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failure. Its author, with a considerable mathematical and mechanical bias, reckoned entirely with the quantity, not with the quality of his units, and relied almost implicitly upon his formulae. It is, however, fair to state that his system was not built entirely upon these muscular variations, but rather upon a more laborious combination of anatomical characters, which were so selected that they presumably could not stand in direct correlation with each other, notably the oil-gland, caeca, carotids, nasal bones and above all, the muscles of the thigh. He was, indeed, the first to show clearly the relationship of the heron-like birds with the Steganopodes; of stork-like birds with the American vultures; the great difference between the latter and the other birds of prey; the connexion of the gulls and auks with the plovers, and that of the sand-grouse with the pigeons—discoveries expressed in the new terms of the orders Ciconiiformes and Charadriiformes. These are instances, now well understood, that almost every organic system, even when studied by itself, may yield valuable indications as to the natural affinities of the various groups of birds. That Garrod has so very much advanced the classification of birds is ultimately due to his comprehensive anatomical knowledge and general insight.

1911 Britannica-Bird-Left thigh-muscles.png
From Newton’s Dictionary of Birds.
Fig. 15.—Left thigh-muscles of a Rail. Outer view after removal of the
Il.fb, ilio-fibularis and Il.tib, ilio-tibialis.
A, Caudal. N, Sciatic nerve.
B, Iliac portion of caud-ilio-femoralis., Ischio-femoralis.
X, Caud-ilio-flexorius. Is.fl, Ischio-fibularis.
Y, “Accessory” portion of the same. Sart. Sartorius.
Pif, Pubischio-femoralis.

To return to these thigh muscles. The most primitive combination, ambiens and A B X Y, is the most common; next follows that of A X Y, meaning the reduction of B, i.e. the iliac portion of the caud-ilio-femoralis; A B X and B X Y are less common; A X and X Y are rare and occur only in smaller groups, as in subfamilies or genera; B X occurs only in Podiceps. But the greatest reduction, with only A remaining, is characteristic of such a heterogeneous assembly as Accipitres, Cypselidae. Trochilidae, Striges and Fregata. This fact alone is sufficient proof that these conditions, or rather reductions, have been acquired independently of the various groups. A B Y, A Y, A B, X Y and B do not occur at all, some of them for obvious reasons. Occasionally there is an instructive progressive evolution expressed in these formula; for instance Phaethon, in various other respects the lowest of the Steganopodes, has A X Y, Sula and Phalacrocorax have A X, Fregata, the most specialized of these birds, has arrived at the reduced formula A. Further, the combinations B X Y and A X Y cannot be derived from each other, but both directly from A B X Y in two different directions. Keeping this in mind, we may fairly conclude that the flamingo with B X Y points to an ancestral condition A B X Y, which is still represented by Platalea and Ibis, whilst the other storks proper have taken a different line, leading to A X Y.

Literature.—Well nigh complete lists of the enormous myological literature are contained in Fürbringer’s Untersuchungen zur Morphologie und Systematik der Vögel, and in Gadow’s vol. Vögel of Bronn’s Klassen und Ordnungen des Tierreichs. Only a few papers and works can be mentioned here, with the remark that few authors have paid attention to the all-important innervation of the muscles. A. Carlsson, Beiträge zur Kenntniss der Anatomie der Schwimmvögel; K. Svensk, Vet. Ak. Handlinger. J. G. No. 3 (1884); A. Alix, Essai sur l’appareil locomoteur des oiseaux (Paris, 1874); H. Gadow, Zur vergl. Anat. der Muskulatur des Beckens und der hinteren Gliedmasse der Ratiten, 4° (Jena, 1880); A. H. Garrod, “On Certain Muscles of the Thigh of Birds and on their value in Classification,” P.Z.S., 1873, pp. 624-644; 1874, pp. 111-123. Other papers by Garrod, 1875, pp. 339-348 (deep planter tendons); 1876, pp. 506-519 (wing-muscles of Passeres), &c.; J. G. de Man, Vergelijkende myologische en neurologische Studien over Amphibien en Vogels (Leiden, 1873), (Corvidae); A. Milne-Edwards, Recherches anatomiques et paléontologiques pour servir à l’histoire des oiseaux fossiles de la France (Paris, 1867-1868), tom. i. pls. ix.-x. (Aquila and Gallus); R. Owen, article “Aves,” Todds’ Cyclopaed. of Anat. and Phys. i. (London, 1835); “On the Anatomy of the Southern Apteryx,” Trans. Zool. Soc., iii., 1849; A. Quennerstedt, “Studier i foglarnas anatomi,” Lunds Univers. Aarsk., ix., 1872 (hind-limb of swimming birds); G. Rolleston, “On the Homologies of Certain Muscles connected with the Shoulder-joint,” Trans. Linn. Soc., xxvi., 1868; R. W. Shufeldt, The Myology of the Raven (London, 1891); M. Watson, “Report on the Anatomy of the Spheniscidae,” Challenger Reports, 1883.

3. Nervous System.

Brain.—The more characteristic features of the bird’s brain show clearly a further development of the reptilian type, not always terminal features in a direct line, but rather side-departures, sometimes even a secondary sinking to a lower level, and in almost every case in a direction away from those fundamentally reptilian lines which have led to the characters typical of, and peculiar to, the mammals.

The forebrain forms the bulk of the whole brain, but the large size of the hemispheres is due to the greater development of the basal and lateral portions (pedunculi cerebri and corpora striata), while the pallium (the portion external to the lateral ventricles) is thin, and restricted to the median side of each hemisphere. As a direct result of this undoubtedly secondary reduction of the pallium—due to the excessive preponderance of the basal and lateral parts—the corpus callosum (i.e. the transverse commissure of the right and left pallium) is in birds reduced to a narrow flat bundle of a few white fibres; it is situated immediately above and behind the much stronger anterior commissure, i.e. the connexion between the corpora striata, or chief remaining part of the hemispheres. Owing to the small size of the olfactory lobes the anterior arms of the latter commissure are wanting. There is very little grey matter in the cortex of the hemispheres, the surface of which is devoid of convolutions, mostly quite smooth; in others, for instance pigeons, fowls and birds of prey, a very slight furrow might be compared with the Sylvian fissure.

The Thalamencephalon is much reduced. The epiphysis, or pineal body, is quite as degenerate as in mammals, although still forming a long stalk as in reptiles. In birds, this stalk consists entirely of blood-vessels, which in the adult enclose no terminal vesicle, and fuse with the membranous linings of the skull. The midbrain is represented chiefly by the optic lobes, the cortex of which alone is homologous with the corpora quadragemina of the mammals. Their transverse dorsal connexion is the posterior commissure; otherwise the whole roof portion of the midbrain is reduced to a thin membrane, continuous with that which covers the Sylvian aqueduct, and this ventricle sends a lateral cavity into each optic lobe, as is the case in reptiles. The right and left lobes themselves are rent asunder (so to speak), so that they are freely visible from above, filling the corners formed by the hemispheres and the cerebellum. The latter is, in comparison with mammals; represented by its middle portion only, the vermis; in a sagittal section it shows an extremely well developed arbor vitae, produced by the transverse, repeated folding of the whole organ. In comparison with reptiles the cerebellum of birds shows high development. Forwards it covers, and has driven asunder, the optic lobes; backwards it hides the much shortened medulla oblongata.

Several futile attempts have been made to draw conclusions as to the intelligence of various birds, from comparison of the weight of the whole brain with that of the body, or the weight of the hemispheres with that of other parts of the central nervous system.

The brachial plexus is formed by four or five of the lowest cervical nerves; the last nerve of this plexus often marks the boundary of the cervical and thoracic vertebrae. The composition of the plexus varies much, not only in different species, but even individually. The most careful observations are those by Fürbringer. The serial number of these nerves depends chiefly upon the length of the neck, the extremes being represented by Cypselus (10th-14th cervical) and Cygnus (22nd-24th), the usual numbers of the common fowl being the 13th-17th nerves.

The Crural Plexus is divided into a crural, ischiadic and pubic portion. The first is generally composed of three nerves, the hindmost of which, the furcalis, issues in most birds between the last two lumbo-sacral vertebrae, and then divides, one half going to the crural, the other to the sciatic portions. The obturatorius nerve invariably comes from the two main stems of the crural. The ischiadic portion consists generally of five or six nerves, which leave the pelvis as one thick system through the ilio-ischiadic foramen. The last nerve which contributes to the ischiadic plexus leaves the spinal column in most birds either between the two primary sacral vertebrae, or just below the hindmost of them, and sends a branch to the pubic portion which is composed of post-ischiadic nerves, partly imbedded in the kidneys, and innervates the ventral muscles between the tail and pubis, together with those of the cloaca and copulatory organs.