The Works of Francis Balfour 3-20

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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Cephalochorda | Urochorda | Elasmobranchii | Teleostei | Cyclostomata | Ganoidei | Amphibia | Aves | Reptilia | Mammalia | Comparison of the Formation of Germinal Layers and Early Stages in Vertebrate Development | Ancestral form of the Chordata | General Conclusions | Epidermis and Derivatives | The Nervous System | Organs of Vision | Auditory, Olfactory, and Lateral Line Sense Organs | Notochord, Vertebral Column, Ribs, and Sternum | The Skull | Pectoral and Pelvic Girdles and Limb Skeleton | Body Cavity, Vascular System and Glands | The Muscular System | Excretory Organs | Generative Organs and Genital Ducts | The Alimentary Canal and Appendages in Chordata
Online Editor 
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This historic 1885 book edited by Foster and Sedgwick is the third of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
Modern Notes:

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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

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Vol. III. A Treatise on Comparative Embryology 2 (1885)

Chapter XX. The Pectoral and Pelvic Girdles and the Skeleton of the Limbs

The Pectoral Girdle

Pisces. Amongst Fishes the pectoral girdle presents itself in its simplest form in Elasmobranchii, where it consists of a bent band of cartilage on each side of the body, of somewhat variable form, meeting and generally uniting with its fellow ventrally. Its anterior border is in close proximity with the last visceral arch, and a transverse ridge on its outer and posterior border, forming the articular surface for the skeleton of the limb, divides it into a dorsal part, which may be called the scapula, and a ventral part which may be called the coracoid.

In all the remaining groups of Fishes there is added to the cartilaginous band, which may wholly or partially ossify, an osseous support composed of a series of membrane bones.

In the types with such membrane bones the cartilaginous parts do not continue to meet ventrally, except in the Dipnoi where there is a ventral piece of cartilage, distinct from that bearing the articulation of the limb. The cartilage is moreover produced into two ventral processes, an anterior and a posterior, below the articulation of the limb ; which may be called, in accordance with Gegenbaur's nomenclature, the praecoracoid and coracoid. Of these the praecoracoid is far the most prominent, and in the majority of cases the coracoid can hardly be recognised. The coracoid process is however well developed in the Selachioid Ganoids, and the Siluroid Teleostei. In Teleostei the scapular region often ossifies in two parts, the smaller of which is named by Parker praecoracoid, though it is quite distinct from Gegenbaur's praecoracoid. The membrane bones, as they present themselves in their most primitive state in Acipenser and the Siluroids, are dermal scutes embracing the anterior edge of the cartilaginous girdle. In Acipenser there are three scutes on each side. A dorsal scute known as the supra-clavicle, connected above with the skull by the posttemporal ; a middle piece or clavicle, and a ventral or infraclavicle (inter-clavicle), which meets its fellow below.

In most Fishes the primitive dermal scutes have become subdermal membrane bones, and the infra-clavicle is usually not distinct, but the two clavicles form the most important part of the membranous elements of the girdle. Additional membrane bones (post-clavicles) are often present behind the main row.

The development of these parts in Fishes has been but little studied.

In Scyllium, amongst the Elasmobranchii, I find that each half of the pectoral girdle develops as a vertical bar of cartilage at the front border of the rudimentary fin, and externally to the muscle-plates.

Before the tissue forming the pectoral girdle has acquired the character of true cartilage, the bars of the two sides meet ventrally by a differentiation in situ of the mesoblastic cells, so that, when the girdle is converted into cartilage, it forms an undivided arc, girthing the ventral side of the body. There is developed in continuity with the posterior border of this arc on the level of the fin a horizontal bar of cartilage, which is continued backwards along the insertion of the fin, and, as will be shewn in the sequel, becomes the metapterygium of the adult (figs. 344, bp and 348, mp). With this bar the remaining skeletal elements of the fin are also continuous.

The foramina of the pectoral girdle are not in the first instance formed by absorption, but by the non-development of the cartilage in the region of pre-existing nerves and vessels.

The development of these parts in Teleostei has been recently investigated by 'Swirski (No. 472) who finds in the Pike (Esox) that the cartilaginous pectoral girdle is at first continuous with the skeleton of the fin. It forms a rod with a dorsal scapular and ventral coracoid process. An independent mass of cartilage gives rise to a prascoracoid, which unites with the main mass, forming a triradiate bar like that of Acipenser or the Siluroids. The coracoid process becomes in the course of development gradually reduced.

'Swirski concludes that the so-called praecoracoid bar is to some extent a secondary element, and that the coracoid bar corresponds to the whole of the ventral part of the girdle of Elasmobranchii, but his investigations do not appear to me to be as complete as is desirable.

Amphibia and Amniota. The pectoral girdle contains a more or less constant series of elements throughout the Amphibia and Amniota ; and the differences in structure between the shoulder girdle of these groups and that of Fishes are so great that it is only possible to make certain general statements respecting the homologies of the parts in the two sets of types.

The generally accepted view, founded on the researches of Parker, Huxley, and Gegenbaur, is to the effect that there is a primitively cartilaginous coraco-scapular plate, homologous with that in Fishes, and that the membrane bones in Fishes are represented by the clavicle and inter-clavicle in the Sauropsida and Mammalia, which are however usually admitted to be absent in Amphibia. These views have recently been challenged by Gotte (No. 466) and Hoffmann (No. 467), on the ground of a series of careful embryological observations ; and until the whole subject has been worked over by other observers it does not seem possible to decide satisfactorily between the conflicting views. It is on all hands admitted that the scapulo-coracoid elements of the shoulder girdle are formed as a pair of cartilaginous plates, one on each side of the body. The dorsal half of each plate becomes the scapula, which may subsequently become divided into a supra-scapula and scapula proper ; while the ventral half forms the coracoid, which is not always separated from the scapula, and is usually divided into a coracoid proper, a praecoracoid, and an epicoracoid. By the conversion of parts of the primitive cartilaginous plates into membranous tissue various fenestrae may be formed in the cartilage, and the bars bounding these fenestrae both in the scapula and coracoid regions have received special names ; the anterior bar of the coracoid region, forming the praecoracoid, being especially important. At the boundary between the scapula and the coracoid, on the hinder border of the plate, is placed the glenoid articular cavity to carry the head of the humerus.

The grounds of difference between Gotte and Hoffmann and other anatomists concern especially the clavicle and inter-clavicle. The clavicle is usually regarded as a membrane bone which may become to some extent cartilaginous. By. the above anatomists, and by Rathke also, it is held to be at first united with the coraco-scapular plate, of which it forms the anterior limb, free ventrally, but united dorsally with the main part of the plate ; and Gotte and Hoffmann hold that it is essentially a cartilage bone, which however in the majority of the Reptilia ossifies directly without passing through the condition of cartilage.

The interclavicle (episternum) is held by Gotte to be developed from a paired formation at the free ventral ends of the clavicles, but he holds views which are in many respects original as to its homologies in Mammalia and Amphibia. Even if Gotte's facts are admitted, it does not appear to me necessarily to follow that his deductions are correct. The most important of these is to the effect that the dermal clavicle of Pisces has no homologue in the higher types. Granting that the clavicle in these groups is in its first stage continuous with the coracoscapular plate, and that it may become in some forms cartilaginous before ossifying, yet it seems to me all the same quite possible that it is genetically derived from the clavicle of Pisces, but that it has to a great extent lost even in development its primitive characters, though these characters are still partially indicated in the fact that it usually ossifies very early and partially at least as a membrane bone 1 .

In treating the development of the pectoral girdle systematically it will be convenient to begin with the Amniota, which may be considered to fix the nomenclature of the elements of the shoulder girdle.

1 The fact of the clavicle going out of its way, so to speak, to become cartilaginous before being ossified, may perhaps be explained by supposing that its close connection with the other parts of the shoulder girdle has caused, by a kind of infection, a change in its histological characters.

Lacertilia. The shoulder girdle is formed as two membranous plates, from the dorsal part of the anterior border of each of which a bar projects (Rathke, Gotte), which is free at its ventral end. This bar, which is usually (Gegenbaur, Parker) held to be independent of the remaining part of the shoulder girdle, gives rise to the clavicle and interclavicle. The scapulocoracoid plate soon becomes cartilaginous, while at the same time the clavicular bar ossifies directly from the membranous state. The ventral ends of the two clavicular bars enlarge to form two longitudinally placed plates, which unite together and ossify as the interclavicle.

Parker gives a very different account of the interclavicle in Anguis. He states that it is formed of two pairs of bones 'strapped on to the antero-inferior part of the prassternum,' which subsequently unite into one.

Chelonia. The shoulder girdle of the Chelonia is formed (Rathke) of a triradiate cartilage on each side, with one dorsal and two ventral limbs. It is admitted on all hands that the dorsal limb is the scapular element, and the posterior ventral limb the coracoid ; but, while the anterior ventral limb is usually held to be the praecoracoid, Gotte and Hoffmann maintain that, in spite of its being formed of cartilage, it is homologous with the anterior bar of the primitive shoulder-plates of Lacertilia, and therefore the homologue of the clavicle.

Parker and Huxley (doubtfully) hold that the three anterior elements of the ventral plastron (entoplastron and epiplastra) are homologous with the interclavicle and clavicles, but considering that these plates appear to belong to a secondary system of dermal ossifications peculiar to the Chelonia, this homology does not appear to me probable.

Aves. There are very great differences of view as to the development of the pectoral arch of Aves.

About the presence in typical forms of the coraco-scapular plate and two independent clavicular bars all authors are agreed. With reference to the clavicle and interclavicle Parker (No. 468) finds that the scapular end of the clavicle attaches itself to and ossifies a mass of cartilage, which he regards as the mesoscapula, while the interclavicle is formed of a mass of tissue between the ends of the clavicles where they meet ventrally, which becomes the dilated plate at their junction.

Gegenbaur holds that the two primitive clavicular bars are simply clavicles, without any element of the scapula ; and states that the clavicles are not entirely ossified from membrane, but that a delicate band of cartilage precedes the osseous bars. He finds no interclavicle.

Gotte and Rathke both state that the clavicle is at first continuous with the coraco-scapular plate, but becomes early separated, and ossifies entirely as a membrane bone. Gotte further states that the interclavicles are formed as outgrowths of the median ends of the clavicles, which extend themselves at an early period of development along the inner edges of the two halves of the sternum. They soon separate from the clavicles, which subsequently meet to form the furculum ; while the interclavicular rudiments give rise, on the junction of the two halves of the sternum, to its keel, and to the ligament connecting the furculum with the sternum. The observations of Gotte, which tend to shew the keel of the sternum is really an interclavicle, appear to me of great importance.

A prascoracoid, partially separated from the coracoid by a space, is present in Struthio. It is formed by a fenestration of a primitively continuous cartilaginous coracoid plate (Hoffmann). In Dromaeus and Casuarius clavicles are present (fused with the scapula in the adult Dromaeus), though absent in other Ratitae (Parker, etc.).

Mammalia. The coracoid element of the coraco-scapular plate is much reduced in Mammalia, forming at most a simple process (except in the Ornithodelphia) which ossifies however separately 1 .

With reference to the clavicles the same divergencies of opinion met with in other types are found here also.

The clavicle is stated by Rathke to be at first continuous with the coracoscapular plate. It is however soon separated, and ossifies very early, in the human embryo before any other bone. Gegenbaur however shewed that the human clavicle is provided with a central axis of cartilage, and this observation has been confirmed by Kolliker, and extended to other Mammalia by Gotte. The mode of ossification is nevertheless in many respects intermediate between that of a true cartilage bone and a membrane bone. The ends of the clavicles remain for some time, or even permanently, cartilaginous, and have been interpreted by Parker, it appears to me on hardly sufficient grounds, as parts of the mesoscapula and praecoracoid. Parker's so-called mesoscapula may ossify separately. The homologies of the episternum are much disputed. Gotte, who has worked out the development of the parts more fully than any other anatomist, finds that paired interclavicular elements grow out backwards from the ventral ends of the clavicles, and uniting together form a somewhat T-shaped interclavicle overlying the front end of the sternum. This condition is permanent in the Ornithodelphia, except that the anterior part of the sternum undergoes atrophy. But in the higher forms the interclavicle becomes almost at once divided into three parts, of which the two lateral remain distinct, while the median element fuses with the subjacent part of the sternum and constitutes with it the presternum (manubrium sterni). If Gotte' s facts are to be trusted, and they have been to a large extent confirmed by Hoffmann, his homologies appear to be satisfactorily established. As mentioned on p. 563 Ruge (No. 438) holds that Gotte is mistaken as to the origin of the presternum.

Gegenbaur admits the lateral elements as parts of the interclavicle, while Parker holds that they are not parts of an interclavicle but are homologous with the omosternum of the Frog, which is however held by Gotte to be a true interclavicle.

1 This process, known as the coracoid process, is held by Sabatier to be the pnecoracoid ; while this author also holds that the upper third of the glenoid cavity, which ossifies by a special nucleus, is the true coracoid. The absence of a praecoracoid in the Ornithodelphia is to my mind a serious difficulty in the way of Sabatier's view.

Amphibia. In Amphibia the two halves of the shoulder girdle are each formed as a continuous plate, the ventral or coracoid part of which is forked, and is composed of a larger posterior and a smaller anterior bar-like process, united dorsally. In the Urodela the two remain permanently free at their ventral ends, but in the Anura they become united, and the space between them then forms a fenestra. The anterior process is usually (Gegenbaur, Parker) regarded as the praecoracoid, but Gotte has pointed out that in its mode of development it strongly resembles the clavicle of the higher forms, and behaves quite differently to the so-called praecoracoid of Lizards. It is however to be noticed that it differs from the clavicle in the fact that it is never segmented off from the coraco-scapular plate, a condition which has its only parallel in the equally doubtful case of the Chelonia. Parker holds that there is no clavicle present in the Amphibia, while Gegenbaur maintains that an ossification which appears in many of the Anura (though not in the Urodela) in the perichondrium on the anterior border of the cartilaginous bar above mentioned is the representative of the clavicle. Gotte's observations on the ossification of this bone throw doubt upon this view of Gegenbaur ; while the fact that the cartilaginous bar may be completely enclosed by the bone in question renders Gegenbaur's view, that there is present both a clavicle and prsecoracoid, highly improbable.

No interclavicle is present in Urodela, but in this group and in a number of the Anura, a process grows out from the end of each of the bars (praecoracoids) which Gotte holds to be the clavicles. The two processes unite in the median line, and give rise in front to the anterior unpaired element of the shoulder girdle (omosternum of Parker). They sometimes overlap the epicoracoids behind, and fusing with them bind them together in the median line. Parker who has described the paired origin of the so-called omosternum, holds that it is not homologous with the interclavicle, but compares it with his omosternum in Mammals.


(463) Bruch. " Ueber die Entwicklung der Clavicula und die Farbe des Blutes. " Zeit.f. wiss. Zool., \\. 1853.

(464) A. Duges. " Recherches sur 1'osteologie et la myologie des Batraciens a leurs differens ages." Memoires des savants etrang. Academic royale des sciences de Finstitut de France^ Vol. vi. 1835.

(465) C. Gegenbaur. Untersuchungen zur vergleichenden Anatomie der Wirbelthiere, 2 Heft. Schultergiirtel der Wirbelthiere. Bmstflosse der Fische. Leipzig, 1865.

(466) A. Gotte. "Beitrage z. vergleich. Morphol. d. Skeletsystems d. Wirbelthiere : Brustbien u. Schultergiirtel." Archivf. mikr, Anat. Vol. xiv. 1877.

(467) C. K. Hoffmann. "Beitrage z. vergleichenden Anatomic d. Wirbelthiere." Niederlandisches Archivf. ZooL,Vol.v. 1879.

(468) W. K. Parker. "A Monograph on the Structure and Development of the Shoulder-girdle and Sternum in the Vertebrata." Ray Society, 1868.

(469) H. Rathke. Ueber die Entwicklung der Schildkrbten. Braunschweig, 1848.

(470) H. Rathke. Ueber den Bau und die Entwicklung des Brustbeins der Saurier, 1853.

(471) A. Sabatier. Comparaison des ceinfures et des membres antMeurs et posttrtturs d. la Serie d. Vertttrh. Montpellier, 1880.

(472) Georg 'Swirski. Untersuch. iib. d. Entwick. d. Schultergiirtels n. d. Skelets d. Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.

Pelvic Girdle

Pisces. The pelvic girdle of Fishes is formed of a cartilaginous band, to the outer and posterior side of which the basal element of the pelvic fin is usually articulated. This articulation divides it into a dorsal iliac, and ventral pubic section. The iliac section never articulates with the vertebral column.

In Elasmobranchii the two girdles unite ventrally, but the iliac section is only slightly developed. In Chimaera there is a well developed iliac process, but the pubic parts of the girdle are only united by connective tissue.

In the cartilaginous Ganoids the pelvic girdle is hardly to be separated from the skeleton of the fin. It is not united with its fellow, and is represented by a plate with slightly developed pubic and iliac processes.

In the Dipnoi there is a simple median cartilage, articulated with the limb, but not provided with an iliac process. In bony Ganoids and Teleostei there is on each side a bone meeting its fellow in the ventral line, which is usually held to be the rudiment of the pelvic girdle ; while Davidoff attempts to shew that it is the basal element of the fin, and that, except in Polypterus, a true pelvic girdle is absent in these types.

From my own observations I find that the mode of development of the pelvic girdle in Scyllium is very similar to that of the pectoral girdle. There is a bar on each side, continuous on its posterior border with the basal element of the fin (figs. 345 and 347). This bar meets and unites with its fellow ventrally before becoming converted into true cartilage, and though the iliac process (il) is never very considerable, yet it is better developed in the embryo than in the adult, and is at first directed nearly horizontally forwards.

Amphibia and Amniota. The primitive cartilaginous pelvic girdle of the higher types exhibits the same division as that of Pisces into a dorsal and a ventral section, which meet to form the articular cavity for the femur, known as the acetabulum. The dorsal section is always single, and is attached by means of rudimentary ribs to the sacral region of the vertebral column, and sometimes to vertebrae of the adjoining lumbar or caudal regions. It always ossifies as the ilium.

The ventral section is usually formed of two more or less separated parts, an anterior which ossifies as the pubis, and a posterior which ossifies as the ischium. The space between them is known as the obturator foramen. In the Amphibia the two parts are not separated, and resemble in this respect the pelvic girdle of Fishes. They generally meet the corresponding elements of the opposite side ventrally, and form a symphysis with them. The symphysis pubis, and symphysis ischii may be continuous (Mammalia, Amphibia).

The observations on the development of the pelvic girdle in the Amphibia and Amniota are nearly as scanty as on those of Fishes.

Amphibia. In the Amphibia (Bunge, No. 473) the two halves of the pelvic girdle are formed as independent masses of cartilage, which subsequently unite in the ventral line.

In the Urodelous Amphibia (Triton) each mass is a simple plate of cartilage divided into a dorsal and ventral section by the acetabulum. The ventral parts, which are not divided into two regions, unite in a symphysis comparatively late.

The dorsal section ossifies as the ilium. The ventral usually contains a single ossification in its posterior part which forms the ischium ; while the anterior part, which may be considered as representing the pubis, usually remains cartilaginous ; though Huxley (No. 475) states that it has a separate centre of ossification in Salamander, which however does not appear to be always present (Bunge). There is a small obturator foramen between the ischium and pubis, which gives passage to the obturator nerve. It is formed by the part of the tissue where the nerve is placed not becoming converted into cartilage.

There is a peculiar cartilage in the ventral median line in front of the pubis, which is developed independently of and much later than the true parts of the pelvic girdle. It may be called the praepubic cartilage.

Reptilia. In Lacertilia the pelvic girdle is formed as a somewhat triradiate mass of cartilage on each side, with a dorsal (iliac) process, and two ventral (pubic and ischiad) processes. The acetabulum is placed on the outer side at the junction of the three processes, each of which may be considered to have a share in forming it. The distal ends of the pubis and ischium are close together when first formed, but subsequently separate. Each of them unites at a late stage with the corresponding process of the opposite side in a ventral symphysis. A centre of ossification appears in each of the three processes of the primitive cartilage.

Aves. In Birds the parts of the pelvic girdle no longer develop as a continuous cartilage (Bunge). Either the pubis may be distinct, or, as in the Uuck, all the elements. The ilium early exhibits a short anterior process, but the pubis and ischium are at first placed with their long axes at right angles to that of the ilium, but gradually become rotated so as to lie parallel with it, their distal ends pointing backwards, and not uniting ventrally excepting in one or two Struthious forms.

Mammalia. In Mammalia the pelvic girdle is formed in cartilage as in the lower forms, but in Man at any rate the pubic part of the cartilage is formed independently of the remainder (Rosenberg). There are the usual three centres of ossification, which unite eventually into a single bone the innominate bone. The pubis and ischium of each side unite with each other ventrally, so as completely to enclose the obturator foramen.

Huxley holds that the so-called marsupial bones of Monotremes and Marsupials, which as shewn by Gegenbaur (No. 474) are performed in cartilage, are homologous with the praepubis of the Urodela ; but considering the great gap between the Urodela and Mammalia this homology can only be regarded as tentative. He further holds that the anterior prolongations of the cartilaginous ventral ends of the pubis of Crocodilia are also structures of the same nature.


(473) A. Bunge. Untersuch. z, Entwick. d. Beckengiirtels d. Amphibien, Reptilien u. Vogel, Inaug. Diss. Dorpat, 1880.

(474) C. Gegenbaur. "Ueber d. Ausschluss des Schambeins von d. Pfanne d. Hiiftgelenkes." Morph. Jahrbuch, Vol. II. 1876.

(475) Th. H. Huxley. "The characters of the Pelvis in Mammalia, etc." Proc. of Roy. Soc., Vol. xxvm. 1879.

(476) A. Sabatier. Comparaison des ceintures et des membres anterieurs et posterieurs dans la Serie d. Vertebrcs. Montpellier, 1880.

Comparison of Pectoral and Pelvic Girdles

Throughout the Vertebrata a more or less complete serial homology may be observed between the pectoral and pelvic girdles.

In the cartilaginous Fishes each girdle consists of a continuous band, a dorsal and ventral part being indicated by the articulation of the fin ; the former being relatively undeveloped in the pelvic girdle, while in the pectoral it may articulate with the vertebral column. In the case of the pectoral girdle secondary membrane bones become added to the primitive cartilage in most Fishes, which are not developed in the case of the pelvic girdle.

In the Amphibia and Amniota the ventral section of each girdle becomes divided into an anterior and a posterior part, the former constituting the praecoracoid and pubis, and the latter the coracoid and ischium ; these parts are however very imperfectly differentiated in the pelvic girdle of the Urodela. The ventral portions of the pelvic girdle usually unite below in a symphysis. They also meet each other ventrally in the case of the pectoral girdle in Amphibia, but in most other types are separated by the sternum, which has no homologue in the pelvic region, unless the praepubic cartilage is to be regarded as such. The dorsal or scapular section of the pectoral girdle remains free ; but that of the pelvic girdle acquires a firm articulation with the vertebral column.

If the clavicle of the higher types is derived from the membrane bones of the pectoral girdle of Fishes, it has no homologue in the pelvic girdle ; but if, as Gotte and Hoffmann suppose, it is a part of the primitive cartilaginous girdle, the ordinary view as to the serial homologies of the ventral sections of the two girdles in the higher types will need to be reconsidered.


It will be convenient to describe in this place not only the development of the skeleton of the limbs but also that of the limbs themselves. The limbs of Fishes are moreover so different from those of the Amphibia and Amniota that the development of the two types of limb may advantageously be treated separately.

In Fishes the first rudiments of the limbs appear as slight longitudinal ridge-like thickenings of the epiblast, which closely resemble the first rudiments of the unpaired fins.

These ridges are two in number on each side, an anterior immediately behind the last visceral fold, and a posterior on the level of the cloaca. In most Fishes they are in no way connected, but in some Elasmobranch embryos, more especially in Torpedo, they are connected together at their first development by a line of columnar epiblast cells 1 . This connecting line of columnar epiblast is a very transitory structure, and after its disappearance the rudimentary fins become more prominent, consisting (fig. 343, &) of a projecting ridge both of epiblast and mesoblast, at the outer edge of which is a fold of epiblast only, which soon reaches considerable dimensions. At a later stage the mesoblast penetrates into this fold and the fin becomes a simple ridge of mesoblast, covered by epiblast. The pectoral fins are usually considerably ahead of the pelvic fins in development.

For the remaining history it is necessary to confine ourselves to Scylliurn as the only type which has been adequately studied.

The direction of the original ridge which connects the two fins of each side is nearly though not quite longitudinal, sloping somewhat obliquely downwards. It thus comes about that the attachment of each pair of limbs is somewhat on a slant, and that the pelvic pair nearly meet each other in the median ventral line a little way behind the anus.

The elongated ridge, forming the rudiment of each fin, gradually projects more and more, and so becomes broader in proportion to its length, but at the same time its actual attachment to the side of the body becomes shortened from behind forwards, so that what was originally the attached border becomes in part converted into the posterior border. This process is much more completely carried out in the case of the pectoral fins than in that of the pelvic, and the changes of form undergone by the pectoral fin in its development may be gathered from figs. 344 and 348.


b. pectoral fin ; ao. dorsal aorta ; cav. cardinal vein ; ua. vitelline artery ; u.v, vitelline vein ; al. duodenum ; /. liver ; sd. opening of segmented duct into the body cavity ; mp. muscle plate ; ;. umbilical canal.

1 I. M. Balfour. Monograph on Elasmobranfh l-'hhes, pp. 1012.

Before proceeding to the development of the skeleton of the fin it may be pointed out that the connection of the two rudimentary fins by a continuous epithelial line suggests the hypothesis that they are the remnants of two continuous lateral fins 1 .

Shortly after the view that the paired fins were remnants of continuous lateral fins had been put forward in my memoir on Elasmobranch Fishes, two very interesting papers were published by Thacker (No. 489) and Mivart (No. 484) advocating this view on the entirely independent grounds of the adult structure of the skeleton of the paired fins in comparison with that of the unpaired fins 2 .

The development of the skeleton has unfortunately not been as yet very fully studied. I have however made some investigations on this subject on Scyllium, and 'Swirski has also made some on the Pike.

In Scyllium the development of both the pectoral and pelvic fins is very similar.

In both fins the skeleton in its earliest stage consists of a bar springing from the posterior side of the pectoral or pelvic girdle, and running backwards parallel to the long axis of the body. The outer side of this bar is continued into a plate which extends into the fin, and which becomes very early segmented into a series of parallel rays at right angles to the longitudinal bar.

1 Both Maclise arid Humphry {Journal of Anat. and Pkys., Vol. v.) had previously suggested that the paired fins were related to the unpaired fins.

2 Davidoff in a Memoir (No. 477) which forms an important contribution to our knowledge of the structure of the pelvic fins has attempted from his observations to deduce certain arguments against the lateral fin theory of the limbs. His main argument is based on the fact that a variable but often considerable number of the spinal nerves in front of the pelvic fin are united, by a longitudinal commissure, with the true plexus of the nerves supplying the fin. From this he concludes that the pelvic fin has shifted its position, and that it may once therefore have been situated close behind the visceral arches. If this is the strongest argument which can be brought against the theory advocated in the text, there is I trust a considerable chance of its being generally accepted. For even granting that Davidoff's deduction from the character of the pelvic plexus is correct, there is, so far as I see, no reason in the nature of the lateral fin theory why the pelvic fins should not have shifted, and on the other hand the longitudinal cord connecting some of the spinal nerves in front of the pelvic fin may have another explanation. It might for instance be a remnant of the time when the pelvic fin had a more elongated form than at present, and accordingly extended further forwards.

In any case our knowledge of the nature and origin of nervous plexuses is far too imperfect to found upon their character such conclusions as those of Davidoff.

In other words, the primitive skeleton of both the fins consists of a longitudinal bar running along the base of the fin,


The skeleton of the fin was still in the condition of embryonic cartilage. b.p. basipterygium (eventual metapterygium) ; fr. fin rays; p.g. pectoral girdle in transverse section; /. foramen in pectoral girdle; pc. wall of peritoneal cavity.

and giving off at right angles series of rays which pass into the fin. The longitudinal bar, which may be called the basipterygium, is moreover continuous in front with the pectoral or pelvic girdle as the case may be.

The primitive skeleton of the pectoral fin is shewn in longitudinal section in fig. 344, and that of the pelvic fin at a slightly later stage in fig. 345.

A transverse section shewing the basipterygium (inpi) of the pectoral fin, and the plate passing from it into the fin, is shewn in fig. 346.

Before proceeding to describe the later history of the two fins it may be well to point out that their embryonic structure completely supports the view which has been arrived at from the consideration of the soft parts of the fin.

My observations shew that the embryonic skeleton of the paired fin consists of a series of parallel rays similar to those of the unpaired fins. These rays support the soft part of the fin which has the form of a longitudinal ridge, and are continuous at their base with a longitudinal bar, which may very probably be due to secondary development. As pointed out by Mivart, a longitudinal bar is also occasionally formed to support the cartilaginous rays of unpaired fins. The longitudinal bar of the paired fins is believed by both Thacker and Mivart to be due to the coalescence of the bases of primitively independent rays, of which they believe the fin to have been originally composed. This view is probable enough in itself, but there is no trace in the embryo of the bar in question being formed by the coalesceace of rays, though the fact of its being perfectly continuous with the bases of the rays is somewhat in favour of this view 1 .

FIG. 345. PELVIC FIN OF A VERY YOUNG FEMALE EMBRYO OF SCYLLIUM STELLARE. bb. basipterygium ; pu. pubic process of pelvic girdle ; il. iliac process of pelvic girdle.

A point may be noticed here which may perhaps appear to be a difficulty, viz. that to a considerable extent in the pectoral, and to some extent in the pelvic fin the embryonic cartilage from which the fin-rays are developed is at first a continuous lamina, which subsequently segments into rays. I am however inclined to regard this merely as a result of the mode of conversion of the indifferent mesoblast into cartilage ; and in any case no conclusion adverse to the above view can be drawn from it, since I find that the rays of the unpaired fin are similarly segmented from a continuous lamina. In all cases the segmentation of the rays is to a large extent completed before the tissue in question is sufficiently differentiated to be called cartilage by an histologist.

Thacker and Mivart both hold that the pectoral and pelvic girdles have been evolved by ventral and dorsal growths of the anterior end of the longitudinal bar supporting the fin-rays.

There is, so far as I see, no theoretical objection to be taken to this view, and the fact of the pectoral and pelvic girdles originating continuously, and long remaining united with the longitudinal bars of their respective fins is in favour of rather than against this view. The same may be said of the fact that the first part of each girdle to be formed is that in the neighbourhood of the longitudinal bar (basipterygium) of the fin, the dorsal and ventral prolongations being subsequent growths.

1 Thacker more especially founds his view on the adult form of the pelvic fins in the cartilaginous Ganoids ; Polyodon, in which the part which constitutes the basal plate in other forms is divided into separate segments, being mainly relied on. It is possible that the segmentation of this plate, as maintained by Gegenbaur and Davidoff, is secondary, but Thacker's view that the segmentation is a primitive character seems to me, in the absence of definite evidence to the reverse, the more natural one.

The later development of the skeleton of the two fins is more conveniently treated separately.

The pelvic fin. The changes in the pelvic fin are comparatively slight. The fin remains through life as a nearly horizontal lateral projection of the body, and the longitudinal bar the basipterygium at its base always remains as such. It is for a considerable period attached to the pelvic girdle, but eventually becomes segmented from it. Of the fin rays the anterior remains directly articulated with the pelvic girdle on the separation of the basipterygium (fig. 347), and the remaining rays finally become segmented from the basipterygium, though they remain articulated with it. They also become to some extent transversely segmented. The posterior end of the basipterygial bar also becomes segmented off as the terminal ray.

FIG. 346. TRANSVERSE SECTION THROUGH THE PECTORAL FIN OF A YOUNG EMBRYO OK SCYLLIUM STELLARE. mpt. basipterygial bar (metapterygium) ; fr. fin ray; m. muscles; hf. horny fibres.

The pelvic fin thus retains in all essential points its primitive arrangement.

The pectoral fin. The earliest stage of the pectoral fin differs from that of the pelvic fin only in minor points, is the same longitudinal or basipterygial bar to which the fin-rays are attached, whose position at the base of the fin is clearly seen in the transverse section (fig. 346, mpf). In front the bar is continuous with the pectoral girdle (figs. 344 and of above, by which the attachment of the pectoral fin to the body wall becomes shortened from behind forwards, the basipterygial bar is gradually rotated outwards, its anterior end remaining attached to the pectoral girdle. In this way this bar comes to form the posterior border of the skeleton of the fin (figs. 348 and 349, mp], constituting what Gegenbaur called the metapterygium, and eventually becomes segmented off from the pectoral girdle, simply articulating with its hinder edge.


bp. basipterygium ; m.o. process of basipterygium continued into clasper; il. iliac process of pectoral girdle ; pit. pubis.

The changes which take place in the course of the further development are however very much more considerable in the case of the pectoral than in that of the pelvic fin.

By the process spoken m p t me tapterygium (basipterygium of earlier stage); me.p. rudiment of future pro- and mesopterygium ; sc. cut surface of scapular process ; cr. coracoid process;/;', foramen;/, horny fibres.

The plate of cartilage, which is continued outwards from the basipterygium, or as we may now call it, the metapterygium, into the fin, is not nearly so completely divided up into fin-rays as in the case of the pelvic fin, and this is especially the case with the basal part of the plate. This basal part becomes in fact at first only divided into two parts (fig. 348) a small anterior part at the front end (me.p), and a larger posterior along the base of the remainder of the fin. The anterior part directly joins the pectoral girdle at its base, resembling in this respect the anterior fin-ray of the pelvic girdle. It constitutes the rudiment of the mesopterygium and propterygium of Gegenbaur. It bears four fin-rays at its extremity, the anterior not being well marked. The remaining fin-rays are borne by the edge of the plate continuous with the metapterygium.

The further changes in the cartilages of the limb are not important, and are easily understood by reference to fig. 349 representing the limb of a nearly full-grown embryo. The front end of the anterior basal cartilage becomes segmented off as a propterygium, bearing a single fin-ray, leaving the remainder of the cartilage as a mesopterygium. The remainder of the now considerably segmented fin-rays are borne by the metapterygium.

The mode of development of the pectoral fin demonstrates that, as supposed by Mivart, the metapterygium is the homologue of the basal cartilage of the pelvic fin.

From the mode of development of the fins of Scyllium conclusions may be drawn adverse to the views recently put forward on the structure of the fin by Gegenbaur and Huxley, both of whom consider the primitive type of fin to be most nearly retained in Ceratodus, and to consist of a central multisegmented axis with numerous rays. Gegenbaur derives the Elasmobranch pectoral fin from a form which he calls the archipterygium, nearly like that of Ceratodus, with a median axis and two rows of rays ; but holds that in addition to the rays attached to the median axis, which are alone found in Ceratodus, there were other rays directly articulated to the shoulder-girdle. He considers that in the Elasmobranch fin the majority of the lateral rays on the posterior (median or inner according to his view of the position of the limb) side have become aborted, and that the central axis is represented by the metapterygium ; while the pro- and mesopterygium and their rays are, he believes, derived from those rays of the archipterygium which originally articulated directly with the shoulder-girdle.

Gegenbaur's view appears to me to be absolutely negatived by the facts of development of the pectoral fin in Scyllium ; not so much because the pectoral fin in this form is necessarily to be regarded as primitive, but because what Gegenbaur holds to be the primitive axis of the biserial fin is demonstrated to be really the base, and it is only in the adult that it is conceivable that a second set of lateral rays could have existed on the posterior side of the metapterygium. If Gegenbaur's view were correct we should expect to find in the embryo, if anywhere, traces of the second set of lateral rays ; but the fact is that, as may easily be seen by an inspection of figs. 344 and 346, such a second set of lateral rays could not possibly have existed in a type . of fin like that found in the embryo 1 . With this view of Gegenbaur's it appears to me that the theory held by this anatomist to the effect that the limbs are modified gill arches also falls ; in that his method of deriving the limbs from gill arches ceases to be admissible, while it is not easy to see how a limb, formed on the type of the embryonic limb of Elasmobranchs, could be derived from a visceral arch with its branchial rays 2 .

Gegenbaur's older view that the Elasmobranch fin retains a primitive uniserial type appears to me to be nearer the truth than his more recent view on this subject ; though I hold that the fundamental point established by the development of these parts in Scyllium is that the posterior border of the adult Elasmobranch fin is the primitive base line, i.e. the line of attachment of the fin to the side of the body.


m.p. metapterygium ; me.p. mesopterygium ; //. propterygium ; cr. coracoid process.

1 If, which I very much doubt, Gegenbaur is right in regarding certain rays found in some Elasmobranch pectoral fins as rudiments of a second set of rays on the posterior side of the metapterygium, these rays will have to be regarded as structures in the act of being evolved, and not as persisting traces of a biserial fin.

2 Some arguments in favour of Gegenbaur's theory adduced by Wiedersheim as a result of his researches on Protopterus are interesting. The attachment which he describes between the external gills and the pectoral girdle is no doubt remarkable, but I would suggest that the observations we have on the vascular supply of these gills demonstrate that this attachment is secondary.

Huxley holds that the mesopterygium is the proximal piece of the axial skeleton of the limb of Ceratodus, and derives the Elasmobranch fin from that of Ceratodus by the shortening of its axis and the coalescence of some of its elements. The secondary character of the mesopterygium, and its total absence in the embryo Scyllium, appears to me as conclusive against Huxley's view, as the character of the embryonic fin is against that of Gegenbaur ; and I should be much more inclined to hold that the fin of Ceratodus has been derived from a fin like that of the Elasmobranchii by a series of steps similar to those which Huxley supposes to have led to the establishment of the Elasmobranch fin, but in exactly the reverse order.

With reference to the development of the pectoral fin in the Teleostei there are some observations of 'Swirski (No. 488) which unfortunately do not throw very much light upon the nature of the limb.

'Swirski finds that in the Pike the skeleton of the limb is formed of a plate of cartilage, continuous with the pectoral girdle ; which soon becomes divided into a proximal and a distal portion. The former is subsequently segmented into five basal rays, and the latter into twelve parts, the number of which subsequently becomes reduced.

These investigations might be regarded as tending to shew that the basipterygium of Elasmobranchii is not represented in Teleostei, owing to the fin rays not having united into a continuous basal bar, but the observations are not sufficiently complete to admit of this conclusion being founded upon them with any certainty.

The ckeiropterygium.

Observations on the early development of the pentadactyloid limbs of the higher Vertebrata are comparatively scanty.

The limbs arise as simple outgrowths of the sides of the body, formed both of epiblast and mesoblast. In the Amniota, at all events, they are processes of a special longitudinal ridge known as the Wolffian ridge. In the Amniota they also bear at their extremity a thickened cap of epiblast, which may be compared with the epiblastic fold at the apex of the Elasmobranch fin.

Both limbs have at first a precisely similar position, both being directed backwards and being parallel to the surface of the body.

In the Urodela (Gotte) the ulnar and fibular sides are primitively dorsal, and the radial and tibial ventral : in Mammalia however Kolliker states that the radial and tibial edges are from the first anterior.

The exact changes of position undergone by the limbs in the course of development are not fully understood. To suit a terrestrial mode of life the flexures of the two limbs become gradually more and more opposite, till in Mammalia the corresponding joints of the two limbs are turned in completely opposite directions.

Within the mesoblast of the limbs a continuous blastema becomes formed, which constitutes the first trace of the skeleton of the limb. The corresponding elements of the two limbs, viz. the humerus and femur, radius and tibia, ulna and fibula, carpal and tarsal bones, metacarpals and metatarsals, and digits, become differentiated within this, by the conversion of definite regions into cartilage, which may either be completely distinct or be at first united. These cartilaginous elements subsequently ossify.

The later development of the parts, more especially of the carpus and tarsus, has been made the subject of considerable study ; and important results have been thereby obtained as to the homology of the various carpal and tarsal bones throughout the Vertebrata ; but this subject is too special to be treated of here. The early development, including the succession of the growth of the different parts, and the extent of continuity primitively obtaining between them, has on the other hand been but little investigated ; recently however the development of the limbs in the Urodela has been worked out in this way by two anatomists, Gotte (No. 482) and Strasser (No. 487), and their results, though not on all points in complete harmony, are of considerable interest, more especially in their bearing on the derivation of the pentadactyloid limb from the piscine fin. Till however further investigations of the same nature have been made upon other types, the conclusions to be drawn from Gotte and Strasser's observations must be regarded as somewhat provisional, the actual interpretation of various ontological processes being very uncertain.

The forms investigated are Triton and Salamandra. We may remind the reader that the hand of the Urodela has four digits, and the foot five, the fifth digit being absent in the hand 1 . In Triton the proximal row of carpal bones consists (using Gegenbaur's nomenclature) of (i) a radiale, and (2 and 3) an intermedium and ulnare, partially united. The distal row is formed of four carpals, of which the first often does not support the first 1 This seems to me clearly to follow from Gotte and Strasser's observations.

metacarpal ; while the second articulates with both the first and second metacarpals. In the foot the proximal row of tarsals consists of a tibiale, an intermedium and a fibulare. The distal row is formed of four tarsals, the first, like that in the hand, often not articulating with the first metatarsal, the second supporting the first and second metatarsals ; and the fourth the fourth and fifth metatarsals.

The mode of development of the hand and foot is almost the same. The most remarkable feature of development is the order of succession of the digits. The two anterior (radial or tibial) are formed in the first instance, and then the third, fourth and fifth in succession.

As to the actual development of the skeleton Strasser, whose observations were made by means of sections, has arrived at the following results.

The humerus with the radius and ulna, and the corresponding parts in the hind limb, are the first parts to be differentiated in the continuous plate of tissue from which the skeleton of the limb is formed. Somewhat later a cartilaginous centre appears at the base of the first and second fingers (which have already appeared as prominences at the end of the limb) in the situation of the permanent second carpal of the distal row of carpals ; and the process of chondrification spreads from this centre into the fingers and into the remainder of the carpus. In this way a continuous carpal plate of cartilage is established, which is on the one hand continuous with the cartilage of the two metacarpals, and on the other with the radius and ulna.

In the cartilage of the carpus two special columns may be noticed, the one on the radial side, most advanced in development, being continuous with the radius ; the other less developed column on the side of the ulna being continuous both with the ulna and with the radius. The ulna and radius are not united with the humerus.

In the further growth the third and fourth digits, and in the foot the fifth digit also, gradually sprout out in succession from the ulnar side of the continuous carpal plate. The carpal plate itself becomes segmented from the radius and ulna, and divided up into the carpal bones.

The original radial column is divided into three elements, a proximal the radiale, a middle element the first carpal, and a distal the second carpal already spoken of. The first carpal is thus situated between the basal cartilage of the second digit and the radiale, and would therefore appear to be the representative of a primitive middle row of carpal bones, of which the centrale is also another representative.

The centrale and intermedium are the middle and proximal products of the segmentation of the ulnar column of the primitive carpus, the distal second carpal being common both to this column and to the radial column.

The ulnar or fibular side of the carpus or tarsus becomes divided into a proximal element the ulnare or fibulare the ulnare remaining partially united with the intermedium. There are also formed from this plate two carpals to articulate with digits 3 and 4 ; while in the foot the corresponding elements articulate respectively with the third digit, and with the fourth and fifth digits.

Gotte, whose observations were made in a somewhat different method to those of Strasser, is at variance with him on several points. He finds that the primitive skeleton of the limb consists of a basal portion, the humerus, continued into a radial and an ulnar ray, which are respectively prolonged into the two first digits. The two rays next coalesce at the base of the fingers to form the carpus, and thus the division of the limb into the brachium, antebrachium and manus is effected.

The ulna, which is primitively prolonged into the second digit, is subsequently separated from it and is prolonged into the third ; from the side of the part of the carpus connecting the ulna with the third digit the fourth digit is eventually budded out, and in the foot the fourth and fifth digits arise from the corresponding region. Each of the three columns connected respectively with the first, second, and third digits becomes divided into three successive carpal bones, so that Gotte holds the skeleton of the hand or foot to be formed of a proximal, a middle, and a distal row of carpal bones each containing potentially three elements. The proximal row is formed of the radiale, intermedium and ulnare ; the middle row of carpal i, the centrale and carpal 4, and the distal of carpal 2 (consisting according to Gotte of two coalesced elements) and carpal 3.

The derivation of the cheiropterygium from the ichthyoptcrygium. All anatomists are agreed that the limbs of the higher Vertebrata are derived from those of Fishes, but the gulf between the two types of limbs is so great that there is room for a very great diversity of opinion as to the mode of evolution of the cheiropterygium. The most important speculations on the subject are those of Gegenbaur and Huxley.

Gegenbaur holds that the cheiropterygium is derived from a uniserial piscine limb, and that it consists of a primitive stem, to which a series of lateral rays are attached on one (the radial) side ; while Huxley holds that the cheiropterygium is derived from a biserial piscine limb by the "lengthening of the axial skeleton, accompanied by the removal of its distal elements further away from the shoulder-girdle and by a diminution in the number of the rays."

Neither of these theories is founded upon ontology, and the only ontological evidence we have which bears on this question is that above recorded with reference to the development of the Urodele limb.

Without holding that this evidence can be considered as in any way conclusive, its tendency would appear to me to be in favour of regarding the cheiropterygium as derived from a uniserial type of fin. The humerus or femur would appear to be the basipterygial bars (metapterygium), which have become directed outwards instead of retaining their original position parallel to the length of the body at the base of the fin. The anterior (proximal) fin-rays and the pro- and mesopterygium must be supposed to have become aborted, while the radius or ulna, and tibia or fibula are two posterior fin-rays (probably each representing several coalesced rays like the pro- and mesopterygium) which support at their distal extremities more numerous fin-rays consisting of the rows of carpal and tarsal bones.

This view of the cheiropterygium corresponds in some respects with that put forward by Gotte as a result of his investigations on the development of the Urodele limbs, though in other respects it is very different. A difficulty of this view is the fact that it involves our supposing that the radial edge of the limb corresponds with the metapterygial edge of the piscine fin. The difficulties of this position have been clearly pointed out by Huxley, but the fact that in the primitive position of the Urodele limbs the radius is ventral and the ulna dorsal shews that this difficulty is not insuperable, in that it is easy to conceive the radial border of the fin to have become rotated from its primitive Elasmobranch position into the vertical position it occupies in the embryos of the Urodela, and then to have been further rotated from this position into that which it occupies in the adult Urodela and in all higher forms.

Bibliography of the Limbs

(477) M. v. Davidoff. "Beitrage z. vergleich. Anat. d. hinteren Gliedmaassen d. Fische I." Morphol. Jahrbuch, Vol. v. 1879.

(478) C. Gegenbaur. Untersuckungen z. vergleich. Anat. d. Wirbelthiere. Leipzig, 1864 5. Erstes Heft. Carpus u. Tarsus. Zweites Heft. Brustflosse d. Fische.

(479) C. Gegenbaur. "Ueb. d. Skelet d. Gliedmaassen d. Wirbelthiere im Allgemeinen u. d. Hintergliedmaassen d. Selachier insbesondere." Jenaische Zeitsckrift, Vol. V. 1870.

(480) C. Gegenbaur. " Ueb. d. Archipterygium." Jenaische Zeitschrift, Vol. vii. 1873.

(481) C. Gegenbaur. "Zur Morphologic d. Gliedmaassen d. Wirbelthiere." Morphologisches Jahrbuch, Vol. II. 1876.

(482) A. Gotte. Ueb. Entivick. u. Regeneration d. Gliedmaassenskelets d. Molche. Leipzig, 1879.

(483) T. H. Huxley. "On Ceratodus Forsteri, with some observations on the classification of Fishes." Proc. Zool. Soc. 1876.

(484) St George Mivart. "On the Fins of Elasmobranchii." Zoological Trans., Vol. x.

(485) A. Rosenberg. "Ueb. d. Entwick. d. Extremitaten-Skelets bei einigen d. Reduction ihrer Gliedmaassen charakterisirten Wirbelthieren." Zeil.f. iviss. Zool., Vol. xxin. 1873.

(486) E. Rosenberg. "Ueb. d. Entwick. d. Wirbelsaule u. d. centrale carpi d. Menschen. " Morphologisches Jahrbuch, Vol. I. 1875.

(487) H. Strasser. "Z. Entwick. d. Extremitatenknorpel bei Salamandern u. Tritonen." Morphologisches Jahrbuch, Vol. V. 1879.

(488) G. 'S wirski. Untersitch. iib. d. Entwick. d. Schultergitrtels u. d. Skelcls d. Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.

(489) J. K. Thacker. "Median and paired fins. A contribution to the history of the Vertebrate limbs." Trans, of the Connecticut Acad., Vol. ill. 1877.

(490) J. K. Thacker. "Ventral fins of Ganoids." Trans, of the Connecticut Acad., Vol. iv. 1877.