groups
of Birds, and is of course correlative with their habits. It has little
connexion with taste, though often of assistance in obtaining nutriment. To
this follows the gullet (_oesophagus_), which in many cases has an
enlargement forming the crop (_ingluvies_), wherein the food may be
temporarily retained before passing into the stomach, the last-named always
having an antechamber (_proventriculus_) where digestion is largely
accomplished, in front of the gizzard (_ventriculus_). This has frequently
strong muscular walls, and its action is often assisted by the mechanical
process of comminution performed by stones, grit or sand, swallowed for
that purpose. The stomach is succeeded by the intestines, which in most
cases have a pair of blind-sacs (_caeca_) attached to them, often acting as
aids to digestion, though these are not always functional, and are absent
in many Birds, while in others they attain a very large size, their
condition being in consequence of some importance as a systematic
character.
10. The organs of voice in Birds have long attracted special interest from
the loud cries which some utter, and the melody with which others are
gifted. Setting aside the part played by {13}the _trachea_ or windpipe in
supplying air to the lungs, its formation is worthy of attention. Its upper
end consists of the _larynx_, and it passes down the neck as a flexible
tube, formed by a continuous succession of bony rings connected by
membrane, until it bifurcates into two _bronchi_, which open into the
lungs. A common feature, found in many groups not nearly allied, is the
dilatation of a portion, generally near the middle, while a remarkable
modification is exhibited by the males of many of the Duck-tribe, some of
the lowest rings being fused together and forming what is known as the
_bulla ossea_ or "labyrinth." In other _Anatidae_ (some of the Swans), and
some of the Cranes, the trachea enters the keel of the sternum; but a not
unfrequent modification, usually confined to the male sex, often occurs
elsewhere, when the windpipe is looped back upon itself. All these
arrangements, however they may affect the sounds uttered by Birds, do not
in themselves constitute the voice organ of most. That is reserved for the
_syrinx_, a peculiarity of the Class _Aves_, consisting of the lower end of
the trachea and the adjoining part of the bronchial tubes; and the varied
modulations are effected by means of muscles attached thereto. These
voice-muscles may be wholly absent or of the simplest character, but they
attain their highest perfection in the _Passeres_, and especially in the
large group of them known as _Oscines_, where there are often five or seven
pairs. In this group the lowest four or five tracheal rings are solidly
fused into a little bony box communicating with the bronchi; the first and
second bronchial rings (or in this part often semi-rings) being closely
attached to the trachea, and the spaces between the second and third and
the third and fourth being generally closed by an outer tympaniform
(drum-like) membrane, while the rest of the semi-rings of the bronchi are
closed by the inner tympaniform membrane. It should be clearly understood
that all the notes emitted by Birds are produced by the above structures
only, and that the tongue has nothing to do with their utterance, except,
possibly, in the case of the sounds that Parrots (but not other birds) are
taught to produce.
CLASSIFICATION.–The Classification of Birds is still in a condition of
uncertainty, notwithstanding the many schemes successively propounded
during more than two centuries. To dwell upon them here would be
impossible, and it is only practicable {14}to trace in the briefest way the
line which has led to the most recent attempts, and to name those whose
researches have produced the results which may be fairly regarded as
attained. First among them is Nitzsch (1806-1840), to whom followed Merrem
(1812-1817), and after a few years L'Herminier (1827). These three worked
quite independently, and in their lifetime little notice was taken of their
labours; for, though there were good ornithologists among their
contemporaries, little value was then set upon internal characters in this
connexion. An improvement took place when the great Johannes Müller (1846,
1847) published his scheme for grouping the _Passeres_, which, though based
on purely anatomical facts, was almost immediately accepted, chiefly
through the simultaneous exertions of Dr. Cabanis, by systematists of the
Old School. For twenty years no advance was made, for the morphological
researches of Parker were not directly taxonomical; but Huxley (1867, 1868)
started what was practically a new line of investigation, though it
subsequently appeared that up to a certain point it had been already
suggested by Dr. Cornay (1842-1847). The impetus thus given was fortunately
sustained, Huxley's example being followed by Dr. Murie, and by two
promising men, A. Garrod and W. A. Forbes, both of whom died at an early
age, leaving their mark in work which, though much of it was crude, was
that of true genius. Mr. Sclater (1880) has tried to bring the results of
the whole four into harmony with pre-existing views, and a similar attempt
was that of Dr. Stejneger (1885); but all were overshadowed by the
monumental performance of Prof. Fürbringer, whose _Untersuchungen zur
Morphologie und Systematik der Vögel_, completed in 1888, must ever remain
a record of unexampled labour, while his considerations on the derivation
of Birds from Reptiles, and of the later groups of Birds from the earlier,
whether his results be right or wrong, are of the utmost importance to the
ornithologist. During the progress of this work the author was in frequent
communication with Dr. Gadow, himself engaged on the ornithological portion
of Bronn's _Thier-Reich_, and thus the opinions of each were in many cases
mutually affected. Dr. Gadow, on the completion of his undertaking,
propounded a scheme of classification, which is followed, with some slight
modifications, in the present volume (see foregoing table)–it being, of
course, understood that a linear arrangement is, {15}strictly speaking,
impossible, since any group may have a decided affinity to more than two
others. This Classification, beginning (as Birds themselves must have
begun) with the lower forms, takes us, except in the _Oscines_, as far as
the Families, which in most cases are fairly distinguishable, though of
very variable value. Coming to Genera, and still more to Species, the
opinions of authorities often differ so widely, that at present an attempt
to reconcile them is hopeless. It cannot be denied that Genera and Species
are merely "convenient bundles," and that divisions of either, if carried
too far, defeat the object for which Classification is intended. Genera are
only more distinct from Species, and Species from Races, because the
intervening links have disappeared; and, if we could have before us the
complete series which, according to the doctrine of Evolution, has at some
time existed, neither Genus nor Species would be capable of definition, any
more than are Races in many cases; while the same remark will apply to the
larger groups.
From these Races or Geographical variations we may not unnaturally turn to
GEOGRAPHICAL DISTRIBUTION. It will always be credited to Ornithology that
the interesting study of the Geographical Distribution of Animals was first
placed on a scientific basis as a result of the study of Birds. This was
effected by Mr. Sclater, whose division of the Globe into Six "Regions"–the
_Palaearctic_, _Ethiopian_, _Indian_, and _Australian_, forming one
group–the "Old World" (_Palaeogaea_); and the _Nearctic_ and _Neotropical_,
forming a second–the "New World" (_Neogaea_); was announced in 1858 (_J.
Linn. Soc._ ii. pp. 130-145). His scheme, being solely grounded on
Ornithological considerations, was accepted with scarcely any modification
by Mr. Wallace in his great work (_Geograph. Distrib. of Animals_, 1876),
and by the majority of zoologists, though some demurred, and among them
Huxley, who, in especial reference to Birds, shewed (_Proc. Zool. Soc._
1868, pp. 313-319) that there was more reason to divide the earth's surface
latitudinally than longitudinally, and that Four Regions were better than
Six–these four being (1) _Arctogaea_, comprising Mr. Sclater's Indian,
Ethiopian, Palaearctic, and Nearctic; (2) Austro-Columbia, corresponding
with the Neotropical; (3) Australasia; and (4) New Zealand–the last three
being combined as _Notogaea_. In 1882 Prof. Heilprin proposed to unite Mr.
Sclater's Palaearctic and Nearctic under {16}the name of Triarctic; but in
the next year (_Nature_, xxvii. p. 606) adopted for that union Prof.
Newton's earlier term Holarctic. Some other general schemes have been
promulgated, as those of M. Trouessart and Professor Möbius; but they have
found little support, and with regard to the Class _Aves_, though certainly
not with regard to other groups as _Pisces_, or _Mollusca_, what is
practically the scheme of Mr. Sclater has met with acceptance, whether with
or without the modifications proposed by Huxley and Professor Newton, there
being really but two important points of difference–(1) the recognition of
New Zealand as a distinct Region, and (2) the union of the Nearctic and
Palaearctic areas into a single Region. It would be impossible here to set
forth the arguments by which these views are maintained or contested, and
it must suffice to trace briefly the outlines of the several districts.
_New Zealand_, if admitted as a distinct Region, consists only of the
islands so named, the smaller Chatham, Auckland, and Macquarie groups,
Antipodes Island, Lord Howe's, Norfolk and Kermadec Islands. The
_Australian_, if the preceding be cut off, will include Tasmania, all
Australia, and the islands to the northward as far as what has been called
"Wallace's Line" (between Lombok and Bali), Celebes, New Guinea, New
Britain, and all the countless groups of tropical islands in the Pacific
Ocean–except the Galapagos, which undoubtedly belong to the next Region.
The _Neotropical_ is made up of all South America, the Antilles and Central
America, the only doubt being whether to draw the northern boundary so as
to exclude or include Mexico, or even the southern part of the United
States. To this naturally succeeds, but with an indefinite southern
boundary, the _Nearctic_, comprising the whole of the rest of North America
to the shores of the Polar Sea, with the addition of Greenland. Its
north-western corner, Alaska, is now known to be largely tenanted by forms
from Asia, not found elsewhere in America, and this is one of the chief
reasons assigned for uniting it with the _Palaearctic_ area, which may be
taken to include Japan and all continental Asia to the north of China
proper, the Himalayas, the Persian Gulf and the east end of the
Mediterranean. Some authorities would add Northern Arabia and Lower Egypt;
but all have agreed to include Tunis and the ancient Mauritania–the Barbary
States lying north of the Great Desert to the Atlantic Ocean about Mogador,
as well as the Canaries, Madeira and the Azores, with the whole of Europe
{17}from Greece to Iceland. What is left of Arabia and Africa, after taking
off the above portions, with the addition of Madagascar and the Mascarene
Islands, is the _Ethiopian_ Region; and all the rest of continental Asia,
with the islands not included in the Australian Region, becomes the
_Indian_, or, as it has lately been called, the _Oriental_. It would be
quite impossible to enumerate here the various Sub-regions and Provinces
into which these several Regions may be divided. The views of Mr. Wallace
are set forth at length in his excellent work, those of Mr. Sclater in _The
Ibis_ for 1891, pp. 514-557, and those of Professor Newton in his
_Dictionary of Birds_. Many writers would assign to Madagascar a higher
rank than that of a Sub-region.
MIGRATION.–Few peculiarities of Birds have excited more general interest
than their seasonal Migration, which in many species is so marked as to
have been observed from very remote times; and it is probable that nearly
all species are subject to periodical movements of varying extent. These
movements are greatest in the Birds which have their breeding quarters in
the northern parts of the Northern Hemisphere; and, with some exceptions,
it may be said that the more northerly is the range of a species the more
extensive are its migratory wanderings. In the Southern Hemisphere the
facts known are as yet insufficient to allow of safe deductions. Absence of
a food-supply in winter is alone enough to account for migration in the
above cases, and the return from the south in spring is probably due to the
desire of Birds to reoccupy their old haunts, or those in which they have
been bred. But just as there are some species which habitually breed within
the Arctic Circle and winter in the Tropics, there are others which may not
go so far in either direction, and yet have their movements governed by
exactly the same principle, with the result that in a temperate zone we
have Birds coming from the north to winter with us, while others, arriving
from the south in spring, spend the summer here, and depart towards autumn.
Others again, the true "Birds of Passage," arriving like the last in
spring, make little or no stay, but pass onward to more northerly lands,
and re-appear for as short a time in autumn on their return journey
southwards. Moreover, observation shews that, in most parts of the
temperate zone, there are many Birds which, though _resident as species,
are migratory as individuals_–that is to say, that while examples of
{18}the species may be met with at certain spots throughout the whole year,
those which occur at one season are not always the same individuals as
those which occur at another–the particular Thrush, Titmouse, or Finch,
appearing in the winter not being identical with that which appears in
summer. Again, among species of which some individuals are constantly
present throughout the year, a great accession to the numbers is made at
the close of the breeding-season by the influx of other individuals of the
same species bred in another district, though this influx generally lasts
for a comparatively short time, and the strangers pass on, accompanied it
may be, by some or even most of those that have been reared on the spot in
the season immediately preceding. These species are the "Partial Migrants."
It would at first seem from the above that the annual migratory movement
would be in a direction due north and south, or south and north, according
to season, and so in a general way it is; but there is no doubt that this
simple movement is disturbed by many causes, chief among which is possibly
the configuration of the land, which is found to give rise to considerable
deviations, and that to an extent which is at present very imperfectly
understood. It may be considered proved that the trend of a coast-line, the
course of a great river, or the intervention of a chain of mountains, has a
very appreciable effect on the direction taken by migrating Birds; but not
one of these, nor all in combination, affords a sufficient explanation of
all the deflexions, and will certainly not account for at least one
remarkable fact, as it may now be regarded–the tendency of many Birds in
Eastern Europe and part of Siberia to travel westward towards the close of
summer or in autumn. This is shewn in several ways, but in none better than
by the almost yearly occurrence in Britain at that season of examples of
species which breed only in the Russian Empire. For, admitting that such
examples are stray wanderers, which have lost their course, their
appearance here is still useful in indicating the existence of the westward
movement; and, with the evidence they furnish before us, we may judge
whence come vast numbers of others–Starlings, Crows, Rooks, Jays, Larks,
and what not, whose origin and starting-point it would be otherwise hard to
trace or even surmise. Much has been written, especially in Europe, on
so-called Lines of Flight, but as yet to little purpose, and indeed
{19}scarcely any writers on the subject have had sufficient _data_ to form
an hypothesis, so that it is not surprising that hardly any two agree in
theory.[10] In other parts of the world there is still less ground for
theorising, though in North America many valuable observations have been
made; and these, in conjunction with those carried on in Europe, will no
doubt in due time lead to satisfactory results as regards the Northern
Hemisphere. Concerning the Southern our ignorance is almost complete.
Of the way in which Migration is performed there is still much to learn–but
one thing is certain, all Birds do not migrate in the same manner. Some
gather in flocks, great or small, others seem to accomplish their northward
journey in pairs, or at any-rate arrive at their breeding-quarters already
paired. Some undoubtedly voyage by night, others may be seen to travel by
day. Of the Birds which in spring arrive unpaired, it is now incontestable
that the males outstrip or precede the females. There is, moreover, equal
diversity in the southward movements towards the close of summer and all
through the autumn. Of some species the earlier broods disappear without
attracting attention, and the later broods as well as the parents slip away
almost as imperceptibly. In one remarkable case, that of the Cuckoo, the
adults leave this country long before the young are fit to follow; but, in
by far the greater number, the young start first, and are followed, often
at an interval of some weeks, by their parents.[11] It is contended by many
that of actual Migration we see very little, since it is constantly carried
on at a height where the Birds are beyond our ordinary observation, and as
regards some species this seems to be true. Moreover, it would seem that
the longest flights are performed by night, and when the sky is clear, so
that only in thick weather do the Birds come near enough to the earth to be
heard–seeing them being of course impossible in the dark, though in a few
cases they have been telescopically observed passing across the face of the
moon. It is certain that many of the smaller land-birds gradually press
{20}onwards prior to leaving our shores, but after that they may possibly
betake themselves aloft to continue their journey.
The speed at which Birds travel during Migration is a matter on which very
diverse opinions have been and are held; but the highest estimates, such as
those of the late Herr Gätke (who would allow even 150 miles an hour), can
scarcely be otherwise than exaggerations; for there is no evidence of any
but exceptional performances at such rates, and there is really no reason
to suppose that Birds can fly faster at a higher elevation than at a lower.
[Illustration: FIG. 6.–A Falcon. To shew the nomenclature of the external
parts.]
TERMINOLOGY.–The annexed figure explains the nomenclature of most of the
outward parts of a Bird, but some further explanations may be given, as
below:–
{21}_Air-sacs._–Membranous receptacles, filled with air, in communication
with the respiratory organs or passages. Pouches are often exaggerated
air-sacs.
_Alar bar._–A coloured bar across the wing (_ala_).
_Allantois._–A vascular sack, growing from the hind-gut of the embryo and
enclosed by the amnion; the two fuse together and form the Chorion, which
lines the egg-shell, and takes upon itself respiratory functions.
_Altrices_ or _Nidicolae_.–Nestlings which, being hatched in a helpless
condition, are fed by their parents or inhabit the nest for a
considerable time.
_Amnion._–A membrane which grows in the developing egg from the ends and
sides of the embryonic area, and encloses the embryo at an early period.
_Bronchi_ (p. 13).
_Bronchial syrinx._–One in which outer tympaniform membranes exist
between two or more successive bronchial semi-rings, while an inner
tympaniform membrane may also be present. In typical cases the trachea
has no sounding membranes.
_Chest._–The same as the upper breast or base of fore-neck.
_Chorion._–See _Allantois_.
_Compressed._–Used of lateral compression as opposed to vertical
(depressed).
_Coverts (tectrices)._–Feathers that cover the base of the quills
(_Remiges_, oar-feathers) of the wing and of the tail (_Rectrices_,
steering-feathers). The wing exhibits several series above and below
(greater, median, and lesser). Unless otherwise specified, "coverts" in
the text refer to upper coverts.
_Cubitals._– See _Secondaries_.
_Cuneate._–Wedge-shaped.
_Decomposed_ (p. 3).
_Depressed._–See _Compressed_.
_Distal._–That end of any part or member which is furthest from the
imaginary centre or axis of the body.
_Dorsal._–The upper side of the body; and hence applied to the
corresponding surface of any part or parts of the structure.
_Filoplumes._–Filamentous or hair-like feathers.
_Flanks._–The portion of the sides near the leg.
_Graduated._–Used of the tail when its feathers diminish in length
gradually.
_Hackles._–Elongated and pointed feathers, as on the neck of Fowls.
_Heterodactylous_ (p. 10).
_Hyoid Apparatus._–The bony and cartilaginous framework of the tongue.
_Hypocleidium._–A projecting median process at the junction (_symphysis_)
of the clavicles.
_Lanceolate._–Used of the tongue, when pointed and (commonly) lengthened.
_Lore._–The space between the bill and the eye.
_Mantle._–The feathers of the upper back combined with the upper
wing-coverts.
_Manuals._–See _Primaries_.
_Nidicolae._–See _Altrices_.
_Nidifugae._–See _Praecoces_.
_Oil-glands._–Secretory organs near the root of the tail, probably used
in oiling the plumage. Some exhibit tufts of feathers, others are naked.
_Operculum_ (p. 11).
_Pamprodactylous_ (p. 10).
{22}_Patella._–The knee-cap.
_Pouches._–See _Air-sacs_.
_Powder-down patches._–Groups of powder-down feathers (p. 3).
_Praecoces_ or _Nidifugae_.–Nestlings which are hatched in a condition
that enables them almost immediately to leave the nest and feed
themselves.
_Primaries_ or _Manuals_.–Those wing-quills (_Remiges_), varying from ten
to twelve, borne by the _manus_ or hand. They should properly be counted
outwards from the _carpus_ or wrist.
_Procnemial._–In front of the knee.
_Proximal._–That end of any part or member which is nearest to the
imaginary centre or axis of the body.
_Racquet-shaped._–Used of feathers with bare shafts and roundish terminal
vanes.
_Rectrices_ and _Remiges_.–See _Coverts_, _Primaries_ and _Secondaries_.
The _Rectrices_ usually number twelve, but vary from four to
twenty-eight.
_Reticulated_ (p. 10).
_Rictal._–Belonging to the gape.
_Roofed._–See _Vaulted_.
_Sagittate._–Used of the tongue, and meaning arrow-shaped.
_Scapulars._–The feathers lying along the _scapulae_ or shoulder-blades.
_Scutellated_ (p. 10).
_Secondaries_ or _Cubitals_.–Those wing-quills (_Remiges_) borne by the
_Ulna_, which often exhibit roughnesses where they grow. They should
properly be counted inwards from the wrist, and vary from six to thirty
or more.
_Spatulate._–Spoon-shaped. Used of the bill or of racquet-shaped
feathers.
_Speculum._–Strictly applied to a band across the wing, more or less
metallic in colour, which occurs in the Duck-tribe.
_Square._–Used of the tail when level at the end.
_Syrinx_ (p. 13).
_Tectrices._–See _Coverts_.
_Tertials._–A mistaken word for the inner secondaries.
_Thighs._–Loosely used in describing plumage to indicate the feathers
falling over the leg.
_Trachea_ (p. 13).
_Tracheal syrinx._–One in which the lower portion of the trachea consists
of thin membranaceous walls, about six of the rings being thin or
deficient. Both inner and outer tympaniform membranes exist in the
bronchi, as well as some vibratory tracheal membranes. The few muscles,
generally but one pair, are wholly lateral.
_Tracheo-bronchial syrinx_ (the normal form).–One which has this
essential feature, that the proximal end of the inner tympaniform
membrane, forming the median wall of each bronchial tube, is attached to
the last pair of tracheal rings.
_Vaulted_ or _Roofed_.–Used of the tail when compressed like that of a
Fowl.
_Ventral._–The lower side of the body, in which lie the heart, lungs and
digestive organs; and hence applied to the corresponding surface of any
part or parts of the structure.
_Zygodactylous_ (p. 10).
{23}CHAPTER II
ARCHAEORNITHES–NEORNITHES RATITAE–NEORNITHES ODONTOLCAE.
The Class AVES is divided by Dr. Gadow (Bronn's _Klassen und Ordnungen des
Thier-Reichs, Aves, Systemat. Theil_, p. 299) into two Sub-classes of like
value, _ARCHAEORNITHES_ and _NEORNITHES_, though some writers prefer to
consider the former as of equal rank only to the several subdivisions of
the latter here adopted, namely, RATITAE, ODONTOLCAE, and CARINATAE
(p. 25). The question is clearly one of degree, and depends entirely on the
amount of weight assigned to the various points of distinction to be
mentioned below.
The Sub-class _ARCHAEORNITHES_ is at present represented by but one member,
the first undoubted fossil Bird, made known in 1861 by Andreas Wagner from
the Jurassic slate formation of Solenhofen in Bavaria, and now preserved in
the British Museum. This he described under the name of _Griphosaurus_; but
as Hermann von Meyer had already bestowed the title of _Archaeopteryx
lithographica_ upon a bird, presumably identical, a feather of which had
been obtained from the above system, the latter appellation has a prior
claim. In 1877 a second example, now at Berlin, was procured from the same
beds,[12] since which date Meyer's specific name has become firmly
established, in place of that of _macrura_ given by Owen to Wagner's
specimen.
This very remarkable animal, about the size of a Rook, is without doubt a
connecting link between Reptiles and Birds; but zoologists are practically
unanimous in regarding it as an Avine form, with Reptilian affinities and
probably arboreal habits.
{24}[Illustration: FIG. 7.–_Archaeopteryx lithographica._ The Berlin
specimen. (After Dames.)]
The sternum was possibly furnished with a weak keel, the strong wide
furcula was U-shaped, the ribs had no uncinate processes, while in all
probability the coracoid and scapula made a right, or even an acute, angle
at their junction, and the centra of the vertebrae of the neck and back
were biconcave. The bill was short and blunt, each side of the upper jaw
possessing about thirteen teeth, of which six seem to have belonged to the
praemaxilla; whereas in each side of the lower jaw only three can be
recognised, and those towards the anterior extremity. These teeth, conical
in shape and of fairly equal size, were fixed in a regular row, in distinct
sockets. The fibula and tibia did not coalesce, the latter exceeding the
metatarsus in length; the toes were four {25}in number, with two, three,
four, and five phalanges respectively, ending in claws, the hallux being
directed backwards. The manus had three free digits, and apparently three
free metacarpals; the pollex consisted of two joints, the index of three
and the third finger of four, while each had a strong hooked claw at the
tip. The hand was furnished with six or seven well-developed primaries,
attached to the third metacarpal and the second and third digits, the
number of secondaries being ten. The long Lizard-like tail had no terminal
pygostyle, but was composed of about twenty-one free post-sacral vertebrae,
of which the first twelve each bore a pair of large feathers, similar to
those of the wing, with the inner webs broader than the outer, and with
decided shafts.[13]
The Sub-class _NEORNITHES_ may be arranged, as above stated, in three
divisions, (A) NEORNITHES RATITAE, (B) NEORNITHES ODONTOLCAE, and (C)
NEORNITHES CARINATAE. The first of these contains the Ratite Birds proper
and possibly part of the so-called _STEREORNITHES_ of Patagonia (p. 43),
with several fossil forms of doubtful position from England, France, and
New Mexico, as will be seen below; the second the _HESPERORNITHES_ of the
Cretaceous Shales of Kansas, the _ENALIORNITHES_ of the Cambridge Upper
Greensand, and _Baptornis_ of the American Chalk; the third the
_ICHTHYORNITHES_ of the aforesaid Kansas deposits, and all other existing
Birds, with various extinct species closely allied to them.
Of the points of distinction between the Neornithes and the Archaeornithes
the most important are that the metacarpals are fused together, the second
digit being the longest, and the third more or less reduced; and that the
number of caudal vertebrae does not, as far as is known, exceed thirteen,
of which the last five or six combine together to form a pygostyle, except
in the Hesperornithes, Ratitae, and Tinamidae, where such is seldom the
case.[14] The _centra_ of the vertebrae also are concave on one side only,
except in _Ichthyornis_, and perhaps in _Enaliornis_. The possession of
teeth is, of course, exceptional, as is the remarkable loss of the keel of
the sternum in the Ratitae.
It is now generally, if not universally, agreed that Flightless Birds were
developed from those that could fly. It does not, however, necessarily
follow that the Neornithes are direct {26}descendants of the
Archaeornithes, as each may be a separate offshoot from the same parent
stem. All we can safely assert is, that the former were in existence about
the end of the Jurassic times, that teeth were still retained in some cases
during the Cretaceous Epoch, and that not only normal forms, but also
flightless forms without keel or pygostyle,[15] had arisen by that date.
(A) The RATITAE are commonly characterised as Birds with no keel to the
sternum; but this will not hold as a definition, since _Hesperornis_ has
also that peculiarity, while such genera as _Didus_, _Stringops_,
_Cnemiornis_, and _Notornis_ are nearly in the same condition. It is no one
point, therefore, but the sum of many, which enables us to draw so clear a
line of demarcation between this primitive group and the remainder of
existing forms; nevertheless it is convenient to preserve the name
unaltered, as it is well understood to what members of the class it is more
especially meant to apply. The rhamphotheca, or horny sheath of the bill,
instead of being simple, is composed of several more or less separate
pieces, as in the Procellariidae, Tinamidae, and Steganopodes; the quadrate
bone, by means of which the lower jaw is articulated to the skull, in place
of two proximal knobs has only one, as in _Hesperornis_, _Ichthyornis_, and
the Tinamidae; the coracoid and scapula are fused together, and meet at an
obtuse, as opposed to an acute or right, angle; and the last six or seven
caudal vertebrae do not coalesce into a pygostyle, or upright triangular
expansion to carry the rectrices, a state of things found elsewhere in
_Hesperornis_ and the Tinamidae.[16] The reduced wings preclude flight; the
tail is functionless, as in the Podicipedidae and Tinamidae; the tongue is
very small; the oil gland is absent; the penis is large and erectile, being
comparable to that of the Anseriformes; while in the adult the feathers are
evenly distributed over the whole surface, as in the Spheniscidae and
Palamedeidae, no down being present. Claws are found on the pollex and
index in _Struthio_ and _Rhea_, or occasionally on the third digit; in
_Casuarius_, _Dromaeus_, and _Apteryx_ they occur only on the index.
Ratite Birds proper are comprised in six groups, STRUTHIONES or Ostriches,
RHEAE or Nandus, MEGISTANES or Cassowaries and Emeus, APTERYGES or Kiwis,
DINORNITHES or Moas, and AEPYORNITHES or Rocs.
{27}I. STRUTHIONES.
Fam. STRUTHIONIDAE.–These birds are distinguished from all others by having
only two toes–the third and fourth–the terminal phalanges of which are
shortened and bear thick stunted claws, that of the outer toe being
commonly absent. The whole foot, including the long scutellated metatarsus,
is exceptionally stout, and the toes are padded beneath. The beak is short,
broad, and depressed, with deeply split gape; the head is small, with large
eyes; the neck very long; the wing- and drooping tail-feathers–the plumes
of commerce–are large and soft, with broad equal vanes. The furcula and
syringeal muscles are wanting, nor is there any aftershaft.
_Struthio camelus_, the Ostrich or "Camel-bird" of North Africa, now
extends from Barbary to Arabia, and even to Mesopotamia, though no longer
found, as of old, in Egypt or Central Asia, its former occurrence in
Baluchistan being somewhat open to question. It is black with white wings
and tail, having a flesh-coloured neck covered with brownish down, and
partially bare tibiae of the same hue. The female and young male are almost
entirely cinereous, while the chicks are clothed with bristly
yellowish-white down with blackish stripes. The eggs of the typical
northern bird have a surface like ivory, while those from Southern Africa
are marked with close-set pits, whence some authorities recognise a
different species (_S. australis_) in the latter region, distinguishable,
moreover, by the bluish colour of the naked parts. Examples from Somaliland
and the adjoining districts of East Africa to Lake Tanganyika are separated
as _S. molybdophanes_, on account of the leaden colour of the unfeathered
portions, coupled with a red patch on the front of the metatarsus. The eggs
are smoother than in the southern species, but similarly pitted. The fossil
forms _S. asiaticus_ from the Pliocene of the Siwalik Hills of India, and
_S. karatheodori_ from the Upper Miocene of Samos complete the family,
while _S._ (_Struthiolithus_) _chersonensis_ has been founded on a
petrified egg from the government of Cherson in South Russia.
{28}[Illustration: FIG. 8.–Ostrich. _Struthio camelus._ × 1/19.]
The Ostrich stands about eight feet high, being the largest of existing
birds; it frequents sandy wastes and dry arid localities, such as are found
in the Sahara and the plains and valleys of Southern Africa, while
districts studded with low bushes are not unfrequently tenanted. Though the
fable of the head being hidden to avoid detection is of course devoid of
foundation, this species is timid and wild in its native haunts, and being
keen-sighted as well as wary, gives an impression of great restlessness.
From the fact that a single stride is said to cover twenty-five feet or
more, it will readily be understood that the speed is very great, exceeding
that of a galloping horse; but, owing to its habit of running in a curve,
it is generally possible to intercept the bird's path at a distance from
the point where it started. In motion the head is held forward, and the
wings are outspread, while both beak and feet are used as weapons of
defence when capture is imminent, the latter delivering strong sideways
kicks, which make close quarters very dangerous. Forty or fifty individuals
may at times be seen in company; the usual parties, however, consist of
five or six at most, especially during the breeding season, when the
polygamous cock escorts a flock of several hens, obtained by battle or
allured by courting performances earlier in the season. A liking for the
companionship of zebras, hartebeests and other antelopes, has been noticed
by various observers. The cry is said to be hoarse and mournful, resembling
the roar of a lion or {29}the lowing of an ox; but Ostriches are, as a
rule, decidedly silent. In a state of nature the food consists chiefly of
herbage, including seeds and fruits; in captivity the diet is of every
description, and even in a wild condition small mammals, birds, reptiles,
and insects are eaten, with a quantity of grit to aid digestion. In
confinement the birds become very tame, and will then swallow bones, nails,
and the like–in fact almost anything they can pick up. They can exist for a
long time without water, but drink regularly when opportunity offers; they
show a liking for salt, and will bathe in the sea or in rivers, immersed up
to the neck. The hens belonging to one cock lay in the same nest, which is
a fairly shallow excavation dug in sand or dry soil, and surrounded by the
material thrown out during the process, or more rarely by an edging of
grass. The spot is hard to discover in the desert, the stride being too
long for tracks to be of much assistance. More than thirty yellowish-white
eggs are sometimes deposited within the pit in circular arrangement, and
many more are dropped around, to serve, it is asserted, for food for the
newly-hatched young; in the wild state, however, the average number is
probably less. The contents, equal to those of some two dozen hens' eggs,
are used for food by the natives, the shells forming convenient pots for
water and so forth. The cock undertakes almost the whole duty of
incubation, being occasionally relieved by the hens during the daytime;[17]
but when the sun is hot no brooding is necessary, though a covering of sand
is superposed to guard the spot from the depredations of marauders. The
chicks, which run from the shell, are hatched in six or seven weeks, and
are accompanied by both parents, the male often counterfeiting wounds to
draw away the intruder, circling around with drooping wings or throwing
himself down as if in extremities.
Ostriches were well known to the ancients, who used the plumes for
ornament, as we do; these were considered emblems of justice from the
equality of the two webs, or were worn in token of victory, as is still
done in some parts of Africa. The words of Aristotle–who was followed by
Pliny in the statement that the Ostrich was part quadruped, part
bird–combine with those of Xenophon to bear witness to this knowledge,
while monuments, inscriptions, and even the Bible tell the same tale. In
the Sahara and elsewhere these birds are hunted with horses and camels,
{30}being stalked or ridden down by means of fresh relays of beasts; the
Namaquas draw a cordon round them; the Bushman, concealed in sand or
disguised in skins, shoots them with poisoned arrows; while the lasso,
pitfall, or other device are used in particular districts. Space will not
permit a detailed account of the Ostrich farms of modern Africa, so well
described in Messrs. de Mosenthal and Harting's _Ostriches and
Ostrich-Farming_, and other books; but it may be mentioned that the tribes
of the north of that continent have long been in the habit of domesticating
the bird, that the value of the sales in South Africa is not far from a
million pounds yearly, and that the plumes are plucked or, preferably, cut
about twice a year, the adults yielding the finest feathers. The flesh is
coarse, and of little use for food.
II. RHEAE.
Fam. RHEIDAE.–The Rheas, or Nandus, have the head, neck, and bill much like
those of Ostriches, the maxilla being somewhat more rounded and terminating
in a nail-like process; the metatarsus is also similar and equally stout in
proportion, but the toes are three in number in place of two, the
mid-phalanges being shortened and the terminal furnished with decided
claws. In _Rhea darwini_ alone the metatarsi are mainly reticulated instead
of scutellated anteriorly, and have the upper portion feathered. The bones
of the wing are comparatively well developed, the feathers being slender
but not ornamental, while there is no apparent tail. The furcula is
wanting, as is the aftershaft to the feathers, but the syrinx is
tracheo-bronchial with one pair of syringeal muscles, a condition
absolutely unique among the Ratitae. The head and neck are feathered, only
the lores, orbits, and ear-openings being naked, and of these the latter
are surrounded by bristles.
_Rhea americana_, the so-called American Ostrich, the Ema of the
Brazilians, the Avestruz, Nandú, or Chueké of Argentina, is found from
Bolivia, Paraguay, and South Brazil to the Rio Negro, if not further; it is
brownish-grey with blackish crown, nape, and breast, white thighs and
abdomen, and yellowish neck. The sub-species _R. macrorhyncha_ of
North-East Brazil is darker, with longer bill and more slender metatarsi.
_R. darwini_, which occurs south of the Rio Negro, and up the Andes to
Tarapaca, is buffish-brown, with whiter underparts and white margins to the
{31}feathers of the wings and back. Hens are not so dark, and Mr. Hudson
says[18] that in _R. darwini_ the young are dusky grey and are hatched with
the legs feathered to the toes. Rheas are shorter than Ostriches by about a
couple of feet, _R. americana_ being the largest form; the feathers are
much rounded, broad, and very soft. Fossil remains occur in the Upper
Tertiary or quite recent deposits of South America.
[Illustration: FIG. 9.–Nandu. _Rhea americana._ × 1/20.]
The members of this family find their favourite haunts on the treeless
flats of the Argentine pampas, the scrub-covered plains of Patagonia, or
the dry open Sertões of Brazil, where their acute vision enables them to
detect the approach of enemies from afar. Small flocks of from three to
seven individuals are met with at certain seasons, and parties of twenty or
thirty at other times–often with deer or guanacos–so it would appear that,
as in the case of the Ostrich, larger companies are formed after the young
are able to provide for themselves. The birds become exceedingly tame when
not molested, but when danger threatens they run at great speed, doubling
upon their pursuers constantly, or crouching down among bushes or other
cover, if they think they can escape observation. In the latter case they
will lie closely until almost trodden upon, and may be shot before they
rise by the hunter who cautiously approaches their hiding-place, as the
head is usually visible above the surrounding vegetation. When moving at
full pace the wings have normally a somewhat drooping position, but they
are raised alternately above the back–apparently {32}to aid progress–when
fresh exertions are necessary. Mr. Hudson tells us[19] that Darwin's Rhea
"carries its neck stretched forward, which makes it seem lower in stature
than the allied species." The diet consists chiefly of grass, roots, and
seeds, but berries of _Empetrum_ are a favourite food, and lizards,
insects, worms, and molluscs are said to be eaten, together with hard
substances to promote digestion. Nandus take readily to the water, and can
swim across a river several hundred yards wide, the body being hardly
visible. In spring the cock utters a deep, resonant, booming noise, a loud
hiss being not uncommonly heard also; while at that season the rival males
attack each other viciously with their beaks, trampling down the ground in
their passion, but not generally using their feet, as they do when wounded.
The hens secured by each of the cocks lay together in a mere depression in
the soil with very little, if any, lining; the eggs numbering from twenty
to thirty, or exceptionally more, besides those scattered about outside the
nest. Here again Mr. Hudson is our authority for stating[20] that the eggs
of _R. americana_ are golden yellow when fresh, those of _R. darwini_ deep
rich green; both however fade quickly to a whitish colour. The male
incubates very closely for about six weeks, often taking up his position,
as the Ostrich does, before the final egg is laid; he afterwards attends
upon the young, and charges intruders who seem dangerous, with outstretched
wings and beak. Rheas may be captured by riding after them in a semicircle,
which closes upon them as they go, or by means of long-winded hounds; but
the most usual method is that of hurling the "bolas" or leaden balls
connected by leather thongs, which wind around the bird's neck or legs, and
thereby hamper its movements or throw it down. The feathers, though
inferior to Ostrich plumes, are much used for brooms and the like, and are
said to be called "Vautour" in the trade. The flesh is very poor. These
birds have bred both on the Continent and in Britain.
III. MEGISTANES.
The MEGISTANES comprise the _Casuariidae_ or Cassowaries, and the
_Dromaeidae_ or Emeus, the following being the chief peculiarities of the
group. The wings are quite rudimentary; {33}the aftershaft of the contour
feathers is extremely large, so that they appear to be double; three front
toes are present, with shortened mid-phalanges and large claws; and the two
clavicles do not meet. The lack of ornamental wing- or tail-plumes, and the
hair-like nature of the coat is also characteristic, while, as opposed to
_Rhea_, there is no indication of syringeal muscles. Within the group
itself the Cassowaries are distinguished from the Emeus by the points next
to be mentioned. The former have a compressed keeled beak and a large
casque of bony tissue upon the bare head, the greater part of the neck
being also naked and in most cases wattled; the remiges are reduced to
thick black barbless quills from four to six in number, and the inner toe
has a particularly long sharp claw. Emeus, on the contrary, have a broad
depressed beak, short feathers on the head and neck, no helmet, wattles, or
spines on the wing, and an ordinary claw on the inner toe. Both Families
have long necks, stout metatarsi covered with coarse roundish scales, and
toes padded below; the tibia being nearly, if not quite, covered by the
plumage.
Fam. I. CASUARIIDAE.–Following Professor Salvadori,[21] Cassowaries may be
divided into two groups: the first with the helmet laterally compressed,
and the second where it is triangular and pyramidal, or even depressed.
They are all large birds, though smaller than Emeus, which are only
surpassed in size among existing forms by the Ostrich; the colour of the
coarse but glossy hair-like plumage is black, and similar in both sexes;
the hen is bigger than the cock, as is also the case in the Dromaeidae and
Apterygidae.
Of the first of the above groups, _Casuarius tricarunculatus_, from Warbusi
in New Guinea, which is possibly a "sport," has two lateral wattles on the
fore-neck and a third small median caruncle at a lower level. _C.
bicarunculatus_, of the Aru Islands, has two long distant reddish-violet
wattles, a black casque, bluish-green head, and blue neck with some red
behind. _C. galeatus_ of Ceram, the species first known to ornithologists,
is similarly coloured, though less brightly, and has the flesh-coloured
throat-wattles close together, and a naked reddish-purple space on each
side of the neck. The larger _C. australis_ of North-East Australia has a
higher helmet, a brighter blue throat, and a few scattered hairs on the
wattles, which Wall, who discovered the species, said were coloured with
blue and scarlet. _C. beccarii_ of the Aru Islands, {34}Middle and South
New Guinea, has the front and top of the casque black, its sides greenish,
and its back yellowish; the head is grey-blue, the throat and sides of the
neck are blue, the hind-neck is red and orange, a yellow streak running
across to the mandible; a bare space on each side of the base of the neck
is flesh-coloured, and the long single neck-wattle of the same colour is
somewhat deeply divided at the tip.
Of the second group, _C. uniappendiculatus_ (Fig. 10), of Salawatti and the
adjoining parts of New Guinea, has the head, throat, and nape blue, the
lower portion of the neck and the median pear-shaped caruncle yellow, the
casque dusky olive, and a longitudinal naked space towards the sides of the
neck flesh-coloured with a yellow margin. _C. occipitalis_ of Jobi Island
is distinguished from the last-named by a large occipital spot of yellow
and a paler helmet; while the remaining three forms have no wattle at all.
Of these, _C. papuanus_, of North-East New Guinea, has a dusky black
casque, blue head, throat, and fore-neck, grey-green occiput and auricular
region, and orange hind-neck changing into rosy flesh-colour towards the
sides. _C. picticollis_ of South-East New Guinea has a black helmet,
grey-blue occiput, violet-blue nape, pale blue hind-neck, red throat and
longitudinal space on the sides of the lower neck; _C. bennetti_ of New
Britain differing in having the head and neck of an almost uniform blue.
Nestling Cassowaries are clothed in rusty brown, relieved by darker
stripes; at a later period they become more tawny, and the black plumage
begins to appear; but a few hair-like feathers remain on the head for some
time, while the helmet is very gradually developed from a flat Coot-like
shield, though the gaudy colours of the neck and wattles are assumed much
earlier.[22]
All the species of this family inhabit wooded country, commonly of the
densest description, though often found in more open scrub and in the
neighbourhood of creeks and watercourses. Naturally shy but inquisitive,
they have been rendered doubly wary by man's persecution since their haunts
have been invaded by colonists. They dislike the sun, and emerge from cover
only in the morning and evening, seeking their favourite spots, where they
feed chiefly on fallen fruit, varying this diet with insects and
crustaceans. Berries, leaf-buds, and bulbs are, however, also eaten, with
grit and pebbles for digestive purposes, and in captivity they are almost
omnivorous.
{35}[Illustration: FIG. 10.–One-wattled Cassowary. _Casuarius
uniappendiculatus._ × 1/14. (From _Nature_.)]
In this state they become extremely tame, and are kept like fowls by the
natives of some districts, who consider the flesh very palatable; while in
Queensland the adults are said to be hunted with dogs. The plumage is used
for the manufacture of mats, rugs, head-ornaments, and the like.
Cassowaries run with wonderful swiftness, though rather heavily, diving
into the bushes at a moment's notice, or aiding themselves by their wings,
and leaping over obstacles as much as six feet high, if shelter is not
readily available. They usually rest on the whole of the metatarsus, but
sleep on the breast, or perhaps occasionally on the side; at other times
they will dance about with contortions of the neck, or roll on the ground
like playful monkeys. Old males become very fierce when driven to bay,
kicking out in front or sideways, ruffling up their feathers and using
their beaks at the same time. In the wet season swimming is a common
practice, wide rivers being {36}crossed with ease, and in the absence of
other bathing-places the sea is often utilised. The note in a state of
excitement is a sort of grunt or snort, the call to the young being of a
lowing nature; but the ordinary voice is loud, guttural, and unearthly,
consisting of quickly-repeated croaking sounds, lasting for as long as
three minutes, and audible at a distance of a mile, or considerably more.
The female is much quieter, while the "Mooruk" (_C. bennetti_) is stated to
utter a low scolding or plaintive whistle. A rough nest of leaves and grass
is formed in a depression of the soil, generally below bushes or tangled
undergrowth, in which from three to six very large eggs are deposited,
placed in the shape of the letter V. These are normally light green in
ground colour, with close-set granulations of dark bright green; but one,
if not more, is ordinarily of a perfectly smooth texture, and is therefore
entirely light green. The cock incubates, it appears, solely, though some
say that the hen takes her turn; and the former tends the young when
hatched, the period of sitting being about seven weeks. The nest is said to
be covered by the parent if left for a time, but this is uncertain, as is
the use of the two or three eggs scattered round the nest, which are
asserted by natives of widely-distant districts to furnish food for the
chicks. After breeding, small flocks are formed in some cases, possibly by
the combination of two families. The Ceram species, which seems to have
been called "Emeu" or "Ema" by the early Portuguese navigators, often lays
in captivity, while _C. bennetti_ has bred in the gardens of the Zoological
Society of London.
Fossil remains occur in Australia. _Hypselornis sivalensis_ is an allied
form from the Pliocene of the Siwalik Hills in India.
Fam. II. DROMAEIDAE.–From about the beginning of this century the name
"Emeu," used, as mentioned above, in varying form for both the Rhea and the
Cassowary, has been restricted to the genus _Dromaeus_, the members of
which stand more than five feet high, though lower on their legs than an
Ostrich. _D. novae-hollandiae_ of the interior of Eastern Australia, which
extended in times past to Tasmania and the islands in Bass's Straits, is
blackish grey, with black tips to the plumage. _D. irroratus_, a more
slender species from West, and probably the adjoining parts of South,
Australia, has each feather transversely barred with dark grey and white,
and a rufous margin to the black patch at the end. Young birds in down are
greyish-white, with longitudinal blackish streaks above, {37}and spots on
the head and lower parts. The sexes are similarly coloured, both possessing
a remarkable tracheal pouch, connected by a slit with the windpipe, and
only fully developed in adults.[23]
[Illustration: FIG. 11.–Emeu. _Dromaeus novae-hollandiae._ × 1/20.]
In their general habits Emeus are not unlike Cassowaries, but they inhabit
sandy plains or open forest districts, being invariably monogamous, though
seen in small parties after breeding. Their sight is keen, they run
strongly and rapidly, rest on the whole metatarsus, and kick out backwards
towards the side. The food is of fruit, roots, and herbage, generally
obtained in the morning or evening; water is freely drunk, and the birds
love bathing, being capable of crossing even a broad river. They utter at
times a hissing or grunting sound, but in the nesting season a peculiar
loud booming or drumming note is produced, probably in connexion with the
tracheal pouch. The nest may be a mere hollow scraped in the ground, with
or without a surrounding ring of grass or plant-stems, or a mound of
bark-scales some three inches high[24]; the eggs are from seven to thirteen
in number, or even more, and are of a dark, or occasionally light,
{38}green colour, while the surface is covered with granulations which give
it the appearance of shagreen. They are small for the size of the bird,
being less than those of the Cassowary. The cock performs the duties of
incubation, and it is very doubtful whether the hen ever assists him; the
chicks break the shell in about eight weeks. The flesh is moderately good
for eating, and the fat below the skin yields a large quantity of oil. The
birds are constantly hunted with dogs or shot on account of the damage they
do to wire fencing and the grass they devour. Emeus are easily
domesticated, and propagate readily in semi-confinement, being perfectly
hardy in Britain and elsewhere.
_D. patricius_ is a fossil species from the Pleistocene of Queensland and
New South Wales. _D. gracilipes_ is another extinct Australian form, but
_Dromornis australis_ of Queensland may indicate a distinct group of
Ratitae.[25] _Dromaeus ater_, of Kangaroo Island, off the south coast of
Australia, is now extinct, though a stuffed skin and a skeleton are in the
Paris Museum.[26]
IV. APTERYGES.
The APTERYGES, or Kiwis, have been recently shown to be much more nearly
related to the Dinornithes than to the remaining Ratite forms, and are
accordingly placed in close proximity to them in the classification here
adopted. Professor T. J. Parker has, moreover, lately formulated a new
system–excluding the _Aepyornithes_, which may commend itself to many
persons as a further improvement.[27] In this, the Order _Struthiones_
contains the family Struthionidae, and the _Rheae_ the Rheidae; but the
third Order, upon which the name _Megistanes_, Vieillot, is bestowed,
includes two Sub-Orders–_Casuariformes_, comprising the Casuariidae and
Dromaeidae, and _Apterygiformes_, with the Dinornithidae and Apterygidae.
In other words, the original stock is considered to have produced three
Ratite branches only, the third of which gives rise to two twigs, each of
these separating again into two smaller twigs representing the Families.
Fam. APTERYGIDAE.–These birds are at once distinguished {39}from all their
allies by their small size, and by the long, weak, decurved bill, which
tapers regularly and has the nostrils placed almost at the extremity. The
head and eyes are comparatively small, as will be seen to be the case in
the Dinornithidae. The legs are very stout and situated backwardly, a small
elevated hallux is present, and the toes are provided with long, sharp
claws. The moderate metatarsus is reticulated in the young, but is clothed
with fairly large scutes in the adult, when it becomes much smoother. The
wings are small-boned and invisible, with functionless quills, the tail is
rudimentary, the aftershaft and furcula are absent, while many elongated
hairs occur on the front of the head.
[Illustration: FIG. 12.–Kiwi. _Apteryx australis._ × ⅐.]
These curious flightless birds are confined to New Zealand, whence a
specimen was brought to England as early as 1813. _Apteryx mantelli_, of
the North Island, is deep red-brown with longitudinal streaks of
yellowish-brown, the head being darker and the lower parts greyer; _A.
australis_, of the South Island, is lighter, and feels soft instead of
harsh when grasped. _A. oweni_, of both islands, is much smaller, and is
light grey-brown, transversely marked with blackish bars. _A. haasti_, also
said to occur in both islands,[28] is a larger and darker form of the last
named; _A. lawryi_, of Stewart Island, hardly differs from _A. australis_;
while _A. maximus_, of Verreaux, is a very doubtful species. Mr.
Rothschild[29] has founded a sub-species (_occidentalis_) {40}on examples
of _A. oweni_ from the North Island and the west of the South Island. In
all these birds the lanceolate feathers have a hair-like texture, due to
the disunited filaments of the upper portion, the lower part being covered
with grey down, and the rhachis more or less exserted. The tibia is
feathered, the bill being yellowish, and the feet brown or black. The
female is similar, but larger, the young blacker. Mr. Lydekker has
described a fossil species, _Pseudapteryx gracilis_, from New Zealand,[30]
and Mr. De Vis _Metapteryx bifrons_ from Queensland.[31]
Kiwis inhabit wooded country and hills up to the snow line; they are still
met with at low elevations on a few islands, but their retreats are now
chiefly on the slopes and in the gullies of the mountains, where a dense
undergrowth of shrubs and tree-ferns shades a carpet of creeping vegetation
and moss. Here parties of from six to twelve used to be seen, though in the
breeding season they separated into pairs, but at the present day flocks
can hardly be hoped for. In the daytime these shy birds hide in burrows in
the ground, or natural cavities under tree-roots or rocks, while towards
dusk they emerge in an animated condition. The direct rays of the sun seem
to dazzle them, and they roll themselves up into a ball, if not disturbed;
when stirred up they are somewhat sleepy and quickly retreat to cover.
Lengthy strides carry them along at a great pace, the body being held
obliquely with outstretched neck; and, if molested, they ruffle up the
plumage and snap the bill, while striking viciously with their feet at the
intruder, the leg being drawn up to the breast and the blow delivered
downwards. Sometimes they rest upright with the point of the bill touching
the ground, sometimes upon the whole metatarsus, but usually they are seen
at feeding time cautiously moving from spot to spot, and tapping the soil
or the walls of their cage with their long sensitive beaks. A sniffing
sound accompanies this operation, and probably the smell of food assists in
its discovery, yet the sense of touch is no doubt the primary agent. The
diet consists chiefly of worms, in search of which the ground is deeply
probed, and shows funnel-like holes scattered over its surface; when a
capture is made the worm is extricated with a gentle wriggling motion, and
is either beaten upon the ground to kill it, or swallowed at once with a
jerk of {41}the head. Grubs, beetles, molluscs, and berries are also eaten,
with grit or pebbles as digestives. The loud whistling note, which gives
the name to the Kiwi, is chiefly heard on light nights, that of the female
being shorter, and the young uttering a chuckling or kitten-like cry.
Growls are emitted by the birds when disturbed, and they have a curious way
of yawning in the daytime. The nest is usually in an enlarged space at the
end of a round tunnel in the soft earth, said to be made by the female
alone, the opening being under a tree-root, a stone, or a tussock of grass;
it consists merely of a little dry fern, herbage, or a few leaves. The
eggs–generally two in number, though one is often found, and three are
recorded–are enormous for the size of the bird, and are equal to a quarter
of its weight; they are pure white, or slightly green in hue, with a smooth
surface, recalling by their appearance those of the Whooper. The Maories
are very fond of the flesh, either roasted or boiled, and hunt Kiwis
systematically with muzzled dogs, while of old the chiefs utilised the
plumage for ornamentation. The cock performs most, if not all, of the
duties of incubation, and attends upon the young. Females lay in captivity,
but no chicks appear to have been hatched as yet under these conditions.
V. DINORNITHES.
The Family DINORNITHIDAE contains those well-known extinct New Zealand
forms the Moas, as they are supposed to have been denominated by the
Maories, some of which were of gigantic size. The larger species must have
been comparatively rare, judging by the fossils obtained, while some seem
to have survived until about four or five hundred years ago, or even a
century later in the South Island. Being flightless, these birds were
easily slaughtered by the natives, who were very fond of the flesh, and
captured them when exhausted by repeated spear-wounds, after they had been
driven from their retreats by burning the grass and vegetation. It was not
until the year 1839 that a femur-shaft was exhibited by Owen to the
Zoological Society of London, that being the first portion of a Moa known
to have reached this country; but since the above date an immense quantity
of bones of all descriptions have been procured in many parts of both the
North and the South Islands, some hidden under the sand or exposed upon
{42}its surface, some in marshes and superficial deposits generally, and
others in caves, hollows of rocks, or cooking places of the former
inhabitants. Footprints have been observed in the sandstone; portions of
muscles, ligaments, and even of skin have been discovered; and, most
remarkable of all, feathers have been met with of fresh appearance and
unfaded colours. Pebbles used to aid digestion, and eggs, both whole and
fragmentary, complete the list.
Moas had comparatively small heads, and also small orbits and eyes; the
bill varied, as will be seen below; the legs were stout, though not always
equally so, a hallux being usually present; the wings were extremely
reduced, or even wanting; the furcula was absent, and the aftershaft of the
larger feathers was of great size. The neck is supposed to have been
partially bare, while the webs of the rounded feathers were disunited and
more or less downy below. Some of the latter were black, with red-brown
bases and white tips, others were blackish-brown or yellowish.
Professor Parker, in his recent memoir,[32] proposes three Sub-families,
_Dinornithinae_, _Anomalopteryginae_, and _Emeinae_; _Megalapteryx_, which
he omits, possibly representing a fourth. The first of these contains only
one genus, _Dinornis_, with wide convex sternum, comparatively slender
limbs, broad skull, and long, wide, deflected beak; the height of _D.
maximus_, the largest of the whole group, being estimated at about twelve
feet. The second Sub-family comprises three genera, _Pachyornis_,
_Mesopteryx_, and _Anomalopteryx_, forms of small or moderate height and
varying bulk, with less broad skulls and pointed beaks, the sternum ranging
from long and narrow to wide and flat. The third possesses a single genus,
_Emeus_, in which the limbs are heavy, the strongly-built skull is narrow,
and the beak short and broad. _Pachyornis elephantopus_ has extraordinarily
stout, short legs, while _Anomalopteryx parva_, perhaps the smallest Moa
known, is said to have been about the size of a turkey. The validity of
some genera and species is, however, questionable. Most writers think that
the female was larger than the male. Mr. De Vis has described a fossil
femur from Queensland as _D. queenslandiae_,[33] but it may belong to the
Dromaeidae. According to native testimony the habits were sluggish, but the
birds were dangerous to approach; they lived in pairs and fed upon green
shoots and roots of ferns, making a nest of a pile {43}of grass and leaves.
We are told that the eggs found with the remains were dark green, light
green, or yellowish, but the last colour at least probably refers to faded
specimens.
VI. AEPYORNITHES.
Quite as remarkable as the Moas are the immense, massive-limbed forms of
the Family AEPYORNITHIDAE, supposed by many to be identical with the "Ruc"
or "Roc" of the Venetian traveller Marco Polo, and of the _Arabian Nights_.
If this is the case, the size of the birds and their eggs must have been
absurdly exaggerated, since the largest species known probably stood about
seven feet high, and the egg is certainly not as big as a butt;
nevertheless, the fact of the Roc being accredited to Madagascar makes it
probable enough that the fables were engrafted upon _Aepyornis_, which was
an inhabitant of that island. The eggs were first brought to the notice of
ornithologists by Strickland in 1849, while soon afterwards Isidore
Geoffroy St.-Hilaire obtained two of them, with some fragments of
bones.[34] These eggs, which exceed all others in magnitude, measuring some
thirteen inches by nine and a half, have now been obtained in considerable
numbers, with a large quantity of fossil remains of the birds themselves;
and in consequence about twelve species have been indicated, and a second
genus, _Mullerornis_.[35] It is supposed that some of them were in
existence not more than two hundred years ago. The most salient points of
their structure are the long, stout legs, with four toes and broad flat
metatarsi, the apparently rudimentary humeri, the absurdly short keel-less
sternum, and the frontal pits, indicating a large crest, comparable to that
supposed to have existed in certain of the Dinornithidae.[36] The shell of
the eggs, some of which contain two gallons, is used by the natives to hold
liquor, and is slightly pitted.
* * * * *
It will be remembered that, in the arrangement here followed, Dr. Gadow
placed the _STEREORNITHES_ under the head of Neornithes Ratitae, though not
under that of Ratitae in the restricted {44}sense; but it should be noted
that their systematic position was not by any means assured, though
justified by what was then known of these extraordinary fossils, of which
the sternum has not even yet been brought to light. Remains of various
forms, chiefly of gigantic size, have been disinterred from the Miocene
strata of Santa Cruz in Patagonia, one of which (_Phororhachos_) was
described in 1887 by Dr. Ameghino,[37] from its mandible as an Edentate
Mammal, though four years later[38] he arrived at the more correct
conclusion that the jaw was to be referred to a bird. In 1891, moreover,
Señores Moreno and Mercerat[39] proposed a new Order with the name of
_Stereornithes_, when publishing a series of fine plates; while Dr.
Ameghino, who criticised their work, reduced the nine genera created
therein to the smaller number of three.[40] Another paper by the author
last named,[41] and two by Mr. Lydekker[42] should be consulted by those
interested in the details of the subject, while an admirable summary will
be found in Professor Newton's _Dictionary of Birds_. In a review of Dr.
Ameghino's paper on these birds,[43] Mr. C. W. Andrews stated that
_Phororhachos_ and others of the "Stereornithes" were not truly Ratite, but
were Carinate forms in which the wings had undergone reduction, and
suggested that possibly they were related to the parent stock of the
Gruiformes, approximating particularly to _Cariama_ (_Dicholophus_).
Shortly afterwards Dr. Ameghino's collection was acquired by the British
Museum, and a study of the specimens themselves has not caused the reviewer
to change his opinion.[44] Some members of the group (e.g. _Mesembriornis_)
are perhaps truly Ratite, and one at least (_Dryornis_) belongs to the
_Cathartidae_. _Phororhachos_ is remarkable for the immense size and heavy
build of the skull, to which the legs, huge though they sometimes are, bear
no proportion; the maxilla is exceedingly compressed, yet very deep, and
ends in a strong hook, while the long massive mandible curves upwards to
meet it. There is a quite or nearly complete interorbital septum in this
case, as opposed to _Apteryx_, and, to a considerable extent, to the
Dinornithidae; {45}while the nostrils are pervious, and the quadrate
articulates with the skull by two heads, contrary to what occurs in the
Ratitae proper. The furcula is existent, but extremely slender; the
metatarsus is more or less elongated, the hallux is present, the wings are
small but well developed, and the tail is said to be long, with a
considerable number of separate vertebrae.
This genus includes the species _P. longissimus_, _P. inflatus_, _P.
platygnathus_, _P. modicus_, _P. gracilis_, and _P. sehuensis_;
_Brontornis_, which has a shorter and wider mandible and smaller but
stouter metatarsi, possesses in _B. burmeisteri_ a form as large as
_Aepyornis maximus_, while _Opisthodactylus_ and other proposed genera are
too imperfectly known to deserve consideration in our limited space.
Besides the above, Dr. Gadow classed with the Stereornithes, _Diatryma_ of
New Mexico, known from a metatarsus; _Dasornis_ of the London Clay,
described from fragments of a skull; _Remiornis_ from the neighbourhood of
Rheims, of which several imperfect bones have been found; and _Gastornis_
of both England and France, of which a fair number of parts have been
unearthed. All occur in the Eocene, but the question of their relationship
is by no means settled, and some writers consider _Gastornis_ to be nearly
allied to the Anseres. This form appears to have been of the size of an
Ostrich, with long leg-bones and short weak wings, and was probably
flightless. Three species have been propounded, _G. parisiensis_, _G.
klaasseni_, and _G. edwardsi_.
* * * * *
(B) With regard to the difficult question of the position in the system of
the NEORNITHES ODONTOLCAE, a few introductory words of explanation are
necessary. In 1872 Professor Marsh bestowed upon two fossils from the
Cretaceous deposits of Kansas the names of _Hesperornis_ and _Ichthyornis_,
which he proposed in the following year[45] to comprise in a Sub-class
_Odontornithes_, so called from the presence of teeth in the jaws.
Subsequently[46] he divided this Sub-class into two Orders, _Odontolcae_
and _Odontotormae_, the former containing _Hesperornis_, with the teeth
arranged in grooves, and the latter _Ichthyornis_, where they were placed
in distinct sockets. His views have been controverted by many writers, but
Mr. Lydekker–an authority of great weight in this connexion–while fully
admitting the affinity of the first form to {46}the Divers, and the
resemblance of the second to the Gull-tribe, proposed in 1891[47] to retain
the term Odontornithes for a series of birds ancestral to the modern series
of toothless Carinatae, for which he adopted the title _Euornithes_, used
in a narrower sense by Dr. Stejneger. It has, however, been decided to
follow Dr. Gadow on this point; while the marks of distinction given below
make it seem at least probable that, whereas _Ichthyornis_ may be referred
to the Carinate division, _Hesperornis_ should be placed in closer
proximity to the Ratite forms. Our Neornithes Odontolcae consequently
contain the HESPERORNITHES, the ENALIORNITHES, and _Baptornis_, all of
which appear to be nearly related.
[Illustration: FIG. 13.–Restoration of _Hesperornis_. (From Huxley, after
Marsh.) × 1/13.]
_Hesperornis regalis_, which stood about three feet high, and _H.
crassipes_, of even larger dimensions, had blunt teeth in the {47}grooves
of both maxilla and mandible, the number being thirty or more below, but
considerably less above, where they did not reach to the anterior
extremity. The bill was long and pointed, the rami of the lower jaw being
entirely separate; the head was rather small, the neck was long, and the
quadrate bone articulated with the skull by one knob only. The sternum was
long, broad, and flat, without keel; the furcula was decidedly reduced, the
metatarsus was moderate and laterally compressed; there were four toes, all
directed forwards and probably webbed; the wing was rudimentary, being
little more than a humerus; the tail was fairly long and broad, but had no
pygostyle. _Enaliornis barretti_ and _E. sedgwicki_ of the Cambridge
Greensand had leg-bones very similar to the above, but being only known
from fragmentary remains, their position is uncertain; while the same may
be said of _Baptornis_ of the North American Cretaceous strata, which, like
the two last-named, is much smaller than _Hesperornis_.
{48}CHAPTER III
NEORNITHES CARINATAE
BRIGADE I–LEGION I (COLYMBOMORPHÆ). ORDERS:
ICHTHYORNITHES–COLYMBIFORMES–SPHENISCIFORMES–
PROCELLARIIFORMES
(C) The NEORNITHES CARINATAE, or birds which, with few exceptions, have a
keel to the sternum, include all the remaining members of the Class. It is
unnecessary to recapitulate the distinctions between these forms and the
Ratitae, to be found on p. 26, but it may be well to reiterate that it is
the sum of all the characters that constitutes the difference, and to point
out that in one or more of the items several of the Carinatae agree with
the members of the aforesaid group, though totally at variance with them in
the aggregate. Claws on the manus are found on the pollex and index in
certain of the Anseres, Cathartae, and Accipitres, and on the pollex alone
in some Anseres, Accipitres, and Galli, with individual instances in other
birds.
[Illustration: FIG. 14.–Head of _Ichthyornis_. (From Geikie, after Marsh.)
× ½.]
ORDER I. ICHTHYORNITHES.
Enough has already been said with regard to the position of the Order
Ichthyornithes, with its sole Family ICHTHYORNITHIDAE; but it remains to
discuss the several members. _Ichthyornis victor_, _I. dispar_, and the
other species were small forms of about the size of a Partridge, with the
habits and appearance, it is presumed, of Terns or Gulls.[48] The head was
extremely large {49}in proportion to the remainder of the skeleton; the
beak was long and pointed, with entirely separate rami to the mandible; the
sharp teeth, fixed regularly in distinct sockets, were inclined backwards,
and occupied the whole of the lower and at least the posterior half of the
upper jaw; the keel of the sternum was large and broad; the dorsal and
cervico-dorsal vertebrae were biconcave, as in _Archaeopteryx_, and perhaps
to some extent in _Enaliornis_; the quadrate articulated to the skull by
one knob, as in the Neornithes Ratitae and Neornithes Odontolcae; the
metatarsus was short and the whole foot small; a furcula was probably
present; the wings were well developed, indicating great powers of flight;
while the tail was comparatively short, and ended in a pygostyle. It will
be observed that of these characters the formation of the jaw and its
teeth, the biconcave vertebrae, and the articulation of the quadrate, are
those that chiefly distinguish the Order from the rest of the Carinatae.
_Apatornis celer_, also from the Cretaceous deposits of Kansas, is probably
to be placed here, but other genera described from the same strata cannot
yet be certainly classified.[49]
ORDER II. COLYMBIFORMES.
The Colymbiformes constitute a very archaic Order of Birds, and hold a
somewhat isolated position. Older writers combined them with the Alcidae as
a group Pygopodes, but recent anatomical investigations make it clear that
Auks have more affinity to Gulls, which again trend to the Limicoline
alliance. As regards structure, the two Sub-Orders COLYMBI and PODICIPEDES,
with their Families _Colymbidae_, or Divers, and _Podicipedidae_, or
Grebes, may be here treated together. They are all water-birds with webbed
or lobed toes and extraordinarily flattened metatarsi. The sternum in the
Colymbidae is much longer than broad, in the Podicipedidae short and wide,
while the furcula is Y-shaped; the neck is more or less elongated; the bill
in the former Family is strong, straight, acute, and compressed, in the
latter moderate and sometimes recurved, being either slender, as in
_Aechmophorus_, or very stout, as in _Podilymbus_. The scutellated
metatarsi are set very far back, and are fairly long, the procnemial
process of the tibia being remarkably elongated, though Grebes alone have a
distinct patella; the hallux is very small and has a small membrane,
{50}but whereas Divers have the anterior toes fully webbed, their allies
have them surrounded by large lobes of skin, connected only at the base.
The claws are abnormally broad and flat in Grebes, the outer margin of the
third being serrated. In the Colymbidae the wing is short, narrow, and
pointed, with eleven primaries and about twenty secondaries; in the
Podicipedidae it is still shorter and concave in form, with twelve
primaries but rarely twenty secondaries; in the latter no true rectrices
can be distinguished, though a tuft of downy feathers exists, while in the
former they are normal though much reduced, and number from eighteen to
twenty. Grebes have bare lores, and are frequently adorned in the breeding
season with crests or tippets of a golden or brownish colour; the dense
glossy plumage being more commonly used for decorative purposes than the
duller coats of Divers. The tongue is always long and pointed, the syrinx
is tracheo-bronchial, the nostrils are pervious, an aftershaft is present,
and both adults and young are uniformly downy. Fossil remains from the
Oligocene of France and southern England, indicating a genus intermediate
between the two Families, have been named _Colymboïdes_.[50]
Fam. I. COLYMBIDAE.–_Colymbus septentrionalis_, the Red-throated Diver of
the Arctic and sub-Arctic parts of both worlds, is brownish black in
summer, with white under-parts and white specks above; the head and neck
are lead-coloured, except the nape, which is black with white streaks, and
the mid-throat, which is reddish-chestnut. _C. arcticus_, the
Black-throated Diver, found in the same regions though with a different
distribution, as for instance in Scotland, is blacker, with white bars as
well as spots; the crown and hind neck being ashy grey, the sides of the
latter striped with black and white, and the throat purplish-black,
interrupted by a semi-collar of white with vertical black lines. _C.
pacificus_ of western North America is barely separable. _C. glacialis_,
the Great Northern Diver, has a much more restricted range, breeding in
Iceland, Greenland, and the Fur Countries as far west as the Great Slave
Lake, where it meets _C. adamsi_ (hardly differing except in the
yellowish-white bill), which extends thence to Northern Asia, and possibly
to Spitsbergen and Jan Mayen. The former is black above, with belts of
white spots making a "chess-board" pattern; the lower surface is {51}white,
and the throat is crossed by two bands of white with longitudinal black
bars, while the head and neck are black with a purplish gloss, changing to
green below. In winter most Divers are found down to the northern tropic,
at which season the throat becomes white, as it is in the young, in which
the feathers of the upper parts are duller with whitish edges. The sexes
are similar; the bill is normally black, and the feet are bluish or
greenish grey. The downy chicks are sooty above.
[Illustration: FIG. 15.–Great Northern Diver. _Colymbus glacialis._ × ⅛.]
Divers are not usually gregarious, and unless driven by stormy weather to
inland waters, are essentially marine, except during the breeding season,
when they ascend the rivers and seek their customary nesting-sites on the
moors, the Black-throated species showing a somewhat greater preference
than the rest for islands in the lakes they frequent, but the Red-throated
often selecting small pools, or even "flows," among the heather. The two
eggs, greenish- or reddish-brown in hue, with blackish and grey blotches
and spots, are laid on a mere depression in the grass or sand close to the
water's edge, or upon a mass of green vegetation which is occasionally
semi-natant. Incubation is said to last four weeks. As a rule the female
performs this duty, lying flat upon her eggs, and gliding or scrambling off
when disturbed, whence a distinct track is often visible upon the turf. On
leaving the land a dive is taken {52}to a considerable distance, then both
parents swim towards the intruder with the body partly submerged, and
finally, if thoroughly scared, they rise heavily on the wing to circle
round with outstretched neck before betaking themselves with rapid but
laboured flight to some neighbouring lake, from which they return at
intervals until the coast is clear. They descend from aloft noisily and
with great impetus, the splashing plunge being followed by a gliding
movement, leaving a broad furrow behind, while on land they move with
difficulty, and rest on the metatarsus. Their croak, or loud, clear,
melancholy cry is often heard before storms, whence the Red-throated Diver
is called Rain-goose in Scotland; the food consists chiefly of fish,
brought to the surface and swallowed with a jerk, but crustaceans,
molluscs, and perhaps aquatic insects vary the diet. The young take to the
water readily, but the female occasionally carries them on her back.
Both Divers and Grebes swim strongly, the flat of the metatarsus meeting
the water during the back stroke, and the thin edge on the return. When
submerged they do not use the pinions.
[Illustration: FIG. 16.–Little Grebe. _Podicipes fluviatilis._ × ¼.]
Fam. II. In the PODICIPEDIDAE both sexes are mainly dusky brown or blackish
grey above, and silvery white below, often with some white on the wing; so
it will only be necessary to note hereafter the distinctive ornaments or
bright colours which are invariably lost in winter. _Podicipes
fluviatilis_, the Little Grebe or Dabchick, ranging over Europe, Africa,
and Asia to the Malay Countries and North Australia, has rich chestnut
cheeks, throat, and sides of the neck, horn-coloured bill, and greenish
feet. In winter the chestnut fades to buff with a white chin. Count
Salvadori[51] considers _P. gularis_ of Australia and Papuasia and {53}_P.
tricolor_ of the Moluccas separable, _P. pelzelni_ of Madagascar being
hardly so. _P. dominicus_, extending from the southern United States to
Patagonia, differs in its black throat. The Little Grebe breeds commonly in
Britain, while _P. cristatus_, the Great Crested Grebe or Loon, only nests
on our largest waters, covering, however, a wide range in Europe, Africa,
Asia, Australia, and New Zealand. It has a bifurcate crest of brown, a
chestnut ruff tipped with black round the cheeks and throat, a red base to
the bill and greenish feet. _P. griseigena_, the Red-necked Grebe, which
wanders to our shores, but breeds in the north of the Palaearctic and
Nearctic Regions, and perhaps occasionally in Morocco, has the foreneck
chestnut, a line above the cheeks white, and the base of the bill yellow.
Some writers denominate the North American and East Asiatic form, _P.
holboelli_. _P. auritus_, the Slavonian Grebe of the sub-Arctic portions of
both worlds, has a tuft of golden chestnut feathers on each side of the
head, an ample black ruff, rufous chest and flanks, black bill and greenish
feet; _P. nigricollis_, the Eared Grebe, of Central and Southern Europe,
Africa, temperate Asia, and western North America, has merely golden
ear-tufts, with a black chest. Both visit us at certain seasons. Finally,
_P. nestor_ inhabits South Australia; _P. rufipectus_ New Zealand; _P.
caliparaeus_, _P. rollandi_, and _Aechmophorus major_ America south of Peru
and Brazil; _Ae. occidentalis_ western North America; _Podilymbus podiceps_
nearly all the New World: and _Centropelma micropterum_ Lake Titicaca only.
The first two have white hair-like filaments on the head, the third and
fourth elongated ear-coverts of golden brown or black and white; while
_Podilymbus_ is remarkable for its stout whitish bill with median black
band and its black throat, _Centropelma_ for its aborted wings and
flightless condition. _Podicipes taczanowskii_, of Lake Junin in Peru,
differs from _P. caliparaeus_ in its longer and lighter bill and feet, and
grey-brown ear-coverts. Grebes in the down are streaked with white or buff
on a dusky ground, while some have a naked red space on the crown.
These migratory birds frequent reedy streams and stagnant waters in summer,
being companionable, though not gregarious; hard weather, however, drives
them to the sea. They walk fairly well, though awkwardly, and sit upon the
whole metatarsus; but the chicks progress on "all fours," using the wings
almost {54}as forefeet.[52] They fly straight and rapidly, with head and
feet extended, but have difficulty in leaving the water; they dive at the
slightest alarm, their quick sight enabling them to vanish below the
surface at the flash of a gun, to reappear, with hardly a ripple, at a
distance. Frequently it requires much patience to obtain a second view, as
their bodies can be submerged to any extent, and at times the bill alone is
exposed. In swimming they jerk the head and often rise vertically to shake
their wings. They descend from the air with a splash and a glide, while in
diving the feet alone act as oars, the young soon equalling their parents
in this respect. The note is a harsh croak in the larger forms, a softer
sound or whit-whit in the smaller; the food consists of fish when
procurable, but small reptiles, amphibians, molluscs, crustaceans, insects,
and vegetable matter are frequently added, and feathers of some size are
constantly found in the stomach. The nest, a pile of aquatic weeds or
rushes of varying bulk, is fixed among reeds, sedges, semi-natant masses of
herbage, or, more rarely, upon low branches of trees or bushes verging upon
the water. Should this rise higher, fresh materials are added. From three
to six bluish-white eggs with a smooth chalky covering are laid in a slight
depression above, but being covered with wet weeds by the female on
leaving, soon become stained with brown. The bill is used in concealing
them, nor does an invader's presence usually hinder the operation.
Incubation lasts from twenty-one to twenty-four days. Both sexes are said
to assist, and the mother carries the nestlings on her back, or even dives
with them in that position.
ORDER III. SPHENISCIFORMES.
The Order Sphenisciformes, with its Sub-Order SPHENISCI, contains only
those remarkable marine birds the Penguins (Fam. SPHENISCIDAE), the life of
which is chiefly spent on the stormy waters of the Antarctic seas. Coupled
by former writers with the Auks, their northern analogues, it has now been
shown that the slight external similarity of the two groups is utterly
misleading, the nearest allies of the primitive forms here treated being
the Petrels on the one hand and the Divers and Grebes on the other. Their
unique structure is correlated with very peculiar habits.
{55}The horny sheath of the maxilla is composed of from three to five more
or less distinct pieces, while the powerful bill may be long, thin, and
slightly decurved, as in _Aptenodytes_ and _Pygosceles_; shorter and pretty
broad, as in _Eudyptes_; or very stout, short, and compressed, as in
_Spheniscus_, where the prominent hook of the culmen overhangs a truncated
mandible. The three metatarsals are not completely fused as in other birds
(p. 10), the scutellated metatarsus itself being shorter and broader than
in any other Family, except the Fregatidae; the legs are set far back, the
tibia is hardly visible, and the short thick toes are directed forwards,
the small hallux alone having no web. Even more striking are the wings,
which are totally devoid of normally-developed quills, though the number of
feathers is very large, the primaries themselves amounting to about
thirty-six; these flippers or paddles have highly compressed bones with no
power of flexure, but work freely from the shoulder in rotatory fashion,
requiring a corresponding increase of strength in the muscles of the
neighbouring parts. The numerous rectrices are fairly long and stiff in
_Aptenodytes_, _Pygosceles_, and _Eudyptes_, but shorter in _Spheniscus_,
having considerably reduced vanes. On the body we find no naked tracts, but
a uniform covering of small scale-like feathers, with or without barbs, and
an equally uniform distribution of down both in adults and young; the
moult, moreover, is accomplished in an exceptional manner, the plumage
being shed in masses, and that of the wing gradually flaking off above the
new coat. The process apparently occupies about ten days.[53] Long
superciliary crests occur in _Eudyptes_, the mandible is more or less
feathered in _Aptenodytes_ and _Pygosceles_, and the metatarsi are clothed
besides in _A. forsteri_. The furcula is U-shaped, the syrinx
tracheo-bronchial, the tongue rudimentary, an after-shaft is present, and
the plentiful subcutaneous fat produces a marketable oil.