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Full text of "The anatomy of the salamander"

THE 

ANATOMY 

OF THE 

SALAMAN D E R 



THE 

ANATOMY 

OF THE 

SALAMANDER 



BY 

ERIC T. B. FRANCIS 

B.SC. LOND.j PH.D. READING 

WITH AN 

HISTORICAL INTRODUCTION BY 

Professor F. J. COLE 

D.SC, F.R.S. 



OXFORD 

AT THE CLARENDON PRESS 
1934 




OXFORD UNIVERSITY PRESS 

AMEN HOUSE, E.C. 4 
LONDON EDINBURGH GLASGOW 
NEW YORK TORONTO MELBOURNE 
CAPETOWN BOMBAY CALCUTTA 

MADRAS SHANGHAI 

HUMPHREY MILFORD 

PUBLISHER TO THE UNIVERSITY 



PRINTED IN GREAT BRITAIN AT THE UNIVERSITY PRESS, OXFORD 
BY JOHN JOHNSON, PRINTER TO THE UNIVERSITY 



PREFACE 

The Salamander is well known to all students of Zoology as the 
animal most commonly dissected to exemplify the structure of the 
Amphibia Urodela. Notwithstanding its general use for teaching 
purposes, the accounts of the anatomy of this animal available in 
text-books are out of date, and even, in some respects, seriously 
misleading. It was with a view to closing this gap in our knowledge 
that the present work was undertaken. Its intention is to provide 
a general account of the anatomy and morphology of a tailed Amphi- 
bian — the Salamander, which shall, within limits, be comparable 
with the work of Ecker and Gaupp on its ecaudate relative — the 
Frog. 

In the main the gross anatomy only has been dealt with, histology 
and embryology being excluded. The practical work has been 
carried out under a Zeiss binocular dissecting microscope, using 
magnifications of x 7-5, and x 20 for the finer details, supplemented 
by a microscopic examination of microtome sections where necessary. 
Any special methods used for the elucidation of particular systems, 
e.g. the lymphatic system, are dealt with in the text. All figures are 
original and have been specially drawn for this work by the author, 
with the following exceptions: Text-figures i, 2, and 3 are taken 
from Willem (1923), Figs. 6^, 66, and 67 have been redrawn (with 
Prof. Hoyer's kind permission) from Hoyer and Udziela, Fig. 53 is 
after Luther, and Fig. 82 has been modified from Bruner. I am 
further indebted to Mr. E. G. Boulenger and the Publication 
Committee of the Zoological Society of London for permission to 
reproduce the coloured plate appearing as the Frontispiece. 

Although this work will probably make its greatest appeal to the 
teacher and advanced student, the requirements of the general stu- 
dent are provided for by means of summaries to the more specialized 
sections, and in the planning of the figures illustrating the more 
important systems, e.g. blood-vessels and cranial nerves. To meet 
the needs of the research worker and comparative anatomist an 
attempt has been made to collate as far as possible all the important 
literature dealing with each subject under review, and to give, when- 
ever possible, tables of the synonyms used by previous authors, 
together with the dates and full details of the papers in question. 

The Bibliography has been compiled with some care, and includes 
not only those publications which have direct reference to the subject- 



vi PREFACE 

matter of this work, but also papers dealing with the Histology, 
Embryology, and Physiology of the type, as well as a certain number 
of works, which, although not dealing specifically with Salamandra, 
have sufficiently important general bearings to justify their admis- 
sion. This list does not pretend to be complete except in so far as 
the anatomy of Salamandra is concerned, and there must inevitably 
be many omissions. Nevertheless it is offered with the confidence 
that it will prove an adequate guide for any future investigation, 
either of Salamandra itself, or of any other closely related Urodele. 
In this connexion I must acknowledge with gratitude the invaluable 
assistance which has been freely given by the Librarian and staff of 
both the Zoological and General Libraries of the British Museum 
(Natural History), of the Science Library, South Kensington, the 
Library of the Royal College of Surgeons of England, and of the 
Royal and Zoological Societies. My importunate demands for books 
have always been met with courtesy, and expert advice in tracking 
down obscure references has been placed ungrudgingly at my disposal. 
To my Tutor, Professor F. J. Cole, F.R.S., in whose laboratory 
the work has been done, my sincerest thanks are due. He is respon- 
sible not only for the original plan of the work, but also for stimu- 
lating its progress in times of doubt and difficulty. Moreover, he has 
closely associated himself with its publication by writing the His- 
torical Introduction. I owe a debt of gratitude to Dr. N. B. Eales, 
whose constructive criticisms have helped to rectify many blemishes, 
for sharing with Professor Cole the arduous task of reading the 
proofs of this work. Their aid has been invaluable. I also desire 
to thank the numerous other Zoologists who have assisted in 
various ways. Among them I would name especially Dr. Chas. H. 
O'Donoghue, of Edinburgh University, who has offered valuable 
suggestions on problems connected with the vascular system, and 
Dr. F. H. Edgeworth, of Bristol, who has very kindly placed at my 
disposal his knowledge of muscle terminology — particularly in 
respect of the hyobranchial apparatus, and finally Professor H. 
Graham Cannon, of Manchester University, who has given very wel- 
come technical advice with regard to the illustrations. I have also to 
acknowledge my indebtedness to the Council of the Royal Society 
and the Research Board of Reading University for generous grants, 
without which the publication of the work in its present form 
would not have been possible. E. T. B. F. 

THE UNIVERSITY, READING. 

Jugust 1933. 



CONTENTS 

SALAMANDRA SALAMANDRA L frontispiece 

HISTORICAL INTRODUCTION, by Professor F. J. Cole, d.sc, f.r.s. 

I. TAXONOMY, LIFE-HISTORY, AND GENERAL 
TAXONOMY 



Characteristics of Class, Order, and Sub-order 

The Family Salamandridae 

The Genus Salamandra 

Salamandra semenovi . 

Salamandra caucasica . 

Salamandra atra 



LIFE-HISTORY 

1. Fecundation and Fertilization ...... 4 

2. Deposition of Larvae ........ 6 

3. Larvae .......... 7 

4. Metamorphosis ......... 7 

5. Habits of the Adult ........ 8 

6. Geographical Distribution ....... 8 

7. Parasites and Diseases ........ 9 

GENERAL LITERATURE, ETC 10 



II. EXTERNAL FEATURES AND VARIETIES 
EXTERNAL FEATURES 



VARIETIES 

1. Forma Typica . 

2. Variety Molleri 

3. Variety Taeniata 



III. THE SKELETON 
THE AXIAL SKELETON 



I. THE VERTEBRAL COLUMN 



I. General ....... 


.16 


2. The Cervical, or 'Atlas' Vertebra 


. 16 


3. The Trunk Vertebrae ..... 


. 18 


4. The Ribs 


. 18 


5. The Sacrum ...... 


. 19 


6. The Caudal Vertebrae .... 


. 20 


46031 






f.../i !•» . ^:\r 



CONTENTS 



II. THE SKULL 

1 . The Skull as a whole . 

2. The Primordial Skeleton 

3. The Investing Bones . 
Summary 

III. THE VISCERAL ARCHES 

1. The Mandibular Arch 

2. The Hyoid Arch 

3. The First Branchial Arch 

4. The Second Branchial Arch 

5. The Copula 

6. The Os triangulare 
Synonyms of the Visceral Skeleton 

THE APPENDICULAR SKELETON 

I. THE PECTORAL GIRDLE AND FORE-LIMB. 

1. The Pectoral Girdle , 

2. The Fore-hmb . 

II. THE PELVIC GIRDLE AND HIND-LIMB. 

I. The Pelvic Girdle 
2. 



The Hind-limb 



32 
34 
35 
35 
35 
35 
36 



37 
40 

44 
45 



IV. THE MUSCLES 
GENERAL LITERATURE .... 



48 



MUSCLES OF THE HEAD 

1. Historical .... 

2. Muscles of the Eye 

3. Muscles of the Fifth Nerve . 

4. Muscles of the Seventh Nerve 

5. Muscles of the Ninth and Tenth Nerves 

6. Muscles of the Head suppHed by Spinal Nerves 



48 
49 

51 
56 
60 

65 



MUSCLES OF THE FORE-LIMB AND SHOULDER 

1. Historical ..... 

2. Shoulder Muscles .... 

3. Muscles of the Arm .... 

4. Extensor Muscles of the Fore-arm and Hand . 

5. Flexor Muscles of the Fore-arm and Hand . 

MUSCLES OF THE TRUNK AND TAIL 

1. Historical ..... 

2. Dorsal Trunk Muscles 

3. Ventral Trunk Muscles 

4. Caudal Muscles .... 



69 

71 
75 
81 
84 



92 
92 
94 
98 



CONTENTS 



MUSCLES OF THE HIND-LIMB AND PELVIS 

1. Historical ..... 

2. Muscles of the Pelvis and Thigh 

3. Extensor Muscles of the Leg and Foot 

4. Flexor Muscles of the Leg and Foot 



• 98 

• 99 
. no 

. 114 



V. THE NERVOUS SYSTEM 

BRAIN AND SPINAL CORD 

1. Historical ...... 

2. Membranes of the Brain and Spinal Cord . 

3. Brain ....... 

4. Spinal Cord ...... 

CRANIAL NERVES 

1. Historical 

2. Description 

N. Olfactorius 

N. Opticus 

Eye-muscle Nerves 

N. Trigeminus 

N. Facialis 

Homology of Chorda tympani 

N. Acusticus 

N. Glossopharyngeus 

N. Vagus 

SPINO-OCCIPITAL NERVES 

SPINAL NERVES 

1. Historical ..... 

2. A Typical Spinal Nerve 

3. The First Spinal Nerve 

4. The Second Spinal Nerve 

5. Brachial Plexus, and Nerves of the Fore-limb 

6. Crural Plexus, and Nerves of the Hind-limb 

SYMPATHETIC 

1. Historical ..... 

2. Description ..... 



123 
123 
128 
132 



132 
134 
134 
134 
135 
138 
144 
146 
149 
150 
151 

156 



156 

157 
159 
161 
163 

170 



179 
180 



VI. THE VASCULAR SYSTEM (Including Lymphatics) 
BLOOD-VASCULAR SYSTEM 



I. HEART. 

1 . Historical 

2. Description 

3. Functional 

4. Pericardium 



185 
186 
191 
193 



CONTENTS 



11. BLOOD-VESSELS. 

1. Historical 

2. General Features 

3. Arterial Arches . 

4. Branches of the Carotid Arch . 

5. Branches of the Systemic Arch 
Branches of the Dorsal Aorta 

6. Branches of the Pulmonary Arch 

7. The Venous System . 
The Venous Trunks . 

8. Description of the Veins 
Vessels of the Brain 

A. Arteries 

B. Veins 
Summary of Vessels of the Brain 

LYMPHATIC SYSTEM 

1. Historical 

2. Technique 

3. Description 

A. Subcutaneous network 

B. Lymphatics of the Limbs 

C. Lymphatics of the Pelvis . 

D. Lymphatics of the Trunk and Viscera 

E. Lymphatics of the Pectoral Region 

F. Lymphatics of the Head . 
G. Lymph hearts, and their Relation to the Blood-vessels 



194 

195 
196 
197 
201 
206 
218 
218 
218 
220 
244 
244 
246 
248 



249 

250 
251 
251 
252 
252 
254 
256 
258 
259 



VII. THE ALIMENTARY TRACT AND ITS APPENDAGES 

TOGETHER WITH THE SPLEEN AND THE 

MESENTERIES 



262 
262 
264 
265 
265 
265 
266 



THE 


ALIMENTARY TRACT 


I. 


General . . . . 


2. 


The Mouth Cavity 


3- 


The Pharynx and Oesophagus 


4- 


The Stomach . 


5- 


The Duodenum 


6. 


The Intestine . 


7. 


The Rectum 



THE GLANDS ASSOCIATED WITH THE ALIMENTARY TR.^CT 

1. The Liver ......... 

2. The Pancreas ......... 

THE SPLEEN 

THE MESENTERIES 

1. The Dorsal Mesentery and its Derivatives . . . . . 

2. The Ligamentum Hepato-entericum . . . . . 



266 
267 

267 



268 
268 



CONTENTS 

3. The Ventral Mesentery 

4. The Ligamentum Hepato-cavo-pulmonale 

5. The Diaphragm 



269 
269 
270 



VIII. THE RESPIRATORY SYSTEM, 
LARYNX 



INCLUDING THE 



I. The Glottis 


. 271 


2. The Laryngo-tracheal chamber 


. 271 


3. The Bronchi ..... 


. 272 


4. The Lungs ..... 


. 272 


Homologies of the Urodele Larynx . 


. 272 


Physiological Considerations . 


• 274 



IX. THE URINO-GENITAL SYSTEM 



GENERAL AND HISTORICAL . 
THE FEMALE SYSTEM 

1 . The Kidneys and Ureters 

2. The Ovaries .... 

3. The Fat-body .... 

4. The Oviducts (Miiller's Ducts) 

5. The Receptaculum Seminis and Cloaca 

6. The Urinary Bladder . 

THE MALE SYSTEM 

1 . The Kidneys and Ureters 

2. The Testes 

3. The Vasa Efferentia, &c. 

4. The Fat-body . 

5. The Cloaca and its Glands 

6. The Urinary Bladder . 



277 



279 

280 
280 
280 
282 
282 



283 
284 
285 
286 
286 
288 



X. THE DUCTLESS GLANDS, ETC. 



r. The Thyroid Gland 


. 289 


2. The Thymus Glands ..... 


. 290 


3. The Parathyroids (Epithelial Bodies) 


. 290 


4. The Ultimo-branchial Body .... 


. 290 


5. Other Glands 


. 291 



XL THE SKIN AND SENSE ORGANS 



THE SKIN 

1. 'General . 

2. Venom Glands 

3. Venom . 



292 
292 
293 



CONTENTS 



THE SENSE ORGANS 

I. THE EAR. 

1. General .... 

2. Skeletal Structures 

3. The Membranous Labyrinth . 

4. Nerve-endings 

5. The Perilymphatic System . 

II. THE EYE. 

1. General . . . . . 

2. Glands of the Orbit . 

3. The Eyelids . . . . 

III. THE NOSE. 

1. General .... 

2. Jacobson's Organ 

3. Opening and Closing Mechanism 

BIBLIOGRAPHY 

INDEX 

ABBREVIATIONS USED IN PLATES 
EXPLANATION OF PLATES . 
PLATES I-XXV . . . . 



294 
294 
296 
298 
299 



300 
301 
302 



• 302 

• 302 

• 303 

• 305 

• 345 

• 367 

• 377 
at end 



HISTORICAL INTRODUCTION 

In a morphological treatise such as the present work it is only neces- 
sary to outline the history of the relevant anatomical and physiolo- 
gical literature. Therefore the numerous medieval legends which 
have condensed around the habits and medical importance of the 
Salamander, and the extensive literature relating to its alleged 
resistance to fire, will be disregarded. On this aspect of the subject 
the investigations of the learned Wurffbain (1683) maybe consulted. 
A typical example of these fables was the belief that the asbestos 
fibre, the incombustible nature of which was soon recognized, was 
the hair of the Salamander, and it was sometimes even referred to as 
Salamander's wool. 

There are several recognizable figures of S. maculosa^ of varying 
degrees of excellence, in the earlier treatises on Natural History, such 
as Mattiolus' Commentary on Dioscorides^ Gesner, Pare, Imperato, 
Aldrovandus, Perrault, Seba, Owen, and Roesel, of which that of 
the last-named naturalist hardly exhibits the accuracy of so careful 
an observer of nature. Of the Amphibia the Frog and Newt were 
investigated before the Salamander, and much more thoroughly. 
The earlier workers, however, included a number of species under 
the term Salamandra, and considered themselves at liberty to ex- 
pound the anatomy of any one of them without further discrimina- 
tion. Hence for example Triton would be dissected and described 
as Salamander, as if the species were anatomically interchangeable. 
This introduces a difficulty of some importance in the interpretation 
of the older literature, and especially in those cases where the figures 
are not sufficiently accurate to make distinction possible. Where, 
however, the figures are good, as in the case of Monro (1755), no 
doubt can exist, and Monro is most certainly describing Triton and 
not the Salamander of his title-page. Jacobaeus also calls Triton a 
Salamander, and here again the Triton determination is beyond ques- 
tion. Indeed nearly all the earlier work labelled Salamander seems 
to be Triton, in spite of the fact that the name Salamandra was applied 
to the true Salamander from Gesner onwards, and the differences 
between S. terrestris^ or the large earth Salamander, and S. aquatica, 
or the small water Salamander (Triton), were usually recognized. 

A statistical survey of the obsolete Salamandra literature shows 
that most of it is grouped between the years 18 15 and 1850. This 
grouping coincides almost exactly with that of Amphibian literature 



xiv HISTORICAL INTRODUCTION 

generally of the same period, which would indicate that the research 
of the time was focused on the phylum as a whole and not on any 
particular member of it.^ 

Aristotle, to whom the terrestrial Salamander must have been well 
known, and who named it UaXanavSpa, does not deal with the anatomy 
of the species. Schneider has suggested that Aristotle's KopBvXos may 
have been an aquatic or a larval Amphibian, and von Siebold actually 
refers to the larval Salamander as Cordylus, but Leuckart (1823) 
rightly questions the accuracy of this determination. Albertus 
Magnus identifies the Salamander with the SteUio of ancient authors 
— the only justification for which seems to be its poisonous nature 
and the spots on its back. 

Belon (1553) appears to have been the first observer to discover 
the viviparous nature of the Salamander, which, he says, like all 
vipers, brings forth its young alive. According to Gesner (copied by 
Aldrovandus and Wurffbain) Belon found that the larvae, to the 
number of forty to fifty, were born without any foetal membranes, 
but the latter passage has not been traced in any of Belon's works. 
Gesner (1554) is the first author to mention S. atra. He says that 
there is a dull black Salamander in the Alps similar in form to the 
spotted Salamander, and secreting the same white fluid from the 
skin, but with a short tail. He adds that Salamandra is the only one 
of its kind to exhibit intra-uterine fertilization. Goiter (1575), the 
first comparative anatomist after Aristotle, describes the skeleton of 
the Lizard in some detail, and he has also investigated the skeleton 
of the Salamander, which he says is similar to that of the Lizard. The 
first confirmation of Belon's discovery comes from Dalecampius 
(1587). In commenting on Pliny's statement that Salamanders have 
no sex and are not the result of normal generation, he says that a 
battered Salamander exuded from all parts of its body a milky 
venom, and from the ruptured abdomen there emerged over thirty 
ova, and more than thirty foetuses, which swam actively in water. 
He also dissected a pregnant female which was full of eggs and 
foetuses, some of which were incompletely developed, but others 
were quite advanced and were able to move and swim in water. He 
therefore concludes that it is false to say, as Pliny does, that Salaman- 
ders are generated from nothing. Imperato (1599) carries the same 
subject farther. He claims that Salamanders pair like Vipers, and 
at the time of bringing forth they congregate in the neighbourhood 
of ditches, and await the time of rainfall. They then enter the water, 
and discharge their foetuses in one day to the number of from 
^ Cf. Cole and Eales, Science Progress, xi, Fig. 6, 19 17. 



HISTORICAL INTRODUCTION xv 

twenty to forty. The membranes in which the foetuses are enclosed, 
after the manner of spiders, are soon thrown off.^ A figure is given of 
the larvae, which clearly belong to S. maculosa^ but no external gills 
are shown. An early dissection of a female Salamander carried out 
by Hoffmann c. 1642, but not published until 1722, included an 
examination of the heart, in which only one auricle is mentioned, the 
abdominal viscera, together with the spleen, gall-bladder, and bile- 
duct, and certain of the larger blood-vessels. He notes that the 
uterus is paired, and from it he took thirteen small living foetuses 
which were black in colour and not yet provided with the yellow 
markings. Somewhat later, in 1673, Steno describes the two ovaries 
and fat bodies and also the two oviducts (with contained ova) which 
he says are not joined at either end. 

The first attempt to work out the general anatomy of a 'Salamander' 
was made by Jacobaeus (1676-86). His little book was severely 
criticized by Swammerdam, whose own monograph on the Frog was 
on a much higher plane. A careful examination of Jacobaeus's text, 
and a comparison of his figures with dissections of Triton, make it 
quite certain that his subject was Triton and not Salamander. In 
general his statements are more or less admissible if applied to 
Triton, but are difficult to understand in relation to Salamandra. He 
gives for the time a fairly complete account of the abdominal viscera, 
and makes the interesting observation that the heart beats for many 
hours after removal from the body, as in Frogs, Toads, and the 
Torpedo. He figures what might be scattered supra-renal nodules, 
but from their distribution are perhaps the Malpighian bodies. 
Almost contemporary with Jacobaeus is the more ambitious and 
scholarly work of Wurff bain (1677, 1683). He claims to have dis- 
sected several terrestrial Salamanders, which hardly accords with the 
fact that his visceral anatomy is taken almost entirely from Jacobaeus, 
and his notes on the skeleton from Goiter. He, however, correctly 
describes the contents of the stomach, which Jacobaeus found to be 
empty, and mentions for the first time the shedding of the epidermis, 
but as he says it takes place in water he can hardly have been dealing 
with Salamandra. He looked for the auditory organ but was unable 
to find it. Previous statements that the terrestrial Salamander is 
viviparous are confirmed, and Wurffbain found thirty-four larvae in 
one specimen. He illustrates the appearance of these larvae, but as 
no external gills are shown or mentioned, we may suspect that his 
figures owe more to Imperato than to nature. 

^ Imperato is quite correct in this. The Salamander is normally ovo-viviparous, but 
the egg membrane may rupture before the egg is laid. 



xvi HISTORICAL INTRODUCTION 

A more serious attempt to anatomize the Salamander, this time 
without any Triton complications, was made by Perrault c, 1680, 
but not published until 1734. He dissected two specimens — a male 
and a female. He noted that the black parts of the skin when viewed 
under the microscope showed a large number of yellow spots which 
were almost invisible to the unassisted eye. There was no external 
auditory opening as in Lizards, but the animal could hear never- 
theless. The tongue, liver, teeth, gall-bladder, and gut are well 
described, but the pancreas is assigned a wrong position, and his 
hepatic duct seems to have been a blood-vessel. Various mistakes 
and omissions occur in the description of the bladder and urogenital 
system, and his version of the female genital ducts is simply incom- 
prehensible. It is perhaps remotely possible that his specimen may 
have been an hermaphrodite. He asserts that, contrary to accepted 
belief, the Salamander is sexual and has distinct male and female 
genital organs, however peculiar they may be. He failed to notice 
that the young larvae had external gills, and he described the heart 
as having only a single left auricle. Vallisneri (17 15) apparently 
regarded Triton and Salamandra as belonging to the same genus. 
His remarks on the poisonous white secretion of the skin, its peculiar 
colour, and the supposed reaction of the animal to fire show that he 
had the true Salamander in mind, but his anatomical work must have 
been based on Triton. Thus he describes long lungs which extend 
the length of the abdomen, he denies that there are four testes, he 
identifies two penes as in snakes and lizards, and in one specimen he 
found the stomach to be full of frog's eggs, and in another of fish 
eggs. Maupertuis (1727) redescribes the 'milk' glands of the skin, 
and from one female he took forty-two, and from another fifty-four, 
well-developed active larvae, but he says nothing of the external 
gills or of the caudal fin. Although the work of Du Fay (1729) 
relates almost entirely to Triton, he deserves mention as being the 
first author to describe those very obvious larval structures — the 
external gills. He found also the 'internal' gills, the gill arches and 
slits, and made an important contribution to the anatomy of meta- 
morphosis. He was likewise the first to describe the double urogeni- 
tal papilla of the male Salamander, which he regards as a penis — a 
structure which, he says, a viviparous animal should possess. He 
'presumes' that the terrestrial and aquatic Salamanders are vivi- 
parous and oviparous respectively. He traces the passage of the 
eggs from the ovary through the ostium into the oviduct and so to the 
exterior. The 'penis' was subsequently described by Latreille in 
1800 and by numerous later authors, but others either failed to 



HISTORICAL INTRODUCTION xvii 

observe it or doubted its erectile nature, and Duvernoy even denied 
its existence. 

The celebrated Monro secundus, in his inaugural dissertation 
(1755), gives us an admirable description of the male genitalia of 
'Salamandra'. He mentions a connexion between the genital system 
and the heart by which the semen is supposed to reach the blood- 
stream. This may be either Muller's duct, which is always present 
in the male, or the post-cardinal vein. Monro's paper is the most 
careful account of the male genitalia and renal organs with their 
ducts, the cloacal glands (which he notes are not developed in the 
female), lung and fat body of a Urodele so far published, but the 
subject was undoubtedly Triton and not Salamander. In the short 
and little known paper by Zinn published in 1757 there are many 
brief anatomical notes covering almost all the organs of the body 
except the nervous system, of which the most important is his dis- 
covery of the operculum which fits into the fenestra vestibuli 
(ovalis) of the ear. He also saw the otolith, and noted the absence of 
auditory ossicles and an external ear duct. In 1758 Roesel, who died 
in the following year, was working on an HistoriaNaturalisSalaman- 
drarum^ which was never completed or published. J. Hermann, in 
1789, reported that he had seen the plates, which compared favour- 
ably with Roesel's beautiful illustrations of the Anura published in 
1758. No descriptions of these plates have been found. Kleeman 
says that Roesel hoped to publish a work on the Lizards and Sala- 
manders of his native country in the same style as that on the Frogs, 
but that he died before he was able to complete it. 

The genus Salamandra was instituted and defined by Laurenti in 
1768, and he was the first to name and figure S. atra from the Alps — 
a species first described by Gesner. The name Salamandra occurs in 
Aristotle, and has been used by many pre-Linnean systematists. It 
was applied indifferently to any tailed European Amphibian, or even 
treated as an abstraction without any objective equivalent. Laurenti 
draws attention to the fact that the Amphibia, being without a 
diaphragm and ribs, cannot breathe like a mammal, but that by 
alternate movements of the throat they can empty and fill the lungs 
with air. He was thus familiar with the idea of the buccal force- 
pump, but was not aware that this had already been described by 
Swammerdam in 1 667, and also that there was an admirable analysis 
of buccal respiration in the posthumous works of Malpighi pub- 
lished in 1697. Malpighi indeed was the first to show that when the 
lower jaw of the frog was severed the lungs could no longer be 
inflated. 

4038 u 



xviii HISTORICAL INTRODUCTION 

John Hunter's work on the Amphibia, which must have occupied 
his attention round about the year 1780, but which was not pub- 
lished until modern times, is characteristic of his original and pene- 
trating mind. There are eleven Hunterian preparations of Sala- 
mandra still to be seen in the Hunterian museum. They illustrate 
the abdominal viscera of male and female in some detail, and a 
S. atra is shown with one of the two foetuses to which it is limited, 
but this last preparation was presented by Buckland and is not 
Hunterian. Hunter attempts an instructive comparison of Salaman- 
dra and Triton in respect of their structure and reproductive habits, 
with which he exhibits a greater familiarity than his predecessors, 
and his interpretations are remarkably sound. As usual, however, he 
did not concern himself with the work of others, and some of his 
points, though original as far as he was concerned, were not new. In 
his classification according to the structure of the heart he institutes 
a group named the Tricoilia (= Reptiles and Amphibians) in which 
the heart has three chambers — two auricles and one ventricle. This 
classification was first printed in the posthumous work on the 
Blood in 1794, but no details are given there, and the complete 
scheme was only published long after Hunter's death. He included 
the Salamanders in the group with the two-auricled heart, and was 
therefore the first to perceive the essential structure of the Amphi- 
bian heart. 

Blumenbach (1787) is the first author to mention all the points 
which characterize the larval Salamander, viz. the external gills, the 
four appendages, and the aquatic, as contrasted with the terrestrial, 
type of tail. This may appear to be observation of an elementary 
character, but we have seen that the external gills were completely 
missed by the earlier naturalists, and even as late as 1800 Latreille 
was denying their existence, whilst Saint-Julien in 1789 interprets 
the gills as a pair of long fins, and failed to discover any traces of the 
limbs. Blumenbach also held that female Salamanders which were 
for months isolated from males nevertheless gave birth to young, 
from which he concludes (i) that the eggs are not fertilized after 
they are laid and that the Salamander is truly viviparous; and (ii) 
that, as in the case of the fowl, a single act of fertilization is effective 
for some time. He found that the Salamander was capable of 
regeneration, and that if a third of the tail was cut off it could be 
completely, but not quickly, regenerated. The first detailed descrip- 
tion of the auditory organ of a Urodele is given by Scarpa (1789), 
but his observations relate to the aquatic Salamander (Triton). 
The essential facts are correctly stated, and he realized that Triton 



HISTORICAL INTRODUCTION xix 

was morphologically not the Reptile it was supposed to be. Townson 
( 1 794— 5) draws attention to the fact (previously known to Aristotle, 
Coiter, and Malpighi) that the lungs of Frogs do not collapse when 
the wall of the cavity in which they lie is opened up. Therefore the 
mechanism of respiration must be very different from that of the 
Mammalia. He then proceeds to investigate buccal respiration in 
Salamandra. He describes and figures a pair of long narrow 'sterno- 
hyoid' muscles,'' which arise from the pelvic girdle and are attached 
to the hyoid. These muscles draw the hyoid downwards and back- 
wards, and thus increase the cavity of the mouth. When they are 
cut respiration ceases. Other muscles, the constrictors of the 
pharynx, elevate the hyoid and draw it forwards — thus contracting 
the cavity of the throat. According to Townson, when the mouth 
cavity is opened up and the muscles of the hyoid left intact, the 
hyoid is still drawn backwards and forwards, and the glottis opened 
and shut, although the lungs remain collapsed; but when the muscles 
of the hyoid are cut the glottis alone is unaffected, and still continues 
to open and shut. Townson claimed that the contents of the bladder 
were as pure as distilled water. It is, therefore, not a urinary bladder, 
but acts as a reservoir for water which is 'further distributed as their 
economy may require'. 

The first comprehensive treatise to collate previous observations 
on the Salamander, and combine them with many original observa- 
tions, is to be found in Cuvier's Lefons. In the first edition of this 
important work the Salamander material relates chiefly to the skele- 
ton and muscles. Between the first and second editions, however, 
the Salamander must have been further investigated, since the latter 
edition, which was posthumous and not prepared by Cuvier, in- 
cludes much additional matter on the species. In the first edition of 
of the Lefons (1805) he states that the Batrachian heart has a single 
auricle and ventricle, and in the second edition (1839) the statement 
is repeated with the editorial qualification, as regards the auricle, of 
the word 'apparently'. The editor of the latter edition, G. L. Duver- 
noy, who had evidently read Davy's paper of 1828, proceeds to add 
that the auricle is divided into two chambers by a very thin partition, 
more membranous than muscular, the free border of which penetrates 
slightly into the cavity of the ventricle. In the tadpole, however, the 
heart is like that of a fish, and has only one auricle. In the second 
edition of the hegons Duvernoy (1846) gives a good description of 
the cloacal glands of Salamandra under the name of 'prostates'. This 
was presumably based on Rathke. In the Ossemens Fossiles of 1 824, 
' = rectus abdominis profundus+ rectus cervicis profundus of this work. 



XX HISTORICAL INTRODUCTION 

which appeared between the first and second editions of the hegons^ 
Cuvier adds and codifies the work already published in 1800 in the 
first and second volumes of the Legons^ and he now gives for the first 
time a very good description with figures of the entire skeleton of 
Salamandra, which is compared with that of the Frog. His interpre- 
tation of the bones of the skull naturally differs from modern usage. 
For example, he describes the posterior nares as entering the mouth 
behind the vomer, but as Cuvier's vomer is a pre-vomer the position 
of the internal nares is not as anomalous as he supposed. The 
Y-shaped or ypsiloid cartilage in front of the pubic symphysis, which 
he compares with the marsupial bone of Mammals, had been already 
described, figured, and named by Townson. The Urodele types are 
somewhat confused in Cuvier's work, and it is not always certain 
to which of them a particular statement refers. 

In Daudin's voluminous treatise on the Reptiles (1803) the Sala- 
manders are correctly located with the Frogs and separated from the 
Lizards and Reptiles. The idea was not a new one, but hitherto it 
had not been expressed in any classification. Thus Cuvier compared 
the Salamanders with the Frogs, but continued to classify them with 
the Reptiles. That the Amphibia were entitled to independent rank 
was emphasized in 1806 by Dumeril, who instituted the groups 
Urodela and Anura under the French names of Anoures (Ecaudati) 
and Urodeles (Caudati) as the two families constituting the Batrachia 
— the latter term with the spelling Batrachii having been previously 
introduced by A. Brongniart in 1 799. The Latin term Urodela was 
coined by Latreille in 1825. 

Important work on the anatomy and reproductive habits of the 
Salamander was published by Gravenhorst in 1808 and 1829. The 
first of these is a short preliminary communication, without illustra- 
tions, in which he refers to thirty-one figures of the anatomy of land 
and water Salamanders which are presumably those published later 
in the folio work. Gravenhorst's volume includes a useful examina- 
tion of most of the preceding literature, the mistakes in which he 
endeavours to correct, and he has very carefully revised the anatomy 
of the gut and urogenital organs of both sexes. The drawings of the 
latter organs are particularly good and accurate, and his figures 
generally are faithfully drawn, and do not include any structure he 
has not seen and closely examined. He discovered Miiller's duct in 
the male, and shows its forward extension, but did not understand 
how the seminal fluid reached the vas deferens, since he found no 
direct connexion between this duct and the testis. Nor does he dis- 
tinguish between Miiller's duct and the vas deferens, but these two 



HISTORICAL INTRODUCTION xxi 

structures are so closely associated in Salamandra that their relations 
were only understood when the histologist Leydig examined them 
microscopically in 1853. Judging from appearances in one of his 
figures Gravenhorst seems to have found the receptaculum seminis 
— later discovered by Rathke, Leydig, and von Siebold. His figures 
of the developing larvae are better than those of Funk and Rusconi's 
earlier drawings. Jacobson (18 17) describes the relations of the 
anterior abdominal and renal portal veins in the Batrachia and Sala- 
mander, and concludes, as Townson had done before him, that the 
bladder in these animals is in no sense a urinary bladder, since its 
structure is entirely different, and the ureters are not related to it as 
they should be in a true urinary bladder. A brief but accurate 
account of the auditory organ of the Salamander was published in 
18 18 by Pohl. He describes the bony capsule, the external vacuity 
closed by the 'cartilaginous operculum' discovered by Zinn, and the 
entry of the auditory nerve. He calls the vacuity the 'fenestra vesti- 
buli' as in modern usage. He describes the three semicircular canals 
and their ampullae, and gives a figure of the operculum and the 
fenestra which it closes. C. A. S. Schultze in 1 8 i 8 was the first to 
reveal a curious osteological point in the terrestrial Salamander, viz. 
that the transverse processes of the vertebrae are bifid, and articulate 
with the bifurcated ends of the ribs. In the same year Meckel pub- 
lished a comparison of the hyobranchial skeleton of Salamandra and 
Triton, which, however, was based largely on Cuvier. The same 
author's treatise on comparative anatomy is a useful summary of 
existing knowledge, but perpetuates many errors. He asserts that 
there is only one auricle of the heart in the Amphibia, and that it 
contains mixed blood, but admits that in Salamandra and Triton the 
auricle is divided by a well-marked constriction into an anterior 
larger and a posterior smaller section. He mentions that he sought 
for a carotid gland in Salamandra, but was unable to find it, although 
he was more successful in other Amphibia, 

The third encyclopedic treatise on comparative anatomy to be 
published in the early years of the nineteenth century was the Lehr- 
buch of C. G. Carus, the first edition of which appeared in 1 8 1 8 and 
the second in 1834. Carus had evidently dissected several examples 
of S. maculosa more or less carefully, so that this work is not entirely 
a compilation. He describes the valve in the nostril^ which he says 
is like an eyelid, and noted its relation to buccal respiration. He 
agrees that the bladder is only a water-holder, and even regards it as 
a possible respiratory organ, since it is analogous to the allantois of 
^ Not, however, present in the Salamander. 



xxii HISTORICAL INTRODUCTION 

Lizards and Birds. The old mistake that the heart has only one 
auricle is repeated, but he states correctly, contra Swammerdam, 
that there are not two carotid glands on each side, but only one. His 
views on the skeleton are dominated by his adhesion to the vertebral 
theory, which he accepted in its most extravagant form. The whole 
work is useful as a restatement and discussion of previous literature. 
A more important contribution by Carus to Salamandra research is 
his paper of 1 8 1 9. In this he describes correctly the parts of the gut, 
and gives details of the female genitalia. He found the ostium of the 
oviduct, and wondered how the large eggs could pass through it. 
Although the eggs develop in the uterus they are entirely free, and 
there is nothing corresponding to a placenta. Neither is there any 
connexion between the foetus and the egg membrane. The anterior 
abdominal vein is interpreted as a persisting foetal structure, and 
comparable with the umbilical vein of the Mammalian foetus. He 
gives a good account of the gross anatomy of the larva, including 
the gut, and believes that the yolk mass is contained within the gut 
wall, and is thus nothing more than an inflated section of the gut. In 
fact both the yolk-sac and the urinary bladder or allantois remain 
within the body wall — unlike the condition found in Mammals. In 
his last paper dealing with the Salamander (1828) Carus publishes 
two original figures of the trunk and limb muscles of S. terrestris^ 
but the only mention of these figures in the text is to be found in the 
description of the plates. They illustrate the more superficial muscles 
of the head, body-wall, limb-girdles, and limbs. 

The genius of Rathke was first exercised in an inaugural disserta- 
tion on generation in Triton which was published in 18 18. Two 
years later it was expanded and reissued, and now included the Sala- 
mander as a closely related form, the two genera being systematically 
compared. He describes the development of the fat-body and its 
relation to the general metabolism of the animal. In starved indi- 
viduals the fat-body dwindles and even disappears, and during the 
winter it is used up. All the fat in the body of the Salamander is 
concentrated in its fat-body.^ The development of the genitalia with 
their ducts, as far as it can be followed with a lens, is attempted for 
the first time. The growth of the eggs in the oviduct is given in 
detail, but he regards them as efflorescences of the ends of the 
ovarian vessels. His mistake that the eggs escaped through an aper- 
ture in the ovary was corrected by Czermak in i 843. A comparison 
of the finer structure of the ovary and testis led him to believe that 
from first to last they are identical structures, the adult testis being 
' In these statements Rathke goes too far. 



HISTORICAL INTRODUCTION xxiii 

nothing more than a higher development of the ovary. The sperma- 
tozoa are not recognized as such, and therefore a comparison of egg 
and sperm is not attempted. He is puzzled to discover how the 
semen passed from the testis to the vas deferens, and had to abandon 
this question unsolved. He describes and figures a 'gland' associated 
with the cloaca of the female Salamander which we now know to be 
the receptaculum seminis. Rathke's paper therefore contains the 
first undoubted description of this structure. He does not mention 
that it contains spermatozoa, but neither does he concern himself 
with the spermatozoa in the testis. The receptaculum seminis of the 
Salamander, as already stated, was rediscovered by Leydig in 1853, 
who found (later) that it contained spermatozoa, but it was not 
adequately investigated until Siebold ascertained its true nature in 
1858. Siebold regarded it as a unique structure only occurring in 
the Salamanders. He overlooked the possibility that the organ might 
be represented in the male by the pelvic gland, as was suggested later 
by Blanchard (i 88 i). If the receptaculum seminis is not comparable 
with the pelvic gland, the cloacal glands of the male have no counter- 
part in the female. Rathke was the first author to study carefully the 
male cloacal glands, and he gives good descriptions of both the pelvic 
gland {Beckendruse) and the anal or cloacal gland {Afterdriise). His 
views on the nature and functions of these glands have naturally 
required revision, and the whole situation was reviewed by Blanchard 
in 1 88 I. Rathke's work is well illustrated, the figures covering the 
anatomy of the Salamander generally. They are most defective in 
respect of the urogenital system of the male, the elucidation of which 
was perhaps beyond the knowledge of his generation. In a later work 
(1829) Rathke describes for the first time the fragmented adrenal of 
the Salamander. 

In 1 8 19 Schreibers publishes a very good description, as far as 
external characters are concerned, of the life-cycle of both species of 
Salamandra. In atra he never found more than two foetuses in each 
female. They possessed very large red gills in the earlier stages, 
which were almost completely lost before birth. In a later paper 
(i 833) he draws a sharp distinction between the two species in struc- 
ture, occurrence, and habits. Maculosa produces periodically in 
water batches of forty to sixty young, the average number being 
forty-six to forty-eight. In captivity they w.ere born within two con- 
secutive days, and all were at the same stage of development and of 
about the same size, and provided with external gills. On the other 
hand atra never gives birth to more than two at a time. Nevertheless 
the ovary is relatively as large as in the other species, and as many 



xxiv HISTORICAL INTRODUCTION 

eggs reach the oviduct together. Only one, however, in each oviduct 
develops, the remaining eggs fusing to form a common yolk-mass 
which serves for the nourishment of the surviving embryo, who is 
thus the 'assassin of his brothers and sisters' (Dumeril). The gills 
are half the length of the entire body and extend backwards to the 
hind limb, but they degenerate before birth, and the young are pro- 
duced in the adult state with scarcely any trace of the gills, and none 
of the gill openings. This is ascribed to the fact that atra lives in 
situations where water may be scarce or not available, and hence the 
metamorphosis must be completed within the uterus. Schreibers 
asserts that he has observed atra in captivity coupling in water as in 
the Frog, and that if the larvae of maculosa are forcibly kept under 
water the metamorphosis may be delayed for many months. 

The spermatozoa of the Salamander were first described by Pre- 
vost and Dumas in 1821. The work of Dutrochet (182 1-2), al- 
though it relates to Triton only, may be mentioned as containing the 
first account of the 'internal' gills of the Amphibian larva. He also 
makes the suggestion that the regeneration of the limbs and tail is 
not a case of preformation, but of the epigenetic development of parts 
much as they were originally produced in the embryo. The ear of 
Salamandra and its semicircular canals were re-described by Blain- 
ville in 1822. He says that there is no trace of a tympanic cavity or 
Eustachian tube, but adds (incorrectly) that these parts are well 
developed in the larva. He shows for the first time that the oper- 
culum which closes the fenestra vestibuli, thereby taking the place 
of the auditory ossicles, can be moved by a component of a shoulder 
muscle (M. opercularis). Two years later Huschke, also working at the 
auditory organs of Salamandra, confirms the absence of a tympanic 
cavity and Eustachian tube, and asserts that the operculum is rather 
a detached portion of the skull (as it is) than a true ear bone. He 
found that the operculum was elastic, and would spring back into its 
place after it had been pulled up. He holds that the columella and 
ear ossicles arise from the anterior gill arches, that the tympanic 
cavity represents the gill cavity of the fish, and that the Eustachian 
tube is an inner gill opening. 

Dr. J. Davy, the younger brother of Sir Humphry Davy, was the 
first author after Hunter to throw any light on the structure of the 
Amphibian heart. He did not examine the Salamander, but his work 
has general bearings which are too important to be disregarded. In 
his first communication of 1826 he finds that the heart of toads and 
frogs consists of two auricles and one ventricle, the auricles being 
separated by a fibrous septum, and he considers it probable that the 



HISTORICAL INTRODUCTION xxv 

same condition prevails in all other members of the group. In the 
full paper, published two years later, but dated 2 July, 1825, he 
states that the auricular septum 'appears' to have muscular fibres, 
and that there are three semi-lunar valves between the auricles and 
the ventricle. There is no direct communication between the two 
auricles. The smaller auricle is associated with the pulmonary veins 
and the larger one with the venae cavae. The two auricles contract 
first, then the ventricle, and finally the conical part of the ventricle 
( = truncus arteriosus), which latter he regards as a second ventricle. 
It is separated from the rest of the ventricle by three semi-lunar 
valves, and its cavity contains a fleshy projection or movable septum 
( = spiral valve). There are further semi-lunar valves between it 
and the aortic vessels. He asserts definitely that the pulmonary 
arteries only accompany the aortic trunks and are not branches of 
them. It is not often that observations of such relative accuracy and 
importance are included in so unpretentious a paper. 

The monograph on the Salamander by Funk published in 1827 
becomes less impressive the closer it is compared with the work 
which preceded it. Funk is at his best on the muscles, which had 
previously not been systematically examined, but the remaining sec- 
tions of the work, which cover more or less the entire anatomy of the 
type, are much less satisfactory, even for the time they were written. 
The historical chapter, however, bears evidence that much labour 
and knowledge were concentrated on its production — indeed Funk 
is a better historian than anatomist. He does not touch the cranial 
nerves, and there are only brief notes on the sense organs, nor did he 
succeed in finding the sympathetic. The otolith is identified as a 
rudimentary auditory ossicle, and he has no observations to off'er on 
the lymphatic system, the investigation of which he admits is beyond 
him. There is a physiological section dealing with the effects of 
various gases and vapours on respiration. The most inaccurate parts 
of Funk's monograph are those which deal with the vascular system 
and the brain. He only distinguishes two chambers in the heart — 
an atrium and a ventricle — an error which induced him to make the 
pulmonary veins discharge into the vena cava. He found no valves 
in the heart, and his account of the distribution of the vessels is de- 
fective both as regards observation and interpretation. He appears 
to have missed completely the carotid arch — h\s first pair uniting to 
constitute the dorsal aorta, and the veins are only partly, and not 
accurately, figured. He even seems to have confused arteries and 
veins. Rusconi later revised this part of Funk's work, and corrected 
his mistakes. As regards the brain, the figures are imaginative and 



xxvi HISTORICAL INTRODUCTION 

have little relation to the objects they represent, and his description 
is too meagre to throw any light on his methods, or to suggest an 
explanation of his errors. Funk's account of the gut and male and 
female genitalia is much better. He describes a three-lobed testis 
(it is usually two-lobed), and he made a serious attempt to unravel 
the relations of the genital glands and ducts — a full understanding 
of which, however, was beyond the technique of his time. Two years 
later Gravenhorst materially improved on Funk's descriptions of the 
genitalia. 

Cachet's paper of 1 827MS an interesting first-hand account of the 
feeding and reproductive habits, metamorphosis, and regeneration 
in the Salamander, but it contains little that is new. A year later 
Siebold published his inaugural dissertation on the Salamander and 
Triton. It is a remarkable and important maiden effort. He is the 
first to trace accurately and in detail the metamorphosis of the larval 
hyobranchial skeleton and musculature into that of the adult. He 
gives a very good description with figures of the larval and adult 
states, and also an excellent illustrated account of the anatomy of 
metamorphosis, especially as regards colour, gills, operculum, and 
tail. It is a very instructive study in the physiology of adaptation, 
and of the phenomena which accompany the change over from aqua- 
tic to terrestrial life. He shows further how metamorphosis may be 
controlled or modified by varying the conditions. A sketch of the 
development of the internal organs is included, and there is a section 
on the myology of the tongue and hyoid, in which the movements of 
the tongue are explained. 

The first statement of the discovery of the posterior lymph hearts 
of the Frog was made by J. Miiller in September 1832. A year later 
the anterior lymph hearts were found, not only in the Frog and Toad 
but also in the Salamander and Lizard. Miiller observed the con- 
traction of the hearts, and noted that the rate was peculiar to the 
organ, and different from that of the heart and lungs, nor were the 
pulsations of the two sides synchronous. The hearts contained a 
clear lymph and not blood. The methods employed by Miiller in the 
investigation were mercury injections and inflations, and he claims 
that a pulsating lymphatic organ had not previously been seen. He 
states that the posterior lymph hearts are difficult to find in the Sala- 
mander, but that they are in the same position as in the Frog. Panizza 
had already seen the lymph hearts of the Frog in 1830, but had not 

^ The volume is dated 1827, but the author refers to an incident which happened 
in April 1828. He states that his plates were being published elsewhere, but so far no 
such publication has been traced. 



HISTORICAL INTRODUCTION xxvii 

observed them in the living animal, and hence was unaware that they 
pulsated and were therefore hearts. 

Windischmann (1831) dissected five Salamanders, and describes 
the 'finer' structure of the auditory organ. He was the first to observe 
that the fenestra vestibuli (ovalis) is closed by a membrane to which 
the operculum is related, the operculum itself not occurring in the 
larval Salamander. His figure of the labyrinth is crude and distorted, 
but he noted the absence of a cochlea and of the fenestra rotunda. 
Bischoff (1832) gives a small and poor figure, but hardly any descrip- 
tion, of the dorsal surface of the brain of Salamandra — a part of the 
animal which had so far been almost entirely neglected. In 1832 
Martin Saint-Ange confirms the existence of two auricles in the 
heart of the Anura, and adds that in the Salamanders also there is 
a small but distinct left auricle. 

The monograph by Panizza on the lymphatic system of Reptiles 
published in December 1833 ^^ one of the most important of the 
earlier works dealing with the comparative anatomy of this system. 
He used injections of mercury, and coloured masses such as size, and 
his material included representatives of most orders of Reptiles and 
Amphibia. He discovered the anterior and posterior lymph hearts 
independently of Muller, and proved by repeated experiments that, 
as the lymphatics did not open into the gut and were peripherally 
distinct from the true blood-vessels, they must therefore be re- 
garded as an independent system. He found that in Salamandra the 
lymphatics were developed to a prodigious extent, and that the net- 
works were of astonishing minuteness. On the other hand the main 
trunks were unusually capacious, and communicated on each side 
with the anterior vena cava by valved apertures. There were further 
openings posteriorly into the pelvic veins, and at this point a con- 
tractile posterior lymph heart driving the lymph into the veins might 
be present. He did not, however, find the latter heart in Salamandra. 
The main lymphatic cistern in Salamandra extended from the anus 
to the pylorus, expanding as it passed forwards. In the abdomen it 
divided into two sinuses associated with the genitalia, which united 
again anteriorly to form the thoracic duct. The latter bifurcated 
in the neighbourhood of the heart, each division receiving the 
lymphatics from the region of the fore-limbs and head of its side, 
and opening finally into the subclavian of the same side by two or 
three small apertures. In a later paper published in 1845 Panizza 
regards the respiratory process in Salamandra as one of deglutition 
of air, and he develops in detail the theory of the buccal force-pump 
and explains the physiology of the muscles and skeleton concerned. 



xxviii HISTORICAL INTRODUCTION 

The backward extension of the sternohyoid muscle to the pubis is 
not associated with the extra length of the lungs, as suggested by 
Haro (1840), but is due to the fact that there is no fixed sternum 
as in the Frogs. The sternal region of the pectoral girdle in Sala- 
mandra moves with the limbs, and therefore offers no point d'appui 
which could serve for muscular attachment. In the following year 
Panizza is engaged on an experimental investigation into the flow and 
velocity of the respiratory current in the larva of the Salamander. He 
describes the action of the cilia in directing and maintaining the 
current, and examines the conditions which affect the vitality and 
irritability of the ciliary mechanism. 

Duges's work on the osteology and myology of the Batrachia 
(1834) includes some observations on the terrestrial Salamander. 
The figures are good and accurate, but not accompanied by adequate 
descriptions. The object of the paper is to compare the condition of 
the skeleton and muscles of larva and adult. Some structures are 
peculiar to the larva and others to the adult, but those which occur 
in both undergo modification during metamorphosis. The figures 
are the most detailed and accurate of these two systems so far pub- 
lished, but the Urodeles have not been so carefully studied as the 
Anura, and Triton figures much more largely in the paper than 
Salamandra. Reichert's work of 1838 also relates to Triton, but it 
is very important from the point of view of the development and 
morphology of the skull and visceral arches in the Urodela. Henle 
is the first author to investigate the anatomy of the larynx of Sala- 
mandra, which he describes in a noteworthy and accurate work pub- 
lished in 1839. 

We now reach the numerous and striking contributions to Sala- 
mandra literature for which Rusconi is responsible. The first was 
published in 1839, and the last, which was posthumous, in 1854. 
Owing to the fact that Rusconi coloured the plates of his papers with 
his own hands, and was unable or unwilling to devote the necessary 
time to this task, several of his most important writings are very rare, 
having been limited to editions of 100 copies, and in one case of only 
24. In his first paper Rusconi states correctly that in S. maculosa the 
sexes couple on land and not in water, and therefore the sperm 
cannot reach the oviducts mingled with water as had hitherto been 
accepted. He further denies (this time wrongly) that only two larvae 
may be produced at a time in a Salamander, thus showing that he was 
not aware of the difference in this respect between maculosa and atra. 
In fact he declines to believe that there can be such a considerable 
distinction between the species. He was greatly puzzled when he 



HISTORICAL INTRODUCTION xxix 

discovered that segmenting eggs occurred in the oviducts before 
pairing had taken place, nor did he succeed in throwing any light 
on the problem of fertilization in the Salamander. In his posthumous 
treatise on the terrestrial Salamander, which may be considered next 
since the work was done before 1 839,^ he does not seem to have been 
aware that the Amphibian heart has two auricles. He definitely says 
it has not in Spelerpes, and as regards Salamandra he always uses the 
term atrium in the singular. He gives a very full account, accom- 
panied by excellent figures, of the development of the external 
characters, from the segmentation of the egg up to the adult stage. 
His outline of the development of the internal organs, such as the 
gut, nervous system, and skeleton, is a very definite advance on the 
standard of such work at his time. The anatomy of metamorphosis 
is accurate and more detailed than any previous description, and he 
is especially admirable on the metamorphosing skull, and on the 
changes undergone by the muscular and vascular systems. He de- 
scribes and figures the carotid gland,^ which he says *is actually a 
consequence of the shortening of the artery at the moment of meta- 
morphosis'. He did not succeed in finding the lymph hearts, but 
describes, and beautifully illustrates, the lymphatics of the gut and 
bladder. Perhaps, however, his most interesting discovery is that of 
the pronephros and segmental duct, which he describes and figures 
under the name of 'Wolffian body'. Leydig also figures these struc- 
tures in the Salamander larva in 1857. In 1841 Rusconi becomes 
involved in a controversy as to whether the blood-vessels of the Sala- 
mander are morphologically inside or outside the lymphatics. For 
some years he adhered to the former view, and insisted that the 
blood-vessels were in direct contact with the lymph right up to their 
smallest branches in the skin. Panizza, on the other hand, main- 
tained that the arteries were only invaginated into the lymphatics, 
and were related to them as the heart was to the pericardium. The 
controversy was prolonged for some years, Rusconi repeatedly claim- 
ing that his double and triple injection methods were superior to the 
mercury injections of Panizza. In fact he was so convinced of the 
soundness of his attitude that he began to get interested in questions 
of priority, and to argue who was the first to discover this mare's nest. 

' This monograph was actually completed in 1839, when it was partly set up in 
type. Rusconi, however, postponed publication in order to revise it, but he did not 
live to complete this task. 

2 The carotid gland of Amphibia was discovered in the Frog by Swammerdam in 
the seventeenth century (published 1738). He, however, describes and figures two 
on each side, the second apparently being the thyroid gland. 



XXX HISTORICAL INTRODUCTION 

In the end, however, he had to admit that he was wrong. First of all 
he found that a vein might be accompanied by a pair of lymphatics, 
and this raised doubts whether he might not be mistaken in some 
cases. Finally he acknowledges that in the Salamander the blood- 
vessels are enclosed by a double wall, and are hence morphologically 
external to the lymph cavities, being related to them as the gut is to 
the abdominal cavity. One valuable result of this otherwise futile 
controversy was that it induced Rusconi to look closely at the 
lymphatic system of Salamandra, which resulted in many valuable 
facts being added to our knowledge of the system. 

It is surprising that up to this time no serious attempt had been 
made to investigate the nervous system of the Salamander. The first 
to do so was Fischer (1843), '^^o describes the cranial nerves in a 
work which is remarkably accurate for the time it was written. He 
did not find the fourth pair of nerves, and believed that its eye-muscle 
was supplied by a branch from the trigeminus, but he was more 
successful with the sixth nerve in spite of its small size. He was 
unable to discover more than the slightest traces of the sympathetic 
in the Salamanders, and believed that it was of much less importance 
in them than in the Anura. This conclusion is not surprising in view 
of the considerable difficulties which confront the investigator of the 
sympathetic of Salamandra. Fischer's description and figure of the 
fifth, seventh, and tenth cranial nerves are detailed and accurate. He 
points out that the fifth and seventh nerves are separate in the Sala- 
mander but fused in the Anura, and he correctly describes the 
association of the glossopharyngeal with the vagus and its anasto- 
mosis with the facial. He also understood the mutual relations of the 
auditory and facial nerves. Meyer's work on the lymphatic system 
of Amphibia (1845) ^^ largely a compilation from Panizza and Rus- 
coni, but he found two pairs of anterior lymph hearts in the Sala- 
mander which had not previously been seen. The longitudinal duct 
coursing parallel with the inner margin of the kidney into which the 
vasa efferentia open was first described by Bidder in 1846, and is 
often referred to as Bidder's duct. The discovery of this duct made it 
possible to solve the old riddle as to how the male sperm escaped from 
the body, which had puzzled so many previous workers. The sperma- 
tozoa themselves were re-described by Czermak in 1848. This 
author, who had previously (1843) explored the differences between 
S. maculosa and S. atra^ gives a remarkably accurate description, with 
figures, of the complex spermatozoon of the Salamander, and he 
even attempts to work out the details of spermatogenesis, as well 
as the mechanics of the tail and its undulating membrane. Many 



HISTORICAL INTRODUCTION xxxi 

subsequent accounts are in fact less reliable than Czermak's. The 
undulating membrane of the Salamander sperm had previously been 
imperfectly observed by Wagner, Siebold, and Dujardin in i 837—8. 
Dujardin interpreted it as a veiy tenuous free spiral thread wound 
round the axis of the tail. This was corrected by Pouchet in i 845, 
but Czermak was the first to give a classic description of the curious 
spermatozoon of the Salamander. In 1853 Leydig described and 
figured the pineal body and sympathetic ganglia of Salamandra, 
and was the first to note that the Miillerian duct of the male was 
not a branch of the Wolffian duct, but ran alongside it, the two 
being enclosed in a common membrane. He was, however, wrong in 
stating that the Miillerian duct opened posteriorly into the Wolffian 
duct. The sense organs of the lateral line were first described in 
aquatic Amphibia by F. E. Schulze in 1861, and again in 1870. 
Leydig (1868) discovered these organs in the larval Salamander, and 
they were re-described by Malbranc in 1875. Leydig gives a good 
figure of the larval S. maculosa showing the number and distribution 
of the neuromasts on the head, body, and tail. The histology of the 
organs is also illustrated, but the methods available at the time were 
inadequate for this purpose. It was only thirty years later that 
Maurer successfully analysed the structure of the amphibian neuro- 
mast, and suspected that on metamorphosis the lateral line organs 
were eliminated from the make-up of the species. 

F. J. Cole. 



BIBLIOGRAPHY OF THE HISTORICAL INTRODUCTION 

1,4, 10,20,35,52,54,58, 103, 104, 105, III, 112, 116, 122, 123, 124, 
126, 127, 129, 132, 133, 147, 148, 149, 150, 156, 157, 158, 159, 187, 206, 
242, 264, 265, 266, 293, 324, 335, 336, 338, 342, 346, 347, 348, 349, 406, 
4i3> 4H> 4i7> 4195 445, 44^, 45o, 45i, 460, 461, 466, 480, 481, 493, 501, 
502, 503, 504, 536, 540, 541, 547, 548, 552, 553, 554, 555, 560, 564, 582, 
586, 587, 599, 600, 601, 603, 608, 623, 636, 637, 638, 639, 640, 641, 642, 
643, 644, 649, 652, 665, 666, 674, 675, 691, 702, 703, 704, 723, 724, 743, 
744, 748, ISS, 804, 822, 823, 824, 835. 



I 

TAXONOMY, LIFE-HISTORY, AND GENERAL 

TAXONOMY 

1 . Characteristics of Class, Order and Sub-Order. 

The Salamander, by virtue of its naked glandular skin devoid of 
scales, its aquatic larva, its three-chambered heart (two auricles and 
one ventricle), two occipital condyles and the absence of any amnion 
or allantois around the developing embryo, is placed in the Class 
Amphibia among the vertebrates. 

The retention of the tail by the adult and the possession of limbs 
of approximately equal size confine it to the Order Urodela or 
Caudata^ while the complete metamorphosis, and the backward 
extension of the pre-vomerine teeth to well behind the internal 
nares, limit it to the Sub-order Salamandroidea.^ 

2. The Family Salamandridae. 

This Sub-order comprises three Families^ the Salamandridae^ the 
Amphiumidae^ and the Plethodontidae (mostly American Urodeles). 
The Salamander belongs to the first of these families, which is 
characterized mainly by the shape of the pre-vomerine teeth, by the 
opisthocoelous vertebrae, and by one or two other skeletal features 
of minor importance. The Family Salamandridae is again roughly 
divided into 'Salamanders' and 'Newts', the former being completely 
terrestrial in the adult stage while the latter show strongly marked 
aquatic tendencies throughout life. All are completely caduci- 
branchiate in the adult phase. The family comprises some eight 
genera, namely, Salamandra, Chioglossa^ Triturus (Triton), 'Tyloto- 
triton^ Pachytriton^ Pleurodeles^ Euproctus^ and Salamandrina, 

3. The Genus Salamandra. 

The Genus Salamandra is distinguished by the 'S' shape of the 
backward prolongations of the pre-vomerine teeth, together with 
the subcircular tongue free at the edges, and the prominent 'para- 
toid' cutaneous glands behind the eyes. The hands are four-fingered 
and the feet are five-toed, while the tail is rounded in section, and 
there is no fronto-squamosal arch. The genus contains four species 
^ The classification of Noble (193 1) is followed here, 

4038 B 



2 THE SALAMANDER 

of which two are European and two Asiatic. The two former are 
well known and are Salamandra salamandra (better known as S. 
maculosa)^ the Spotted Salamander, and S. atra, the Black or Alpine 
Salamander. The Asiatic species are S. caucasica and S. semenovi, 
both of which inhabit mountainous districts and are known from 
a few specimens only — particularly the last named. It is to the first 
named of these four species that the present work is confined, but a 
brief account of the general characteristics of the other three may not 
be out of place, and they will accordingly be discussed in the reverse 
order to that given above. 

4. Salamandra semenovi. 

Salamandra semenovi was first described by Nesterov in 1 9 1 6. It 
is of approximately the same size, or perhaps slightly smaller than 
the Spotted Salamander, and inhabits the mountains of Kurdistan 
and Perso-Turkey at an altitude of about 5,000 ft. (PI. XXV, fig. 83). 
Nothing appears to be known of its anatomy. It is characterized by 
the markings which are small and greenish-yellow. 

5. Salamandra caucasica. 

Salamandra caucasica was first described by Waga (1876) under 
the name Exaeretus caucasicus and its inclusion in the genus Sala- 
mandra is due to Boulenger (1882). It is rather more slender than 
the Spotted Salamander, while its tail has a very much greater 
relative length and contains some fifty-three vertebrae against 
twenty-two to twenty-five for salamandra. The markings are deep 
orange in colour on a pure black ground and show a strong resem- 
blance to those of S. salamandra^ forma typica (see p. 1 3). The male 
is characterized by the possession of a prominent dorsal hook at the 
base of the tail. As its name implies this species inhabits the Cauca- 
sian mountains, to which it appears to be confined (PI. XXV, fig. 8 3). 

6. Salamandra atra. 

Salamandra atra has been known from early times. It is restricted 
to the European Alps over a vertical range extending from about 
2,500 to 9,000 ft. (PL XXV, fig. 83). It is rather smaller than 
S. salamandra and is distinguished from it by the absence of any 
yellow or orange markings to interrupt its uniform black colour, and 
by the relatively shorter tail. Normally this species bears only two 
young at a time, and these are devoid of gills and are not deposited 
in water as in the case of the other species, but are fitted from the first 
for a terrestrial existence. Schwalbe (1896) gives an excellent 



TAXONOMY, LIFE-HISTORY, AND GENERAL 3 

account of the 'intra-uterine' development of this species, and shows 
how the gills have become adapted to effect a gaseous exchange with 
the maternal capillaries in the 'uterine' wall and to absorb nourish- 
ment from the surrounding yolk mass formed by the degenerating 
eggs which fail to develop. 

7. Salamandra salamandra. 

The name Salamandra is of ancient origin and is probably derived 
from the Arabic samandar^ by which term the animal is still popularly 
known in Arabia and Persia. It was applied to this species as a 
generic term in quite early times (see Historical Introduction). 
Nevertheless the tenth edition of the Linnaean Systema Naturae 
(1754) from which all modern nomenclature starts — or is supposed 
to start — describes the animal under the name Lacerta salamandra. 
In 1768 Laurenti quite rightly removed it from the genus Lacerta 
and re-established the older Salamandra^ but unfortunately the same 
author also at the same time, and without any justification, changed 
the specific name to maculosa^ and this name seems to have become 
inseparably associated with the animal, since it still persists, although 
by the rules of priority the Linnaean species should stand. A number 
of authors have pointed out this discrepancy. One of the earliest of 
these is Lonnberg (1896), and one may well echo his lament when, 
in describing the Linnaean type-specimens of Birds, Batrachians, 
and Fishes in the Zoological Museum of the Royal University in 
Upsala, he says under the heading Salamandra maculosa^ Laurenti, 
*It would seem provoking to change such an old name, but according 
to the rules the name ought to be Salamandra salamandra (Linnaeus)'. 
The species is characterized by the shape of the pre-vomerine teeth 
coupled with the yellow markings on the body. A detailed considera- 
tion of the external characters and chief varieties of the species is 
given in the next section (p. 1 3). 

LIFE-HISTORY 

The general features of the life-history and development of the 
Salamander, such as its habit of bringing forth fully developed 
larvae (instead of eggs), which subsequently pass through an aquatic 
phase before metamorphosing into the terrestrial phase characteristic 
of the adult, have been fairly well known since early times. Never- 
theless some puzzling details remained unexplained for a long time, 
and, even now, further observations concerning fecundation would 
not be superfluous. 



4 THE SALAMANDER 

I. Fecundation and Fertilization. 

One of the earliest writers to make observations on the develop- 
ment of the Salamander was von Schreibers (1819 and 1833), who 
correctly observed the method of deposition of the young, but was 
mistaken as to the mode of impregnation of the female by the male 
which he thought took place in water. Rusconi (1839) was the first 
to observe the correct method of copulation, which normally takes 
place on land, although he was quite at a loss to explain how 
segmenting eggs came to be present in the oviducts before copulation 
had taken place — a paradox which took many years to solve. 
Rusconi's paper seems to have been overlooked by most subsequent 
observers. The discovery of the receptaculum seminis in the female 
by von Siebold (1858), although only indirectly concerned with 
fecundation, nevertheless proved to be of great importance in 
furnishing the ultimate explanation of the mystery which baffled 
Rusconi. The next observer to discuss the mating of Salamanders 
was Zeller (i 890-1), who confirmed the observations of von Siebold 
that one act of coition apparently sufficed to fertilize more than one 
batch of eggs. It was thought that the function of the receptaculum 
seminis was to hold a reserve of sperms for future use, i.e. that it was 
analogous to the receptaculum seminis of an insect. This view was 
also supported by Paratre (1894). Zeller believed that copulation 
took place at the time of deposition of the young because he found 
that the receptacula of the females were then full, and he also found 
spermatophores in the water with the larvae. Five years later 
Schwalbe (1896) published a very careful and detailed account of 
the life-history and development of the two European Salamanders, 
prefaced by an equally clear and lucid review of the previous 
observers. Schwalbe's account is the first really adequate statement 
of the facts, which subsequent observers have confirmed, but to which 
they have added only minor details. Schwalbe finds that locality and 
external conditions have some influence on the actual times of deposi- 
tion, copulation, &c., but that on the average fully developed em- 
bryos are found in the oviducts of the females from May until June, 
and then again in October and December. He also showed that 
sperms may be found in the receptacula in May but not in June^ and 
then again from July to November, copulation having taken place 
during July. Thus he established the important fact that the sperms 
must winter in the receptaculum seminis of the female. He also showed 
that fertilization occurs high up in the oviducts. The key to the 
mystery which Schwalbe has thus provided is that the sperms received 



TAXONOMY, LIFE-HISTORY, AND GENERAL 5 

by any female in a given season do not immediately pass into the 
oviducts to fertilize the eggs, but pass into the receptaculum seminis 
and remain there through the winter to fulfil their ultimate purpose 
In the following May or June. Thus the segmenting eggs which 
Rusconi found to be present in the oviducts at the time the sexes were 
pairing were those fertilized by the sperms received during the 
amplexus of the previous year. 

An excellent account of the pairing is given by van Leeuwen 
(1907), who observed the act in his vivarium. He states that it may 
occur at any time from June to September (or even in October if the 
weather is warm) and he only saw it occurring in the evening, and on 
land. Van Leeuwen describes the male pushing his nose under the 
hind-legs of the female and working his body forwards until his head 
is directly under hers. He then embraces the fore-limbs of the female 
with his own fore-limbs and, with a quiver of the body, deposits a 
spermatophore on the moss. The male then rotates the posterior 
part of his body through 90° but still retains his hold of the female, 
who then lowers her cloaca on to the spermatophore and receives 
it therein. The spermatophores are described by the same author 
as transparent yellowish pyramids of gelatinous consistency, about 
8-10 mm. high by 4-6 mm. broad, and are deposited with the 
sharply pointed apex directed vertically upwards. 

Kammerer (i 907) agrees with van Leeuwen, but has also seen the 
male approximate his cloaca to that of the female and pass the sper- 
matophore directly into it. He also claims to have seen copulation 
occur in water as well as on land, and says that then no such approxi- 
mation of the cloacae occurs. His observation remains unsupported 
by any other observer. 

The segmentation of the o.^^ and the early development were 
first studied by Kupffer (1879), Benecke (1880), and Gronroos 
(i 895 and 1898), the work of the last-named being the most detailed. 
In spite of the large amount of yolk present, segmentation is holo- 
blastic but is unequal. 

To summarize the essential facts that have been thus laboriously 
unravelled we may say that the larvae are born in early spring from 
April to May or June, while they have been observed as early as the 
1 6th of January (Paratre, 1 894), the exact period depending on local 
conditions. Ovulation then occurs, and the eggs thus released are 
fertilized high up in the oviduct by sperms which have wintered in 
the receptaculum seminis of the female. Shortly afterwards, during 
July to September, pairing takes place, and the sperms then received 
enter the receptaculum seminis and are stored for use in the following 



6 THE SALAMANDER 

year. The amplexus thus normally occurs while there are seg- 
menting eggs in the oviduct, but whether this is always the case is 
uncertain, neither is there any evidence as to whether the sperms are 
able to fertilize the eggs immediately they leave the male, or whether 
the period of rest in the receptaculum seminis is essential, and if so 
what changes, if any, they then undergo. The embryos developing 
from the eggs fertilized in June have attained full maturity by the 
autumn, but they are normally carried over in the 'uterus' until the 
following spring, i.e. for a total period of ten to twelve months. 
Under favourable circumstances, however, two batches of larvae may 
be deposited in the same year, and both copulation and deposition of 
larvae may occur at almost any period of the year except during the 
winter. 

2. Deposition of Larvae. 

The female deposits the larvae in water, usually in the springs and 
small streams which abound on the lower slopes of the mountains of 
central Europe. She does not enter the water bodily, since the adult 
Salamander is unable to swim and would speedily drown if totally 
immersed in water beyond its depth, but merely remains at the 
water's edge with her cloaca and tail submerged. Deposition may 
occur at intervals over a period as long as two days. The total num- 
ber of larvae deposited by any female during this period varies from 
twelve up to as many as seventy-two (Fatio, 1872). Such high 
numbers are, however, rare and the average may be taken in the 
region of thirty, and they occur in approximately equal numbers 
in each oviduct. Dead larvae in various stages of development, and 
of degeneration, are by no means uncommon and are deposited 
together with the living ones. The embryos remain within the 
vitelline membrane until the very moment of deposition, and may 
in fact actually be 'laid' within it, but normally the act of extrusion 
ruptures the membrane and allows the larva to swim free. The 
empty membrane is then pushed out from the cloaca by the suc- 
ceeding larva. The Salamander is thus ovoviviparous rather than 
strictly viviparous, that is to say, it lays eggs but the eggs have pre- 
viously developed within its body to hatching-point before their 
deposition. The embryo feeds exclusively on ^^o^ yolk. Kaufman 
(19 1 3) finds that the number of degenerating larvae is proportional 
to the length of the 'uterus' and the number of embryos, and she 
therefore concludes that the death of these larvae is due to pressure. 
She also finds that the weight of a full-grown embryo is i -6 times the 
weight of the egg from which it developed, and since it is independent 



TAXONOMY, LIFE-HISTORY, AND GENERAL 7 

of the mother and does not leave the vitelline membrane, she con- 
cludes that the increase in weight is due to the absorption of water. 
Monsters are not uncommon (cf. Kaufman, 1913; Stadtmiiller, 
1930, and others). Deposition often occurs at night, and Schreiber 
(19 1 2) supposes that the act must often be fatal to the females, 
since he frequently finds dead adults near the water containing 
young. 

3. The Larvae. 

The larvae are about 25-30 mm. long at birth, of greyish-brown 
colour dorsally and greyish white ventrally, flecked with arachnoid 
patches of black pigment. They possess four fully developed limbs, 
and external gills, and are adequately fitted for an independent free 
existence. They are very active and commence to feed at once. 
Their normal diet consists of small Crustaceans, e.g. Daphnia, 
Cyclops, and aquatic insect larvae, worms, &c. In captivity they 
flourish well on the Enchytraeid worms sold by aquarist dealers if their 
natural food is not available. They take these from forceps with 
great avidity, and will also take small earthworms cut up, or small 
shreds of meat, provided that the food offered is made to move. The 
hyoid and four branchial arches are represented in the visceral skele- 
ton, and they possess four gill-slits in addition to the external gills. 

4. Metamorphosis. 

They grow slowly and attain a length of ^^-^^ mm. in about i^-K^ 
months, when they metamorphose and leave the water. The chief 
external changes noticeable just prior to metamorphosis are the 
appearance of yellow pigment, the approach to adult coloration, 
and the dwindling of the external gills. The loss of the swimming 
membrane on the tail follows immediately, and the head assumes the 
adult shape through the advent of the paratoid glands. At the same 
time profound internal changes have been going on. The cartilages 
and muscles of the two most posterior branchial arches are normally 
lost entirely, but the arteries and nerves remain. The tongue also 
changes completely, the primary larval tongue being replaced by the 
secondary adult structure (cf. p. 263), while the eye has had to become 
adapted to see in air instead of in water. It is probably on account 
of these considerable changes in the organs associated with gustation 
and deglutition that the larvae show great reluctance to eat for a short 
time just preceding and immediately following metamorphosis. Of 
the internal factors which affect the inception and rate of metamor- 
phosis the secretion of the thyroid glands takes the first place, while 



8 THE SALAMANDER 

the principal external factor is temperature, which has a marked 
positive influence. Larvae born in the autumn may remain unchanged 
right through the winter. 

The Salamander does not reach sexual maturity until 4-5 years 
after metamorphosis, by which time it will have attained a length of 
rather less than 200 mm. Ahl (1930) gives 285 mm. as the maxi- 
mum length, but this must be regarded as exceptional, 150-200 
mm. being much more common. 

5. Habits of the Adult. 

The adults are rarely seen during the day-time except after a warm 
thunder shower, when they may suddenly appear in vast numbers. 
Otherwise they spend the day under stones, in holes, or under moss or 
fallen tree-trunks, and come out at dusk to feed. Their normal diet 
consists largely of insects, e.g. beetles and beetle larvae, and also 
slugs and worms, but Jacobshagen (19 14) reports finding con- 
siderable quantities of moss and other plants in the stomachs of some 
individuals. Schreiber (19 12) says that Salamandra salamandra 
usually occurs on sandstone and marl, and is rarely found on sandy 
or chalky bases. They frequently occur in pine forests on the lower 
slopes of the mountains below 3,000 ft., and rarely up to 4,000 ft. 
During the winter they congregate in large numbers, often several 
hundreds together, under an overhanging rock, or in a hollow tree- 
trunk or some similar place. They get sluggish in cold weather, but 
their hibernation is only slight, and they not infrequently come out 
during a warm spell. In general they favour moist places and a 
humid atmosphere with moderate temperature, and shun the sun- 
light. No data are available concerning the length of life in nature, 
but records have been made which show that individuals have lived 
in captivity for as long as 1 8 and 20 years (Ahl, 1 930 ; Mehlsheimer, 
1911). 

6. Geographical Distribution. 

The geographical distribution of the species is fairly well indi- 
cated in the next section which deals with the several varieties. 
Schreiber (19 12) gives 53j° N. as the northern limit, i.e. a line 
drawn from Lauenburg through Liineburg and Bremen to Olden- 
burg. From here it spreads south-westwards through Holland and 
Belgium, over France, Spain, and Portugal. Southwards it is found 
all over Germany and Austria, in Italy and on the Balkan peninsula, 
in Greece and Turkey. It does not extend eastwards of the Carpa- 
thians. Isolated instances of its occurrence have been reported from 



TAXONOMY, LIFE-HISTORY, AND GENERAL 9 

as far south as Palestine, Algeria, and Tunis. Very favourite locali- 
ties are the Hartz mountains and the Heidelberg district of Germany, 
as well as many parts of the Black Forest, where it is very common 
(PL XXV, fig. 83). 

Good accounts of the life-history and habits of the species have 
been given by Gadow ( 1 90 1 ), Werner ( 1 9 1 2), and Schreiber ( 1 9 1 2), 
the last named being particularly detailed. 

7. Parasites and Diseases. 

The most troublesome disease to which Salamanders are subject 
in captivity is that caused by a fungus, Monilia batrachea^ Scott, 
which first attacks the lips (the hands and feet are first attacked in 
Frogs and Toads), causing open sores to appear. If unchecked 
the fungus finally ramifies throughout the body, even to the 
internal organs, e.g. liver, kidneys, &:c., and is fatal. It is very con- 
tagious and the only effective method of sterilization of vivaria, 
apparatus, &c., is the painter's blow-lamp, or if accumulator jars are 
used (and these make by far the best vivaria and are least likely to 
become infected) repeated scrubbing with absolute alcohol has 
proved effective. The disease is curable, if taken in time, by painting 
the parts affected with 'Mercurochrome' applied with a camel-hair 
brush about three times a week. Cases must of course be isolated 
immediately. I The aetiology of the disease has been worked out by 
Scott (1926). 

Of the animal parasites of the Salamander the Protozoa are 
represented by a coccidian, Eimeria salamandrae^ Steinhaus, which 
infests the epithelial cells of the gut. Among the Helminths repre- 
sentatives of the Trematodes, Cestodes, and Nematodes all occur. 
Von Linstow (1878-89) gives the following: 

TREMATODES. 

Distomum cygnoides^ Zed. in the urinary bladder 
Distomum crassicolle^ Rud. in the intestine and rectum 
Distomum endolobum., Duj. in the intestine 
Distomum glabrum^ Creplin. in the intestine 

nematodes: 

Oxysoma brevicaudatum^ Zed. in the intestine 
Strongylus auricularis^ "Zed. in the intestine 
Nematoxys commutatus^ Rud. in the intestine 

' The author is indebted to the late Dr. Joan Procter, formerly Curator of Reptiles 
at the Zoological Society's Gardens, Regent's Park, London, for the practical notes 
on this fungoid disease, which are the outcome of experiments carried out in her 
laboratory. 



lo THE SALAMANDER 

Among the Cestodes Taenia dis-par has been reported by Fuhrmann 
(1896). Von Linstow's nomenclature is of course obsolete. For 
example his Oxysoma hrevkaudatum is almost certainly the worm 
now known as Aplectana hrevicaudata^ and Nematoxys commutatus is 
probably Cosmocerca commutata (Dies.), since specimens referable 
to these genera have been found during the present investigation, 
together with the trematode Brachycoelium salamandrae Frolich, which 
certainly represents v. Linstow's Distomum crassicolle. Ecto-parasites 
seem to be entirely absent. 

General Literature. 

So far there have been two monographs devoted to the anatomy 
of the Salamander — the first by Funk (1827) which has been dealt 
with in the Introduction, and the second by Rusconi, commenced in 
1839 but not published until 1854, after the death of the author. 
These two works must be considered as to a large extent complemen- 
tary. Rusconi corrects some of Funk's mistakes — notably with 
regard to the vascular system — and fills in some of his omissions. 
Of these latter the most noteworthy is the account of the anatomy of 
the larva and of the changes which take place during metamorphosis. 
He also deals at some length with the lymphatic system, with the 
life-history, and with the habits of both larva and adult — topics 
scarcely touched upon in Funk's account. 

Of the other literature of a general nature must be mentioned the 
'Amphibia' in Bronn's Klassen und Ordnungen des Thierreichs^ written 
by C. K. Hoffmann (1873-8), in which frequent references are 
made to Salamandra, but which is rather a compilation of existing 
knowledge than an original contribution. Most of the original 
material it does contain had been published previously elsewhere. 
Of a similar nature is Noble's recent work on the Biology of the 
Amphibia (1931). Osawa's account of the anatomy of Crypto- 
branchus (1902) and Gaupp's Anatomie des Frosches (1896) have 
been found to be of considerable use for comparative purposes. 

BIBLIOGRAPHY OF SECTION I 

3, 38, 50, 68, 72, 73, 74, 75, 125, 126, 153, 155, 193, 197, 205, 206, 217, 
222, 241, 246, 252, 253, 271, 281, 282, 283, 300, 322, 358, 366, 373, 382, 
384, 386, 387, 402, 416, 424, 425, 447, 461, 463, 464, 465, 468, 492, 51 1, 
512, 521, 522, 542, 558, 559, 580, 585, 588, 604, 621, 624, 651, 662, 663, 
664, 673, 674, 675, 687, 690, 696, 71 1, 712, 730, 734, 760, 761, 767, 768, 
776, 111, 778, 779. 780, 799. 820, 832, 833, 834. 



II 

EXTERNAL FEATURES AND VARIETIES 

EXTERNAL FEATURES 

The external form of the Salamander is one of the simplest which it 
is possible for a terrestrial, tetrapodous vertebrate to have. The 
head is broad and flattened like that of the Frog, but the snout is 
rather more rounded. The eyes stand up prominently from the sur- 
face of the head, while the small external nares may be seen in front 
of them on either side of the snout. The naso-lacrymal duct opens to 
the exterior at the anterior corner of the eye by means of a fine pore 
which is visible only by the aid of a lens. The apparent gape (see also 
p. 2 62) extends to a little behind the posterior corner of the eye, and the 
upper jaw slightly overhangs the lower. On the dorsal surface of the 
head, just behind the eyes, is a pair of large reniform protuberances, 
the paratoid glands (see also p. 292). They are very characteristic 
of the animal. There is no true neck but there is a slight narrowing of 
the contour behind the head, due largely to these paratoid glands, 
while ventrally there is a distinct fold in the skin marking off the 
head from the pectoral region. This is th^gularfold^ and represents the 
line of fusion between the larval gill cover, or operculum, and the skin. 

The trunk is roughly cylindrical, while the sides of the body pre- 
sent a wrinkled appearance due to the fact that the skin is closely 
adherent to the myosepta but free in between. 

The tail tapers, at first rapidly, and then more gradually, and 
finally terminates in a blunt point. It is very slightly flattened laterally. 

Extending from behind the head to the tip of the tail, in the mid- 
dorsal line, is a double row of cutaneous poison glands of a similar 
nature to those forming the paratoids. These series of venom- 
secreting glands are dealt with in more detail on p. 292. 

The limbs are relatively small, particularly the fore-limbs. Thus 
mechanically, as well as in many other ways, the Salamander is truly 
intermediate between the piscine structure, where practically the 
whole of the animal — apart from the skeleton — is composed of 
segmental body muscles and viscera, and that of the typical land 
vertebrate, in which the muscles associated with the limbs tend to 
predominate over those of the trunk and tail. 

The total body length of adult specimens, including the tail, 



12 THE SALAMANDER 

varies from about 120 mm. to 170 mm., according to age. Of this 
the tail, or post-sacral region, accounts for rather less than one-half, 
the proportion being something like 75 : 80. The males tend to be 
rather smaller than the females, but the sexes are more easily dis- 
tinguishable externally by the appearance of the cloaca, which is 
quite flat in the female, but swollen and tumid in the male. This 
difference is more marked in the breeding season, but a little practice 
makes it readily distinguishable at any period. 

The gait of the Salamander is very blundering and clumsy, and 
when the fore-foot is lifted from the ground the whole body is tilted 
so as to assist the movement. Then the limb having been moved 
forwards, the animal 'flops' over on to that side and repeats the pro- 
cess on the other. In this way the movement at the shoulder-joint, 
in a vertical plane, is very slight. The pelvic limb shows rather more 
movement in this direction — nevertheless there is a strong tendency 
for the back of the foot to be dragged along the ground as it moves 
forward. The sector described by the femur in a horizontal plane 
lies almost entirely anterior to the transverse plane across the aceta- 
bula; in other words, when the femora are fully retracted they are 
approximately at right angles to the trunk — while when they are 
drawn forwards to their fullest extent the limb lies close alongside the 
body. In spite of its apparent clumsiness when seen under ordinary 
conditions stalking around a vivarium, the Salamander can at times 
move with astonishing rapidity, this being particularly so with young 
metamorphosed individuals. Seizing of prey, too, is performed at 
lightning speed, and, if the range is a long one, the animal may jerk 
its whole body in the direction of the object to be seized, thus assisting 
the movements of its head. The Salamander is quite capable of seizing 
a passing 'blue-bottle' in flight, and has been observed to do so. The 
climbing powers of some individuals are also rather astonishing, and 
it has been found necessary to keep a piece of wire-gauze over the 
top of a glass accumulator jar some 18 inches high, which was being 
used as a vivarium, in order to prevent the escape of the inmates. 
This they attempted by climbing up the angles, chimney fashion. 

The colour markings are subject to great variation within certain 
limits, some individuals being almost entirely yellow, while others 
again are nearly black. Some have thought that the intensity of 
colour is controllable by varying the environmental conditions, e.g. 
background, temperature, and humidity, and further that such 
adaptive changes are transmissible to future generations. This ques- 
tion is one of experimental zoology rather than of anatomy, and it is 
not proposed to discuss the matter here. The works of Kammerer, 



EXTERNAL FEATURES AND VARIETIES 13 

MacBride, Knoblauch and others given in the bibliography should 
therefore be consulted by those interested in this subject. 

VARIETIES 

In spite of so much variation in the relative proportions of the 
yellow and black pigments, three main and fairly constant varieties 
stand out clearly. They are based on differences of pattern, and 
appear to be correlated, to a very large extent, with geographical 
distribution. These varieties have been carefully investigated by 
E. G. Boulenger (191 1) and the following description is a resume 
of his work (cf. Frontispiece). 

1. Forma Typica. 

The first of these varieties may be called the forma typica, since it 
is the type on which Laurenti first established the genus. In this 
variety the black pigment nearly always predominates over the yel- 
low, while the latter pigment is disposed over the body as markings 
of various shapes, often in three or five alternating series, or with 
a median series forming a sinuous or zigzag vertebral stripe. Very 
exceptionally the dorsal markings may appear to form two longi- 
tudinal series, but in such a case they never hang together in regular 
chains continuous with the spots on the paratoids. The upper eyelids 
and paratoids are usually entirely, but sometimes only partially, 
yellow, and these usually remain distinct from each other, but they 
may fuse. A yellow spot occurs nearly constantly above the angle 
of the mouth, while the snout is usually entirely black. There are 
sometimes a few markings in the gular region, but they may be 
absent. The ventral surface is either quite black or marked by a few 
small spots. The limbs are black with a characteristic and constant 
yellow patch on the proximal portion of the dorsal surface of the 
arm and thigh, and a second similar patch may or may not occur 
on the distal portions of the limbs. The spots on the tail may be 
single or paired, or they may form a stripe. The ventral surface of 
the tail is nearly always black. The colour of the markings varies 
from chrome-yellow to deep orange. 

With very few exceptions all specimens occurring to the east and 
south of a line drawn through the Erz mountains, the Danube, the 
Alps, and the Rhone belong to this form (PL XXV, fig. 84). 

2. Variety Molleri. 

The second form occurs in the Spanish peninsula. It is the 
variety molleri of Bedriaga (i 896), and includes Seoane's var. galliaca 



14 THE SALAMANDER 

(Seoane, 1884). This might almost be described as a sub-variety 
of the forma typica, since a practically uninterrupted series can be 
traced between the two forms, both of which occur on the Peninsula. 
Two descriptions of this type may be quoted. The first, by Bedriaga, 
states that the colour and pattern of this variety is rather variable. 
The ground colour is usually greyish-brown, sometimes of a more 
dirty grey, sometimes more brownish-black or even black, and is 
broken up above and below by pale yellow spots with the addition 
of grey or greyish-brown spots, into which the yellow passes gradu- 
ally. The yellow spots on the sides of the body, the limbs, the tail, 
the paratoids, the throat and on the eyelids, are as if powdered over 
with red dust, or washed with red, or even blood-red. The throat 
may sometimes be a deep red colour, and the dorsal spots show here 
and there a red dot. The yellow spots are very variable both in 
number and size. The second description by G. A. Boulenger 
concerns some specimens kept at the Zoological Gardens. He 
says that some specimens were black, variegated with various tints 
of grey, brown, pale yellow, and crimson. The latter colour was 
particularly conspicuous on the upper eyelids and the paratoid glands, 
the base of the limbs and the throat, but it appeared as small patches 
within the area of the more or less irregular pale yellow spots with 
grey centres which were dispersed very irregularly on the body and 
tail. One of the specimens was pale olive-grey above, and on the 
sides freckled with black and pale greenish-yellow spots, the black 
appearing as an irregular vertebral stripe, a dorso-lateral stripe, and 
bars on the flanks. The upper eyelids, the paratoids, and the throat 
were claret-red. The coloration of such a Salamander has a lichen- 
like appearance more suggestive of assimilation to surroundings than 
of warning to enemies. E. G. Boulenger discussing this type says: 

'On careful examination of the red and grey markings in these specimens 
I have come to the conclusion that these are due, not to special pigments in 
addition to or in combination with the yellow, as has been supposed, but to 
the absence of pigment, the pigmentless flesh highly flushed with blood being 
exposed on certain patches and the grey colour resulting from the absence of 
yellow combined with a small quantity of black pigment.' 

Forma typica also occurs all over the Peninsula side by side with this 
variety. 

3. Variety Taeniata. 

The variety which is commonly imported into this country and on 
which the present investigation has been carried out is the variety 



EXTERNAL FEATURES AND VARIETIES 15 

taeniata, so called on account of the arrangement of the dorsal 
markings in two parallel series continuous with the patches on the 
paratoids, which not infrequently form two uninterrupted stripes. 
This peculiarity distinguishes it unmistakably from the other two 
types. Even when the two stripes are only quite fragmentary they 
retain their individuality and parallel disposition, not encroaching 
over the mid-dorsal line except occasionally above the occiput and in 
the posterior region of the body, where they may be connected by 
cross bars. Another point of difference is that the yellow markings 
are usually present on the snout, while the spot situated above the 
angle of the mouth in forma typica is frequently absent in taeniata. 
On the limbs black usually predominates, and there is a similar 
characteristic yellow patch in each variety near the base of the arm 
and thigh, but the second patch on the fore-arm and leg which is 
variable m forma typica is constant in taeniata^ and usually larger than 
in the typical form. Not infrequently yellow is in excess of the black 
and often extends to the ventral surface of the tail, which is rarely 
the case in the other forms. 

The colour varies from sulphur- or lemon-yellow to a deep orange. 
Fr. von Schweitzerbarth (1909) obtained some specimens near 
Stuttgart with markings almost vermilion red in colour. She named 
t\\Qm'var. coccinea but they clearly represent only individual peculiari- 
ties. Albinism has been reported in several cases (see Schreit- 
miiller, 1 909, and Wolterstorff, 1909). 

The habitat of var. taeniata is restricted — with the exception of 
one doubtful specimen from Rome — to France, Spain, Portugal, 
south Holland, Germany, and Switzerland (PI. XXV, fig. 84). 

Excellent accounts of the external characteristics as well as of the 
life-history and ecology of the Salamander have been given by 
Bruno Diirigen (1897) and by Schreiber (19 12). The arrangement 
of the varieties adopted by these two authors is different in each case 
and also differs from that adopted here, the last-named author 
describing as many as thirteen varieties and sub-varieties. Such 
elaboration seems unnecessary, since it concerns only the relative 
amounts of the yellow and black pigments present and whether the 
spots are confluent or discrete, and so on. 

BIBLIOGRAPHY OF SECTION II 
30, 31, 50, 67, 73, 76, 151, 155, 197, 217, 219, 301, 312, 357, 359, 360, 
362, 363, 395, 436, 468, 558, 673, 677, 683, 688, 689, 692, 693, 710, 730, 
734, 776,778, 779> 784, 821. 



Ill 

THE SKELETON 

The literature dealing with the osseous skeleton of the Salamander 
is very extensive, so that anything approaching a complete resume 
would be too lengthy to include here; neither would any particular 
advantage be attained by so doing. All the chief original contribu- 
tions to our knowledge of the structure and homologies of the various 
elements, as well as those accounts definitely needing correction, are, 
however, referred to in the text. 

As an aid to the study of the skeleton in general, and particularly 
in differentiating between cartilage and bone, it is advantageous to 
steep the macerated and semi-cleaned bones in a saturated solution 
of alizarin in alcohol for a few hours. Subsequent immersion in 70 
per cent, alcohol will extract the dye from the soft tissues and from 
the cartilage, and leave the osseous parts stained a bright magenta. 

, THE AXIAL SKELETON 

I. THE VERTEBRAL COLUMN (PLATE III). 

1. General. 

The total number of vertebrae is somewhat variable, the most 
variable region being the tail, the average being about forty. Funk 
(1827) gives forty-two as a definite number, but a specimen with as 
few as thirty-eight has been counted, while List (1883) mentions an 
example having forty-three, and Hoffmann (1873-8) gives a total 
of forty-four. Four regions are distinguishable, namely, the cervical 
with one vertebra, the trunk with thirteen to fifteen, the sacrum with 
either one or two vertebrae, and the caudal region with some twenty- 
two to twenty-five. Funk further distinguishes a thoracic region, 
but the term 'thorax' is a misnomer when applied to an Amphibian. 

2. The Cervical or Atlas vertebra. 

The Cervical or Atlas vertebra (Figs. 10 and 11) differs rather 
widely from the general pattern. The centrum is almost non-existent, 
being present only at the posterior end at the articulation with the 
second vertebra. The neural arch is high and bears a well-formed 
neural spine (nl.sp.). It slopes off obliquely at the anterior end so as 
to leave a gap between the occipital region of the skull and the arch 



THE SKELETON 17 

of the vertebra which is closed by a strong tough membrane. The 
anterior end bears two large concave facets for articulation with the 
occipital condyles (fa. con.), while there is also a well-marked median 
process, the so-called odontoid process (pr.o.), which fits into the fora- 
men magnum of the skull and articulates, by means of two small 
lateral facets, with the bony side-walls of this foramen. Each side of 
the neural arch is perforated by three foramina, one near the anterior 
end for the emergence of the first spinal nerve (fo.n.sp.I.), another 
near the posterior end for the second spinal nerve (fo.n.sp.II.), and 
the third near the middle, the. foramen nutritium (fo.nt.), for the passage 
of blood-vessels. The foramen for the second spinal nerve may not 
always be completely surrounded by bone but may merge into the 
intervertebral foramen, which is normally closed by a membrane. 

The homology of the first cervical vertebra of Urodeles has caused 
a considerable amount of discussion. The earlier workers, e.g. Hoff- 
man, Hyrtl, Wiedersheim, &c., considered it to be of a double 
nature, since they regarded the 'odontoid' process as the centrum 
of a vertebra which possessed no neural arch, and they thought 
that the 'atlas' probably represented the second vertebra of Amniota. 
On this assumption they call the first spinal nerve N. spinalis II. 
Parker (1882), while agreeing as to its multiple nature, regards the 
vertebra as representing a portion of the occipital region of Amniota, 
and therefore thought that the second vertebra of Urodeles was the 
equivalent of the atlas of Amniotes, and with this view Stohr (1880)^ 
agrees. Albrecht (1878) supposes the vertebra to be the homologue 
of the pro-atlas of Reptiles, while Peter (1895) agrees with Parker 
and Stohr that it corresponds to the occipital segment of higher 
vertebrates, but disagrees with them in that he thinks that it is a 
single vertebra, and not two fused together. In favour of the older 
view Froriep (19 17), working on the development of Salamandra 
atra^ claims that the first vertebra comprises vertebrae I and II and 
that the first spinal nerve of the adult is the morphological second, 
since at one stage he finds a transitory vestige of a nerve emerging 
between the occiput and the vertebral column. More recently still 
Mookerjee (1931) has reinvestigated the problem and finds that 
there is a pair of intercalated cartilaginous arches between the occi- 
pital arch and the atlas (first) arch proper, and further that the 
occipital joint arises from these arches, the cups fusing with the atlas 
arch and the balls with the occipital arch. He also showed that the 
'odontoid' process is the intervertebral cartilage of this arch. The 
cups and the 'odontoid' process Mookerjee terms the pro-atlas. 

^ Stohr deals only with Triton {Zeitschr. f. zviss. ZooL, Bd. 33, pp. 477-526). 

4038 c 



1 8 THE SALAMANDER 

To sum up then, there would seem to be some truth in each of the 
two earHer and apparently conflicting views, since the first vertebra 
of the adult has a double origin while a part of the intercalated arch 
also fuses with the skull. 

3. The Trunk Vertebrae. 

The trunk vertebrae are all essentially similar. The actual one 
figured (Figs. 13—17) is the last pre-sacral and may be taken as 
typical of the whole group. The centrum (cen.) — or corpus — is well 
formed and rather skittle-shaped. It is oval in cross-section and very 
strongly opisthocoelous, the posterior concavity tending to become 
rather deeper towards the ventral side. The ends of the centrum 
remain cartilaginous, but the body is ossified and hollow, the cavity 
being filled with a fatty marrow. The neural canal (can.nl.) — or 
foramen vertebrale — is somewhat pentagonal, the zygapophyses — or 
processi articulares — are well formed and wing-like, especially those 
at the posterior end, which are very broad and form a roof over the 
spinal ganglia. The obliquity of the sliding surfaces is very slight. 
The neural spine — ox processus spinosus — forms a well-defined ridge 
anteriorly but fades away posteriorly. The mesial portion of the 
neural arch between the posterior zygapophyses is thickened, and 
bears on its posterior surface a pair of small but well-marked depres- 
sions, one on each side of the middle line. From these concavities 
the MM. inter spinales arise (see p. 93). The anterior edge of the 
neural arch is V-shaped, so that a triangular space exists on the dorsal 
side between one vertebra and the next which is closed by membrane. 
The transverse processes (pr.tr.) which are stout and very deep dorso- 
ventrally protrude in a postero-lateral direction and bear ribs (rb.). 

4. The Ribs. 

The question of the articulation of the Urodele rib with the trans- 
verse process, and of the homology of the ribs with those of Amniota, 
has long been a somewhat vexed problem, but it has recently been 
cleared up by Gray (1931), who worked on the development of the 
ribs in Triton. An examination of the adult condition only would 
lead one to suppose that both heads of the rib articulate with the bifid 
transverse process, and indeed this was the view generally held until 
the appearance of Gray's paper. It was supposed that the attachment 
of the rib to the chorda, i.e. the capitulum of the rib, was represented 
by a transitory vestige only, but Gray showed that the reverse is 
actually the case, and that the chondrification of the rib rudiment is 
a centripetal process, the capitular process growing inwards towards 



THE SKELETON 19 

the centrum after the remainder of the rib has been formed. This 
capitular process passes ventral to the vertebral artery, thus enclosing 
it in a bony canal. Hence it is apparent that the Urodele rib is 
strictly homologous with the Amniote structure both with regard to 
its insertions on the vertebra and its relation to the artery, but since 
the capitulum fuses with the vertebra in the adult a secondary articula- 
tion is formed with the transverse process, thus masking the true 
relations. 

A detailed review of the earlier literature would be too lengthy to 
be included here, but the works of Goppert (1895) ^^^^ Mayerhofer 
(1909) stand out prominently, and the latter includes a good his- 
torical review up to that date. 

The ribs are better developed in the anterior region of the trunk, 
particularly those of the second and third vertebrae (Fig. 12), while 
towards the pelvic region they become reduced to mere acuminate 
appendages of the transverse processes. The first four or five ribs 
are bent more or less sharply downwards at about their middle, and 
from the point of bending there projects a sharp spine pointing in 
a postero-dorsal direction. The ribs of the second vertebra, and to 
some extent those of the third also, bear at their lateral extremities 
large spatulate expansions composed mainly of cartilage. From these 
expansions arise the MM. thoraci-scapulares suspending the scapula 
(see p. 74). 

The spinal nerves do not emerge from the neural canal through 
the intervertebral foramina as is the case in most animals, but through 
a special foramen situated immediately posterior to the transverse 
process. The intervertebral foramen is nevertheless quite large, but 
is permanently closed by a tough fibrous membrane (mem.iv.). 
Murray (1897) has investigated Spelerpes and Plethodon and has 
come to the conclusion that the phenomenon, which is fairly general 
among Urodeles, does not lend support to the theory that Urodele 
vertebrae are double structures as had been suggested. List (1883) 
describes one case of abnormal fusion of the seventh and eighth trunk 
vertebrae of a Salamander which was so complete as to simulate a 
single vertebra, but they disclosed their dual nature by the fact that 
they each bore a pair of ribs. 

5. The Sacrum. 

The sacral vertebra is normally the sixteenth, but Claus (1876), 
who has made a special investigation of this region of the vertebral 
column, finds that either the fifteenth or the seventeenth vertebra 
may exceptionally articulate with the pelvic girdle, and mentions also 



20 THE SALAMANDER 

three asymmetrical cases in which the ilium was attached to the 
rib of the sixteenth vertebra on one side and of the seventeenth 
on the other. The vertebra itself conforms to the general pattern 
of the trunk vertebrae already described, but it is slightly larger 
and its transverse processes and ribs are very stout and long. The 
extremities of the ribs are pointed and not flat as are those which 
support the shoulder. The ilia are closely bound to them by 
fibrous tissue. 

6. The Caudal Vertebrae. 

The caudal vertebrae (Figs. 1 8 and 1 9) exhibit a gradual reduction 
in the transverse processes which, however, never entirely disappear 
even at the tip of the tail. The zygapophyses are also reduced in 
accordance with the diminishing size of the vertebrae as a whole, but 
they likewise are never entirely absent. As a rule only the first caudal 
vertebra bears ribs and they are always very small. The most note- 
worthy departure from the typical form exhibited by this section of the 
vertebral column is the possession by all but the three anterior verte- 
brae of a ventral ov haemal 2irch (h.ar.), through which passes the caudal 
artery and vein. These arches are doubtless homologous with the 
chevron bones of Reptiles and some Mammals, but they are com- 
pletely fused with the centra of the vertebrae. The posterior end 
of each haemal arch is spread out fan-wise and is slightly lipped, 
while the anterior end narrows considerably and is splayed off very 
obliquely on the ventral side so that it does not extend quite to the 
anterior end of the centrum on the dorsal side. There is a low crest 
running along the mid-ventral line, while the sides are perforated just 
below the transverse processes by a foramen (fo.nt.) which permits 
the exit and entrance of the segmental blood-vessels of the tail. The 
whole series forms a bony canal as complete as the neural canal, and 
quite unlike that formed by the slight 'Y'-shaped chevrons of the 
Lacertilian type found in some Urodeles, e.g. Necturus. 

II. THE SKULL (Pis. I and II). 
I. The Skull as a Whole. 

One of the earliest investigators to describe the skull in any detail 
was Duges (i 834). His account is very condensed and is not much 
more than a catalogue of the several bones, but he was the first to 
show that the os triangulare is of branchial origin and has nothing to 
do with the shoulder girdle. Then follows Parker's well-known 
work (1882) in which the primordial skeleton is distinguished from 



THE SKELETON 21 

its investing bones, and some account is given of the development. 
There are unfortunately several inaccuracies in this work which 
will be referred to in detail in the subsequent description. Gaupp 
(1898) gives an excellent summary of the work done up to that date 
but adds little to it. The same author published a similar work in 
1905 dealing with the development of the skull. In recent years 
Stadtmuller (1924) has undertaken a complete and detailed rein- 
vestigation of the development of the skull of the Salamander, and 
his work has been accepted as a basis for the following investigation 
of the adult skull. 

As an introduction to this important part of the skeleton it may 
be well to quote the words of William Kitchen Parker when describ- 
ing the skull of Salamandra maculosa in 1882 (p. 172): 

'The skull of the adult of this species shows at once the likeness and the 
unlikeness of this type to that of an average Batrachian; yet in reality this 
skull differs as much from that of Rana temporaria or of Bufo vulgaris as it 
did in its early conditions and in the mode of its metamorphosis. But scattered 
up and down the great Batrachian "order" there are remarkably generalized 
types whose skulls, now in this and now in that agree with this of the highest 
kind of "Caducibranchiate" Urodele. I shall, therefore, carefully set down 
the characteristics of this skull, and then it can be used as a measure, being 
"perfectissimum in suo genere", of the lowness or height, in type, of the 
skulls of other Urodeles, and also as a test of what is normal or aberrant in the 
skulls of the Batrachia. This species and Rana temporaria, therefore, will be 
taken as convenient, and yet worthy, representative Salamandrian and Batra- 
chian types. They are culminating forms, being most perfectly specialized 
according to their kind.' 

These sentiments may well be extended beyond the skull to the 
animal as a whole, for, just as the Frog is an excellent basic type of 
Anuran specialization, so the Salamander, considered as a whole, is one 
of the best examples of Urodele organization, towards which other 
types may be regarded as leading, or from which they may be derived. 

The general outline of the skull as seen from above or below is 
roughly that of an equilateral triangle, of which the base-line is 
drawn through the quadrates. The occipital region extends only 
slightly behind this line. The jaws are widely spread laterally, thus 
giving the broad flattened appearance characteristic of Amphibia in 
general. The orbital rim is incomplete, since the maxilla and quad- 
rate are connected by a jugal tendon only; there being no osseous 
bridge. Thus in general shape the skull of the Salamander bears 
fairly close resemblance to that of the Frog, but in detailed structure 
they differ widely. 



22 THE SALAMANDER 

Any description of a Urodele skull falls naturally into two clearly 
defined sections, first, the primordial skeleton, comprising both 
cartilage and endochondrial ossifications, and second the investing 
bones. 

2. The Primordial Skeleton. 

The primary cranium of the Salamander is in many respects 
less developed than either the Frog or Dog-fish, and forms an excel- 
lent 'ideal' type. Both the roof and the floor of the brain-case 
are almost uncovered, the bridges between the right and left sides 
being entirely confined to the regions of the sense capsules. Thus 
the chondro-cranium consists roughly of a pair of olfactory capsules 
and a pair of otic capsules connected together by two longi- 
tudinal bars of partially ossified cartilage, while with the otic 
capsules are associated the occipital ring posteriorly, and the 
palato-quadrate laterally. As a protection to the eye a sclerotic 
ring is developed around that organ in the larva, but disappears 
during metamorphosis. Four regions of the primary skull are thus 
distinguishable. 

{a) The Ethmoidal Region consisting of the nasal capsules and 
their connectives. The accurate and detailed study of this region is 
practically impossible except by one or other of the reconstruction 
methods ; any attempt to raise the investing bones being almost sure to 
damage the thin cartilages lying beneath. Bremer (1924)1 adopted 
the wax-plate method of reconstruction and studied the development 
of the nasal capsules of Salamandra very thoroughly by this means. 
The following description of the ethmoidal region is based largely on 
his account. 

This portion of the primary skeleton forms about 30 per cent, of 
the total length of the skull. The walls of the nasal capsules are by 
no means complete, but are perforated by several large fenestrae as 
follows : 

(i) The fenestra basalis nasi (fen.b.nas.) in the floor is covered 
ventrally by the pre-vomer and, to a slight extent, by the pars pala- 
tina of the maxillary. The internal nares (nar.i.) or choanae enter the 
posterior portion of this fenestra. It is bounded posteriorly by the 
cartilago antorbitalis (c.a.or.), laterally by the cartilago ectochoanalis 
(c.ec.ch.), anteriorly by the cornu trabeculae, and mesially by the 

^ Bremer's account has not been published under his name but is incorporated in 
Stadtmiiller's (1924) paper on the skull. By the kindness of the Librarian of the Uni- 
versity of Gottingen the author has been enabled to consult the original typescript of 
Bremer's thesis. 



THE SKELETON 23 

trabecular to which is fused the planum internasale (c.pl.in.) — a flat 
cartilaginous plate connecting the nasal capsules ventrally. 

(ii) On the dorsal side of the nasal capsule is the fenestra dorsalis 
nasi (fen.d.nas.) lying opposite to the fen. basalis. The nasal, pre- 
frontal and frontal all assist in covering it. It is approximately quad- 
rangular in shape and is bounded mesially by the columella ethmoi- 
dalis (c.col.eth.) (Higgins), or the dorsal arch of the trabecula, while 
postero-laterally it is bounded by tht planum conchale (c. pi. con.), and 
antero-laterally by the cart, obliqua (c.obl.). 

(iii) Two fenestrae perforate the lateral wall of the nasal capsule. 
The anterior and larger is xh.^ fenestra narina (fen.nar.) which sur- 
rounds the external narial opening and its associated muscles. It is 
bordered dorsally by the cart, obliqua^ posteriorly by the cart, retro- 
narina (c.rt.nar.), and anteriorly by the cart, cupullaris (c.cup.). 

(iv) Posterior to the fenestra narina and separated from it by the 
cart, retro-narina lies tht fenestra infra-conchalis (fen.i.ch.) (or lateralis, 
Bruner). This opening lies at the extreme postero-lateral angle of 
the nasal capsule and is considerably smaller than the others already 
described. 

(v) The fenestra olfactoria (fen.ol.) penetrates the nasal capsule 
from the cranial cavity and serves for the passage of the olfactory 
nerves. 

The cartilago antorbitalis (c.a.or.) which separates the nasal capsule 
from the orbit is perforated by a pair of foramina, the for. orbito- 
nasale laterale (fo.o-nas.l.) and mediale (fo.o-nas.m.), through which 
pass the terminal rami of N. ophthalmicus profundus V. 

Thus the primary olfactory capsules of the adult Salamander con- 
sist of a series of cartilaginous bridges, while the intervening spaces 
are roofed over by the investing bones of the secondary skeleton. In 
addition to the cartilages already named in connexion with the 
fenestrae there are the following small projections. From the 
cart, ectochoanalis is a small backwardly projecting process which 
Seydel calls the Gaumenfortsatz (pr.S.). From the anterior end of 
each nasal capsule a small cartilaginous horn protrudes antero- 
mesially. This is the processus prenasalis inferior lateralis (pr.nas.) and 
represents the remains of the cornu trabecular Just below this pro- 
cess is a small opening for the exit of a nerve. It is thcforamen apicale, 
and, according to Bremer, it is not entirely constant. 

The olfactory capsules of either side are connected together ven- 
trally by a flat plate of cartilage which projects backwards under the 
anterior end of the brain, and extends forwards to about the centre 
of the ethmoidal region. It is the planum internasale (c.pl.in.). The 



24 THE SALAMANDER 

anterior portion of the space between the nasal capsules is thus open 
ventrally. It is called the cavum internasale (ca.in.) and houses the 
intermaxillary gland. The posterior wall of the cavum internasale is 
formed by the septum internasale^ which connects the nasal capsules 
posteriorly. It forms the cartilaginous partition between the cavum 
cranii and the cavum internasale and bounds the mesial borders of the 
jenestrae olfactoriae. The tectum internasale (c.t.in.) unites the capsules 
dorsally and thus forms a roof to the cavum internasale. It is pro- 
longed anteriorly in the middle line to a short beak, the rostrum 
■prenasale (Parker) (c.r.). 

{h) The next region of the primary skull is the Orbito-temporal. 
It consists of a pair of vertical lamellae joining the olfactory capsules 
to the otic capsules, and forming the sides of the brain-case, so that 
they separate the cavum cranii from the orbits. They are perforated 
by two foramina of which the larger is the for. opticum (fo.op.) and 
the other, immediately posterior to it, is the/or. oculomotorium (fo.oc). 
The optic foramen would perhaps be better termed the optic fenestra^ 
since its diameter is much greater than the optic nerve, and it is 
largely closed by membrane. In the adult the part of each lamella 
anterior to the vertical diameter of the for. oculomotorium becomes 
ossified, and the resulting bone is most usually termed the orhito- 
sphenoid (o.o-s.), although Broom holds that it is the homologue of 
the Mammalian pre-sphenoid and Noble refers to it as the spheneth- 
moid. The older and more widely used name is retained here. 

{c) The Otic Capsules. The membranous labyrinth of the audi- 
tory capsule is entirely enclosed by the primary skeleton which, in the 
adult, ossifies completely with the exception of the operculum. The 
capsules are relatively large and somewhat flattened dorso-ventrally, 
especially towards the posterior region. On the dorsal surface may 
be seen the convex ridges marking the position of the semicircular 
canals of the labyrinth. The lateral end of the ridge covering the 
horizontal canal is somewhat roughened where it meets the squa- 
mosal and it is often referred to as the crista muscularis (cr.m.) from 
the fact that certain spino-occipital muscles are attached thereto. 
The ventral and anterior surfaces of the capsule are much smoother. 
On the postero-lateral face, and directed somewhat ventrally, is an 
oval opening in the capsule — the. fenestra vestibuli. The fenestra is 
closed by a tough fibrous membrane, to the outer surface of which 
is attached the operculum (cop.). The operculum is a heavy plate of 
cartilage, strongly convex on its external surface and slightly concave 
internally, so that in transverse section it appears as a thick crescent. 
The M. opercularis attaches this element to the suprascapula. The 



THE SKELETON 25 

probable function and modus operandi of these structures are dis- 
cussed on p. 295. 

The lateral wall of the capsule is attached at three points to the 
suspensorium (palato-quadrate), and the details of this attachment 
will be considered in dealing with that part of the skull. The mesial 
wall of the otic capsule, separating the labyrinth from the cavum 
cranii, is perforated by five foramina, namely, the/or. perilymphaticum 
(io.Tp\.)^t\icfor.endolymphaticum(Jo.t\.\ and three/or. acustka(f O.3.U.). 

The for. perilymphaticum (fo.pl.), through which the duct of the 
same name passes into the cavum cranii from the labyrinth, is 
situated close to the base of the capsule, towards the posterior end. 

The /or. endolymphaticum (fo.el.) serves a similar purpose for the 
ductus endolymphaticus and lies a little anterior to the former — at 
about the middle of the capsule — near the dorsal ^2S\.. 

The three /or. acustica (fo.au.) in the adult lie almost in the floor 
of the capsule rather than in the mesial wall, owing to growth changes 
in that region. They serve for the entry of the auditory nerve and 
hence correspond to its three rami. They lie very close together and 
immediately anterior to \kie for. perilymphaticum. 

The cavity of the capsule is not quite simple but is subdivided 
by bridges which develop around the semicircular canals, the septa 
semicircularia (Stadtmiiller). The septum semicircularium anterior^ pass- 
ing from the antero-mesial to the dorsal wall, is very well developed, 
while the sept, semicirc. later ale is considerably weaker and is not always 
complete. The sept, semicirc. posterior is usually only faintly indicated 
and never develops completely. 

The otic capsules are completely fused to the cranium and ample 
traces remain of the parachordal plate. In the occipital region the 
capsules are joined together by a broad hypochordal commissure (c.hyp. 
com.), and, at their anterior ends, by a narrow bar of cartilage, the 
crista retrosellaris (c.cr.rs.) (Stadtmiiller). Between the two bridges 
the parachordals are represented only by slight ridges along the 
ventro-mesial borders of the capsules. The auditory capsules are also 
joined together dorsally by a narrow commissure at the posterior 
end, the tectum synoticum (c.t.syn.). There are thus two fenestrae in 
the floor of the primary cranium, an anterior and a posterior. The 
fenestra basicranialis anterior (fen. be. a.) is the larger and is bounded 
anteriorly by the planum internasale, laterally by the orbito-sphenoids 
(representing the trabeculae), and posteriorly by the crista retrosel- 
laris. The fenestra basicranialis posterior (fen. be. p.) lies between the 
crista retrosellaris anteriorly and the hypochordal commissure pos- 
teriorly, and is bounded laterally by the remains of the parachordal 



26 THE SALAMANDER 

plate. Both fenestrae are of course covered in the adult skull by the 
parasphenoid bone. Practically the whole of the ear capsule except 
the operculum ossifies in the mature adult, although the rim of 
the fenestra vestibuli and the inner surfaces bounding the cavum 
vestibuli tend to retain their cartilaginous nature. The combined 
glossopharyngeus-vagus nerve passes through X^o, foramen post-otkuni 
(fo.p-ot.) between the ear capsule and the occipital segment. 

The occipital ring remains almost entirely cartilaginous, and it is 
only the deeper parts of the condyles which ossify and fuse with the ear 
capsules. The composite ossification thus formed is best called the 
occipito-petrosal (Stadtmialler). 

{d) The Palato-quadrate (c.p-q.). The suspensorium is attached 
to the lateral aspect of the ear capsules by three processes, tripod- 
like, so that a cavity with three exits is enclosed between the palato- 
quadrate and the capsule. This cavity is the antrum petrosum laterale 
(Druner), and through it pass the Art. and Vena petrosa lateralis and 
the facial nerve. 

Of the three attaching processes the processus ascendens (pr.asc.) 
is the smallest. It is continuous with the lateral skull wall and 
separates the N. ophthalmicus profundus V from other two trige- 
minus branches. It also separates the artery from the vein. Normally 
the process remains unossified, but in old animals a certain amount 
of perichondrial ossification may occur. The processus otkus (pr.ot.) 
(proc. lateralis dorsalis, Druner) is the dorsal process and lies beneath 
the squamosal. It is rather larger than the proc. ascendens, but, when 
compared with the larval condition, it shows a tendency to lose its 
connexion with the ear capsule through the ingrowth of connective 
tissue (Stadtmuller, 1924). It remains cartilaginous. "XYit, processus 
basalis (pr.ba.) (proc. lateralis ventralis, Druner) is by far the largest 
of the three, and, as its name suggests, lies ventrally directly opposite 
the proc. oticus. According to Stadtmuller it first fuses completely 
with the larval skull and then separates from it again by the ingrowth 
of connective tissue, but this secondary segmentation occurs a little 
lateral to the point of the original fusion, so that a portion of the 
primitive proc. basalis is left fused to the capsule and forms the base 
of the canal through which the facial nerve emerges — the facialis 
canal. The now secondary proc. basalis — which is the only part 
recognizable in the adult — is thus separated from the primary portion 
by connective tissue, and the most convenient way to dissect the 
antrum petrosum laterale, in order to examine its contents, is to 
disarticulate this junction from the ventral side, when the whole sus- 
pensorium may be turned back, and, after breaking through the 



THE SKELETON 27 

other two processes, removed entire. The quadrate (o.qu.) forms the 
direct lateral continuation of the basal process and points almost at 
right angles to the skull axis in an obliquely downward direction. 
It articulates directly with the lower jaw and always ossifies to a 
greater or lesser extent, being in fact the only portion of the suspen- 
sorium to do so normally. Th.Qprocessus pterygoideus^ or 'cartilaginous 
pterygoid' (c.pt.), arises from an independent chondrification in the 
2,^ mm. larva (Stadtmuller) but is continuous with the palato-quad- 
rate in the adult, from which it protrudes antero-laterally towards the 
posterior end of the maxilla, pointing somewhat obliquely down- 
wards. It is only a narrow rod of cartilage and lies in a groove in the 
osseus pterygoid (which belongs to the investing bones of the skull) 
and usually protrudes a short distance anterior to it. 

The suspensorium is attached to the maxilla by two ligaments, 
the jugal ligaments^ namely lig. quadrato-maxillaris externum and the 
lig. quadrato-maxillaris internum^ which pass laterally and mesially of 
the M. levator mandibulae respectively. 

3. The Investing Bones. 

In contrast with the slow and tardy ossification of the replacing 
bones the investing bones arise very early, and are already well 
formed at birth, so that it is advisable to treat any larva with either 
a decalcifying fluid or an acid fixative before attempting to section 
it. Nevertheless they always remain thin and leaf-like. The follow- 
ing bones are present: on the dorsal side, pre-maxilla, maxilla^ nasal^ 
frontal^ pre-frontal, parietal^ squamosal — all are paired. On the ventral 
side there is an unpaired median parasphenoid and paired pre-vomers 
and pterygoids^ while both the pre-maxillae and maxillae have palatine 
processes. According to Stadtmuller a quadrato-jugal occurs, appear- 
ing first in a larva of 27 mm., but it always remains small and, in 
the adult, fuses homocontinuously with the palato-quadrate so as to 
be indistinguishable. 

The pre-maxillae (o.p-m.) are paired in the Salamander and form 
the skeleton of the snout. They are sometimes called intermaxillaries 
(Funk, Bolkay). Three regions are distinguishable in each bone 
which, according to Stadtmuller, arise as three distinct ossifications. 
These are : {a) the pars prenasalis (or dorsalis) which ascends over the 
dorsal side of the nasal capsule and ends in a sharp slender process 
which to some extent overlaps the nasal and frontal, (b) The pars 
den talis forms the anterior border of the upper jaw and bears teeth — 
it is in fact formed by the fusion of tooth sockets. It is joined to 
the pars prenasalis dorsally and to the pars palatina postero-mesially. 




2 8 THE SALAMANDER 

(c) The pars palatina is a narrow but strong bar of bone which helps 
to form the osseous palate. According to Stadtmiiller it arises partly 
by the fusion of tooth sockets and partly as a separate ossification direct 
from the membrane. 

The maxillae (o.m.) complete the arch of the upper jaw and sur- 
round the latero-ventral edge of the cartilaginous nasal capsule like 
a gutter. Like the pre-maxillae each bone has three parts : (a) The 
pars facialis borders the postero-lateral margin of the external nasal 
aperture and covers the side of the face as far as the antero-lateral 
angle of the orbit. Dorsally it extends on to the edge of the nasal, 
and ventrally it fuses with (b) the pars dentalis which, like the corre- 
sponding part of the pre-maxilla, arises by the fusion of tooth-sockets. 
The teeth only extend backwards to about the level of the anterior 
margin of the optic fenestra. (<:) The pars palatina corresponds en- 
tirely with the similarly named portion of the pre-maxilla, against 
which it abuts anteriorly. On the mesial side it meets the pre-vomer. 
The maxilla extends posteriorly to about the level of the hinder 
border of the optic fenestra. Its posterior extremity lies lateral, and 
slightly caudal, to the anterior extremity of the pterygoid, and is 
joined to the lateral extremity of the suspensorium by a pair of tough 
ligaments — tho. jugal ligaments — but there is no bony connexion as in 
the Frog. 

The nasals (o.nas.) are paired bones of a rather irregular shape 
serving to roof over the mesial portion of the nasal capsules. Their 
antero-lateral edges curve round the mesial border of the external 
nasal aperture.^ Mesially they meet, and overlie, the antero-lateral 
edges of the frontals and the pre-nasal processes of the pre-maxillae, 
while posteriorly they adjoin the pre-frontals, and laterally meet the 
dorsal edges of the facial portions of the maxillae, and are slightly 
covered by them. The nasals are distinctly lateral in position and 
are widely separated from one another in the middle line. They are 
perforated by a variable number of foramina for the passage of the 
ultimate twigs of the mesial branch of the N. ophthalmicus profundus 
V. According to Stadtmuller their development is relatively late. 

ThQ pre-frontals (o.p-fr.) {fronto-lacrymaux^ Duges; ectethmoid V^iV- 
ker) are also paired and occur on each side of the skull immediately 
anterior to the orbit. They are more or less quadrangular in shape, 
and are comparatively small. Their anterior borders are covered by 
the nasals, mesially they overlie the edges of the frontals, while 
laterally they are in turn pushed under the dorsal edges of the facial 

' It should be noted that the aperture formed by the edges of the pre-maxillary, 
maxillary, and nasal bones is larger than the actual cutaneous opening. 



THE SKELETON 29 

portions of the maxillae. The posterior edge of the pre-frontal is free 
and forms a bony rim to the antero-dorsal border of the orbit. The 
ductus naso-lacrimalis (d.nas-lc.) enters a foramen at the postero- 
lateral angle of the bone, and passes antero-lateralwards under it and 
the pars facialis of the maxilla to enter the nasal capsule near the 
external narial opening. 

W. K. Parker (1882) describes and figures a septo-maxillary at 
the antero-lateral angle of the apertura nasalis externa, between the 
maxilla and the pre-maxilla, but he is the only author to do so. 
Stadtmuller (1924) declares himself totally unable to find it in any 
of his sections, and Lapage (1928) also failed to detect any such bone. 
Bruner (1902) describes the bone in certain other Urodeles but says 
it is wanting in Salamandra. The present investigation confirms the 
result of the later workers. It therefore seems that Parker must have 
been mistaken or that he encountered a very abnormal specimen. In 
view of Lapage's work the latter alternative is unlikely. 

The brain-case proper is roofed over by two pairs of investing 
bones, namely the frontals and parietals; they remain separate 
throughout life and do not fuse as in the Frog. 

ThQ frontals (o.fr.) are large and extend far forwards under the 
prenasal processes of the pre-maxillae, while their antero-lateral bor- 
ders also underlie the nasals for some distance and meet the mesial 
edges of the pre-frontals. The lateral edge of each frontal turns down 
ventrallyand meets the orbito-sphenoid, while posteriorly the frontals 
overlap the anterior edges of the parietals. The sagittal suture is very 
irregular and is by no means a straight line. There is no post-orbital 
process as in some Urodeles. 

^ht parietals (o.par.) lie immediately posterior to the frontals and 
have approximately the same superficial area. They complete the 
roof of the cranial cavity, while their posterior ends spread out later- 
ally over the mesial portions of the ear capsules. N. trochlearis may 
penetrate the parietal, but, in older animals at least, this nerve usually 
passes through a foramen in the orbito-sphenoid. The parietals do 
not extend quite to the dorsal rim of the foramen magnum but leave 
a portion of the tectum synotkum exposed. The dorsal surface of the 
frontals and parietals is very flat, or even slightly concave as seen in 
transverse section, and while both are relatively thick bones they are 
very transparent, so that it is quite easy to distinguish the main 
features of the brain through them, even in the adult. 

The squamosals (o.sq.) (jympanicum^ Wiedersheim ; paraquadrate^ 
Gaupp) are paired splint-like bones lying on the dorsal side 
of the quadrates, to which they become very closely fixed. Each 



30 THE SALAMANDER 

squamosal extends from the quadrato-mandibular joint along the 
processus oticus of the palato-quadrate and becomes more or less fused 
with the periotic bone above the horizontal canal. While there is a 
distinct tendency for the bone to be 'T'-shaped it is not so definite as 
in the Frog. 

A quadrato-jugal has been described by Stadtmiiller. According 
to him it appears first in a larva of about 27 mm. and always remains 
small. In the adult it fuses homocontinuously with the ossified por- 
tion of the quadrate, from which it becomes indistinguishable. 

The investing bones of the ventral surface are the parasphenoid, 
the pre-vomers, and the pterygoids. 

'Th.Q parasphenoid {sphenoid^ Duges; Kei/hin, ¥loffm3.nn; vomer ^m 
part, Broom; vomer -^ pterygoids^ Kesteven). This bone is the largest 
in the skull and is characteristic of Amphibia. It forms at one and 
the same time a bony floor to the cavum cranii and a roof to the 
mouth, and is the only median bone in the skull. Posteriorly it 
extends for some distance under the ear capsules on either side, but 
hardly so far, comparatively, as in the Frog. In the orbital region 
the edges turn upwards to meet the orbito-sphenoid, so that the 
bone becomes gutter-shaped in cross-section. The internal carotid 
artery passes into the cranial cavity through a canal — the carotid canal 
(can. car.) — between the parasphenoid and the otic capsule. 

Until quite recent years the parasphenoid has been looked upon 
as a typically amphibian bone, which was also present in Reptiles, 
but in a very reduced state. Recently, however, Broom has sought 
to identify the anterior part of the amphibian, and the whole of the 
reptilian, parasphenoid with the mammalian vomer., while Kesteven 
(19 1 6 and 1926) claims that the posterior part of the amphibian 
parasphenoid is represented in Reptiles and Mammals by the ptery- 
goids. Both these authors base their views mainly upon palaeonto- 
logical evidence. 

The pre-vomers (o.p-v.) (Broom) are perhaps better known as the 
vomero-palatines, but, if one accepts the above-mentioned conclusions 
with regard to the homology of the mammalian vomer, the bones 
now to be described can be none other than the pre-vomers. Whether 
any palatine element is represented will be considered later. The 
paired pre-vomers form a bony floor to the nasal capsules. Each bone 
articulates with the pre-maxilla and maxilla laterally, while mesially 
its anterior border bounds the large internasal space, behind which 
it is approximated to its fellow on the other side for a short distance. 
The edge of the bone then diverges from the middle line and follows 
the curve of the dentigerous portion. The bone also surrounds the 



THE SKELETON 31 

anterior rim of the internal narial opening and a part of the orbit. 
It carries a very characteristic S-shaped dentigerous ridge which 
extends far backwards in the mouth and turns lateralwards under the 
ear capsule. This posterior extension is quite free from the overlying 
part of the skull (parasphenoid), and is separated from it by soft 
tissues, including the M. retractor bulbi, which passes dorsal to the 
posterior end. The pattern of this dentition is of taxonomic im- 
portance (cf. p. i). 

As already suggested, the older workers, e.g. Parker and 
Wiedersheim, regarded it as representing both the vomer and the 
palatine elements fused together, and named it accordingly vomero- 
palatine. In 1 9 10 Wintrebert, as the result of an investigation of the 
metamorphosing skull, came to the conclusion that the palatine en- 
tirely aborts during the change from larval to adult life, and that the 
bone under consideration should therefore be regarded as the vomer. 
In 1924 Stadtmiiller, whose careful and detailed work on the de- 
velopment of the skull has already been mentioned, declares that in 
all his examples the vomer appears to fuse with the palatine, but 
that it is extremely difficult to decide whether this is really the case, 
or whether the latter is entirely resorbed, as Wintrebert claims. He 
criticizes Wintrebert's work on account of the absence of any 
figures, details of methods, and size of the larvae investigated, 
and says that he prefers to agree with the older investigators. 
He nevertheless confirms Wintrebert's statement that the vomerine 
part has a double origin, while he does not discuss the possibility of 
an homology with the pre-vomer. He had not seen Wintrebert's 
chief paper (1922) against which his criticism certainly could not 
be levelled. 

The whole question therefore remains sub judice and its solution 
lies outside the scope of the present work, but in view of the 
evidence of palaeontology the newer terminology has been adopted 
here. 

The pterygoids (o.pt.) are a pair of more or less ploughshare-shaped 
bones investing the pterygoid cartilages of the primary cranium, 
the cartilages lying in a groove on the dorsal surface of each investing 
bone. The posterior end of the bone lies under the quadrate, and 
extends laterally almost to the jaw articulation and mesially over the 
processus basalis. The anterior extremity points antero-laterally to- 
wards the posterior extremity of the maxilla but does not quite meet 
it, and hence the bone forms the posterior margin of the orbit. 

Kesteven identifies this bone with the reptilian ectopterygoid or os 
transversum rather than with the reptilian and mammalian pterygoid, 



32 THE SALAMANDER 

which, as already mentioned, he considers homologous with the pos- 
terior portion of the amphibian parasphenoid.^ 

Summary. 

Reviewing then the skull as a whole, particularly in comparison with 
that of the Frog, the following important differences may be noticed. 

(i) The skull generally is more compactly built than in the Frog. 

(ii) The nasal capsules are more completely roofed over, since the 
nasals are pushed forwards by the pre-frontals, which are not present 
in the Frog. 

(iii) 'X'\\Q pre-vomers have a much greater expansion and thus form 
a complete bony floor to the nasal capsules, while their dentigerous 
portions extend far back into the mouth. 

(iv) Thcfrontals 2ind parietals remain distinct elements and do not 
fuse as in the Frog, neither do they extend so far ventrally round the 
sides of the brain case. 

(v) The brain-case is relatively wider in Salamandra than in Rana, 
when compared with the width of the skull as a whole. 

(vi) The feature just noticed is due largely to the much greater 
development of the suspensorium in the Frog. In Salamandra the 
squamosals are mere splints on the dorsal surfaces of the palato- 
quadrates, while the quadrato-jugal sends no process to meet the 
maxilla. The pterygoid likewise does not join the maxilla so that 
the arch of the upper jaw is incomplete. 

(vii) On the ventral surface the parasphenoid is relatively broader 
in Salamandra, so that the sides of the brain-case — orbito-sphenoids 
— are more nearly vertical. 

(viii) The anterior rim of the orhit on the ventral side is formed by 
the edge of the pre-vomer in Salamandra, and by the palatines in Rana, 
the latter elements being probably absent in the adult Salamander. 

(ix) In the ear capsule the columella auris is vestigial in Sala- 
mandra, its place being taken by the cartilaginous operculum^ which 
has no direct connexion with the skin, while in Rana the columella 
is attached to the tympanic membrane and there is no operculum. 

III. THE VISCERAL ARCHES. 

I. The Mandibular (First Visceral) Arch. (PI. II, fig. 7, and PL 

III, figs. 8 and 9.) 

The shape of the lower jaw as a whole is a simple bony arch, 
almost semicircular in plan. At the median symphysis there is a 

J See also Watson's criticism of this hypothesis, Journ. Anat. London, vol. liii, 
pp. 239-40. 



THE SKELETON 33 

prominent posterior mental -process. The anterior two-thirds of each 
ramus bears a single row of functional teeth, mesial to which a row 
of reserve teeth can be demonstrated in transverse sections. (For 
details of the teeth, see p. 262.) 

The structure of the mandible, in both the larva and adult, has 
been thoroughly investigated by Gaupp (191 1) and Stadtmuller 
(1924). Both accounts are in agreement and correct that of Parker 
(1882) by showing that the j-/>/^;?zW (Operculare of the German 
authors) is present in the larva only, and is resorbed entirely in the 
adult. There are therefore two investing bones only in the lower jaw, 
namely, a dentary and a pre-artkular (Goniale), while two replacing 
ossifications also occur in the persisting Meckel's cartilage, viz. 
the articular at the proximal end and the mento-Meckelian at the 
symphysis. 

Meckel's cartilage (c.M.) persists as the primary skeleton of the 
lower jaw throughout life, and is unossified in the major part. It 
remains exposed at the proximal end between the two investing bones 
— dentary and pre-articular, while the anterior end tapers to a point 
and terminates a short distance from the middle line, its mesial ex- 
tremity having become ossified as a mento-Meckelian (Mento-mandi- 
bulare, Stadtmuller), so that the median symphysis is solid bone. 
This ossification occurs quite late, well after metamorphosis, and 
becomes fused homocontinuously with the dentary, which is probably 
the reason that lead W. K. Parker to say (p. 177) 'the Salamander 
has no mento-Meckelian bone'. The extreme proximal end of Mec- 
kel's cartilage is the only element of the lower jaw to come into 
contact with the suspensorium. Of this the actual surface remains 
cartilaginous, but the deeper portions become transformed into a 
small bony knob, the articular (o.art.). According to Stadtmuller 
this ossification also occurs very late and is first discernible in a 
metamorphosed individual of about 43 mm. long. 

Investing Bones. The dentary (o.den.) forms the principal ele- 
ment of the adult mandible. It covers almost the whole of Meckel's 
cartilage laterally and most of its dorsal and ventral surfaces, and 
bears teeth (see p. 263). In transverse section it is more or less C- 
shaped, with the opening of the *C' towards the mesial side. Anteriorly 
it forms an almost complete circle around Meckel's cartilage, but it 
opens out posteriorly and tapers at the same time, so that it terminates 
a little anterior to the articulation as a thin lath on the lateral aspect 
of Meckel's cartilage. Its fusion with the mento-Meckelian has 
already been mentioned. W. K. Parker describes its dorso-mesial 
portion as a separate element, the splenial^ but according to 

4038 n 



34 THE SALAMANDER 

Stadtmiiller, the dentary, during its development, grows round and 
envelops the larval splenial, which is eventually resorbed, although 
he thinks that a few of its anterior teeth may occasionally remain 
fused with the dentary. Thus the bone as above described is 
equal to the dentary and splenial of Parker's account. Enclosed 
between the dentary and Meckel's cartilage are branches of nerves 
V and VII, and the blood-vessels for the nutrition of the teeth, lower 
lip, &c. 

The pre-artku/ar (o.p-art.) is a wedge-shaped splint of bone which 
is pushed in between the free edges of the dentary on the mesial side of 
the jaw. It expands dorsally into a fairly pronounced coronoid pro- 
cess which is inflected mesially, and thus turns away from the dentary 
so as to expose Meckel's cartilage and the articular dorsally. It does not 
become fused with the articular as it does in some forms (e.g. Triton, 
Gaupp), and is perforated near its posterior end by a foramen for the 
R. alveolaris of N. VII and the alveolar artery (fo.i.den.). This 
element represents the arlicu/aroflp2Lrker, and is called the Goniale by 
Gaupp, Stadtmiiller, and most German authors. 

The Hyobranchial Skeleton. (PI. II, fig. 6 and PI. VI, fig. 38.) 
In the larval Salamander the hyoid and portions of four branchial 
arches are all present. In the adult, however, only portions of the 
hyoid and the first two branchial arches normally persist, the re- 
mainder disappearing during metamorphosis. The whole skeleton 
is cartilaginous with the exception of the small os triangulare (q.v.). 

2. The Hyoid (Second Visceral) Arch. 

This arch consists, in the adult, of two elements on either side, 
namely the hypo- and cerato-hyah. In the larva these two articulate 
with one another, but in the adult they become separated. The hypo- 
hyals (c.h.hy.) retain their connexion with the median longitudinal 
portion of the hyo-branchial apparatus, namely the copula (c.cop.) 
(or basibranchial^ W^iedersheim), from which they protrude as a pair of 
short horns pointing in an antero-dorso-lateral direction, being more 
or less embedded in the substance of the tongue. They are often 
referred to as the anterior radials of the copula (see below). The 
cerato-hyals (c.c.hy.) form the anterior cornua of the hyoid. Their 
anterior ends are greatly expanded into broad spatulate sheets of 
cartilage which underlie the hypo-hyals ventrally, and also overlap 
one another in the middle line. Their posterior ends are much 
narrower and more rounded in section, and curve dorsalwards to 
about the level of the jaw suspension, where they are attached to the 
quadrate by a strong elastic ligament, the ligamentum hyo-quadratum. 



THE SKELETON 35 

Their posterior ends give origin to the M. sub-hyoideus, while the 
anterior ends serve for the insertion of M. subarcualis rectus i. 

3. The First Branchial (Third Visceral) Arch. 

This consists of a narrower arcuate bar articulating with the copula 
on either side and comprising hypo- and cerato-branchial elements 
fused together homocontinuously (i.e. without interruption). Each 
bar is a flat oval in cross-section and its posterior end curves dorsal- 
wards approximately parallel with the cerato-hyal, thus forming the 
posterior cornu of the hyoid. From the posterior end of this cartilage 
the M. subarcualis rectus i takes its origin. 

4. The Second Branchial (Fourth Visceral) Arch. 

The second branchial {^fourth visceral) arch of the adult is repre- 
sented only by the paired hypo-branchial elements (c.h.br. II.). These 
are slender curved bars articulating mesially with the posterior end 
of the copula, and fusing laterally with the previous arch at a point 
about two-thirds along its length. The space between the two arches 
is covered by a tough membrane, the memhrana intercartilaginea 
(Driiner) (me. i.e.), except over the mesial third, where the membrane 
is incomplete to allow for the passage of the M. rectus cervicis 
profundus. 

5. The Copula. 

The copula (c.cop.) is the median cartilage with which those of the 
visceral arches articulate. It probably represents the fused basi- 
branchials, but it never shows signs of segmentation even in the larva. 
Its anterior end bears two pairs of small horn-like processes — the 
radials. As already indicated the anterior pair (c.h.hy.) of the latter 
are the hypo-hyals, but the posterior pair (c.rad.) have been shown by 
Driiner (1901) to arise secondarily during metamorphosis, and are 
therefore not strictly a part of the visceral skeleton. They, like the 
hypo-hyals, serve to attach the copula to the tongue. 

6. The Os Triangulare. 

The OS triangulare (o.t.) is a small, broadly triangular bar of bone 
lying across the ventral side of the pharynx in an inscriptio tendinea 
at the junction of the M. genio-hyoideus and M. rectus cervicis 
superficialis. It is the only portion of the hyo-branchial skeleton to 
ossify. It represents the expanded end of the uro-branchial, and, in 
the larva, it is connected with the copula by a longitudinal rod which 
disappears during metamorphosis, although very occasionally small 



36 THE SALAMANDER 

vestiges may persist as little cartilaginous knobs between the right 
and left portions of the M. genio-hyoideus. 

The chief abnormality met with in the visceral skeleton is the 
occurrence of a rudiment of a hypo-branchial^ element in the third 
branchial (fifth visceral) arch. Four instances of such a condition 
occurring in larvae have been reported, namely, one case of a rudi- 
ment occurring on the left side (Kallius, 1901), two instances of a 
rudiment of the right side (Driiner, 1901), while Helena Tarapini 
(1909) finds in a 30 mm. larva of Salamandra atra a pair of small 
cartilages which she regards as rudimentary third hypo-branchials. 
During the present investigation one case has been found in which 
cartilages apparently representing these elements occurred in the 
adult. This has been fully described elsewhere (Francis, 1931). 
Druner also reports cases where very small vestiges of the cerato- 
branchials III and IV have been found in the adult, and, as mentioned 
above, small traces of the uro-branchial shaft have occasionally been 
noticed. 

The visceral skeleton of Salamandra, both larval and adult, has 
received a considerable amount of attention from time to time, and 
its several parts have been given names indicative of widely varying 
interpretations of their homology (a list of the more important 
synonyms is appended below). Rusconi (1854) was one of the first 
to undertake a detailed investigation of this region in the larva as 
well as in the adult, and his classic account of the metamorphosis of 
both skeleton and muscles remained without any substantial addi- 
tion until 1 90 1, when Druner, applying more modern methods, 
reinvestigated the whole problem. Rusconi was of the opinion that 
both pairs of radials were of secondary origin, while Walter (1887) 
considered that they represented two pairs of hypo-hyals. Druner, 
however, showed that the anterior pair only belong to the hyoid arch 
while the posterior pair are new structures. Wilder, Druner, and 
others also include the laryngeal cartilages in the visceral skeleton, 
but Edgeworth (1920) has shown this to be erroneous (see also under 
'Larynx', p. 272). 

Synonyms of the Visceral Skeleton. 
Cerato-hyal = Stylo-hyal, Duges (1834). 
Both branchial arches together = Corne thyroidienne, Duges. 
Hypo-branchials = Kerato-branchials, Wiedersheim. 
Cerato-branchials = Epi-branchials, Wiedersheim. 

^ It should be noted that only the cerato-branchial element is normally present in_ 
the larval Salamander. 



THE SKELETON 37 

Hypo-branchial I + Cerato-branchial I = Epi-branchial I, Parker. 

Hypo-branchial II = Cerato-branchial II, Parker. 

Copula = Basi-hyal, Duges; Basi-branchial I, Wiedersheim, Parker. 

Radials = Corne stylo'idienne, Duges; Kleine Horner, Wiedersheim. 

Anterior pair only = Hypo-hyal, Parker. 

Posterior „ = Cerato-branchial I, Parker. 

Os Triangulare = Os thyroideum, v. Siebold, Duges, Walter, Wiedersheim. 

= Sternum, Funk. 

= Basi-branchial II, Parker. 

= Os triquetum, Driiner. 

THE APPENDICULAR SKELETON 

I. THE PECTORAL GIRDLE AND FORE-LIMB. (PI. IV and PI. VII, 

figs. 39-42.) 

I. The Pectoral Girdle. (Figs. 20 and 21.) 

Each half of the shoulder-girdle of the Salamander is best re- 
garded as a single skeletal element which is incompletely divided into 
three regions, rather than as an aggregate of three fused elements. 
The major portion remains cartilaginous, the only bone present being 
in the neighbourhood of the glenoid cavity. 

The glenoid cavity (gl.) is ovoid in shape, and is guarded by pro- 
minent faceted lips on the pro-coracoid and coracoid borders. Its 
periphery is largely ossified except at the post-axial border. 

(a) The Scapular region (Engler, 1929). 

Scapula et adscapulum ...... Duges (1834) 

pars Scapularis ...... Stannius (1854-6) 

Scapula et suprascapulare Gegenbaur (1865); Wiedersheim (1892) 
Scapula et suprascapulare . Parker (1868); Hoffmann (1873-8) 

Scapulum et episcapulum .... Sabatier (1880) 

Scapulum ....... Perrin (1899, &c.) 

The scapular region forms the dorso-lateral portion of the girdle and 
comprises first a proximal osseous portion which is the scapula proper 
(o.sc), and secondly a distal cartilaginous plate forming the supra- 
scapula (c.s.sc). The whole structure is fan-shaped and is curved in 
a dorso-ventral direction, so that the convexity is outwards. The 
scapula is elliptical in section and expands only slightly towards its 
distal border, where it merges into the supra-scapula. This latter 
portion expands into a broad fan, comparatively thick at its ventral 
border, but thinning out very considerably dorsally. It is attached to 
the axial skeleton by the MM. thoraci-scapularis and opercularis, as 
well as by connective tissue which extends over the M. dorsalis trunci 
to the neural spines of the second and third vertebrae. The scapula 



38 THE SALAMANDER 

ossifies earlier than the bony region of the ventral portions. ^ Its 
anterior border fuses to a considerable extent with the bony portion 
of the pro-coracoid, but its posterior edge only just meets the coracoid 
at the edge of the glenoid cavity. The angle between the anterior 
edge of the scapula and the lateral edge of the pro-coracoid is nearly 
a right angle. 

(J?) The Pro-coracoid region (c.pr.cor.). 

Clavicle ou fourchette ...... Duges (1834}. 

Pars acromialis (of scapula) .... Stannius (1854-6). 

Procoracoid . Gegenbaur (1865); Parker (1868); Furbringer (1873); 
Engler (1929). 

Precoracoide et epiprecoracoide .... Sabatier (1880). 

Clavicula ....... Wiedersheim (1892), 

Together with coracoid portion = 

Sternum ........ Funk (1827). 

CoracoTde ........ Perrin (1899). 

This region of the pectoral girdle forms the anterior part of the 
ventral plate. It projects almost directly anteriorly from the glenoid 
cavity, and extends nearly as far forwards as the level of the os tri- 
angulare on the floor of the pharynx. It is spatulate in shape and 
slightly dished, with the convexity ventralwards. It is ossified in the 
region of the glenoid cavity but remains cartilaginous anteriorly, and 
is only imperfectly demarcated from the coracoid portion by a deep 
notch — the incisura coracoidea (in. cor.) — which separates the cartila- 
ginous portions of the two regions. The osseous part of the pro- 
coracoid is entirely continuous with that of the coracoid except for 
the foramen supracoracoideum (fo.s.cor.), through which passes the 
N. supracoracoideus and the corresponding artery and vein. A line 
drawn from the glenoid cavity through this foramen to the centre 
of the incisura coracoidea may be taken to represent the boundary 
between the pro-coracoid and coracoid portions, which may thus be 
regarded as in an incipient state of separation. The free edges of the 
cartilage are exceedingly thin. The pro-coracoid forms a prominent 
lip over the anterior face of the glenoid cavity which presents a 
definite facet to the head of the humerus. The free edge of this lip is 
cartilaginous. 

{c) The Coracoid Region (c.cor.). 

Coracoidien ........ Duges (1834). 

Coracoid .... Gegenbaur (1865); Furbringer (1873); 

Wiedersheim (1892); Engler (1929). 
Coracoid and epicoracoid . . Parker (1868); Sabatier (1880). 

I Cf. Parker (1868); Perrin (1899); and Engler (1929). 



THE SKELETON 39 

Together with the pro-coracoid region = 

Sternum ........ Funk (1827). 

Coracoide ........ Perrin (1899). 

This is by far the largest portion of the pectoral girdle and extends 
mesially from the glenoid cavity across about two-thirds the width 
of the body, so that there is an overlapping of the right and left sides. 
It is apparently immaterial in Salamandra maculosa whether the 
right is ventral to the left or vice versa, since both arrangements are 
found to occur with about equal frequency, although in Triton and 
S. atra the former appears to be the more frequent condition. In 
maculosa the controlling factor is probably mechanical pressure 
during intra-uterine life.' The coracoid portion forms a more or less 
semicircular plate, deeply hollowed, with its convex surface ventral. 
Only a very small portion, in the region of the glenoid cavity, is 
ossified. The remainder is cartilaginous. Like the pro-coracoid its 
free edges are exceedingly thin, while the postero-mesial edges are 
attached by connective tissue to the sternal plate (q.v.). It also forms 
a faceted lip to the posterior border of the glenoid cavity which is 
entirely cartilaginous but is not so pronounced as that formed by the 
pro-coracoid. The overlapping portions of the coracoids lie im- 
mediately ventral to the heart, which they protect. 

Parker designates as 'epicoracoid' the antero-mesial portion which 
projects slightly from the general contour, while Sabatier applies the 
same term to the whole cartilaginous portion. Engler, however, 
shows quite clearly that a true epicoracoid is absent from Urodeles.^ 
It should be noticed in this connexion that the overlapping of the 
coracoids — known as arcifery — is not identical with the Anuran 
arcifery (e.g. Bufo), since in these animals it is the epi-coracoids which 
overlap and not the coracoids. 

{d) The Sternum (most authors) (Fig. 22). 

Cartilaginula quadrata intermedia .... Funk (1827). 

The sternum of the Salamander is a small median cartilaginous 
plate, lying behind the coracoids. Its shape is roughly that of an 
equilateral triangle with the apex directed forwards, and a short pro- 
jection from the middle of the base pointing posteriorly. The sides 
are grooved to receive the postero-mesial edges of the coracoid 
cartilages. These grooves are fairly deep and do not merge into one 
another anteriorly, but are divided by a septum, so that one groove 

1 Cf. Engler (1929). 

2 Except in Siren lacertina and Cryptobranchiis allegkaniensis. 



40 THE SALAMANDER 

passes dorsally and the other ventrally across the apex of the triangle. 
Which side is ventral and which dorsal depends on the relation of 
the coracoids to one another, since the grooves in the sternum occupy 
the corresponding relative positions. The sternum is attached to 
the coracoids by a membrane, memhrana sterno-coracoidea (Engler), 
which arises from the linea alha^ and is also closely attached to the 
pericardium dorsally, its dorsal surface being slightly concave so as 
to fit this structure. Although the exact shape is subject to consider- 
able fluctuation, the present writer has never seen a sternum with the 
long anterior process such as is figured by Perrin, and it would seem 
that his figure must have been prepared from a very unusual speci- 
men. The sternum serves for the attachment of the pectoralis muscle 
ventrally, while superficial rectus muscles of the abdomen and neck 
are attached to its edges, dorsal to the coracoids. A very good and 
detailed account of this structure and of its development is given by 
Engler. 

As may be judged from the above synonyms, the several parts of 
the pectoral girdle have been homologized with widely different 
structures. It is perhaps unnecessary to review the somewhat 
involved discussions here, since the identities of the elements as 
expressed above are now generally accepted. The most noteworthy 
papers are those of Gegenbaur (1865), and Parker (1868), while 
Engler (1929) has recently worked out the ontogenetic history in 
considerable detail, particularly with regard to the sternum. 

2. The Fore-limb (figs. 23—8 and 39—42). 

This comprises the usual elements found in a tetrapod fore-limb, 
namely, the humerus in the arm, the radius and ulna in the fore-arm, 
carpals and metacarpals in the wrist, ?in6. phalanges in the digits. 

The chief points of interest are: (i) the peculiar shape of the 
humerus, and particularly of its proximal end, (ii) the morphology 
of the carpus and the homology of the digits. Each of these features 
will be dealt with in the appropriate place. The epiphyses of all the 
long bones, and all articular surfaces, are cartilaginous. 

{a) The Arm (Figs. 23, 24, and 42). The long axis of the humerus 
is almost straight. The head is expanded dorso-ventrally in a peculiar 
manner and merges into the crista ventralis humeri (cr.v.h.) The 
cartilaginous epiphysis at this end when seen in side view is therefore 
shaped like a comma (fig. 42). The crista ventralis is short, occupy 
ing only about a third the total length of the bone, but it is of con- 
siderable expansion and rises steeply from the shaft. It fits into the 
notch, already referred to, formed in the rim of the glenoid cavity 



THE SKELETON 41 

by the lips of the coracoid and pro-coracoid, and serves for the 
attachment of MM. pectoralis and supra-coracoideus. At about the 
same level on the other side of the bone is the crista dorsalis humeri 
(cr.d.h.). This is a smaller hooked process and does not extend on 
to the head of the humerus. M. subscapularis is inserted on it. 
Immediately distal to the point from which these crests arise the 
humerus is approximately circular in cross-section, but it very soon 
expands in an axial plane, i.e. at right angles to the expansion of the 
head. On the pre-axial border of the distal end is the epicondylus 
lateralis humeri (co.l.h.), a fairly pronounced ridge, partly osseous 
and partly cartilaginous, which gives rise to the MM. extensor 
antebrachii et carpi radialis, and extensor digitorum communis and, 
in part, to the M. extensor antebrachii et carpi ulnaris. On the 
postaxial border is a similar ridge, the epicondylus medialis humeri 
(co.m.h.), from which arise the flexor muscles antagonistic to those 
above mentioned. The radial condyle (co.r.) (Eminentia capitata^ 
Gaupp) is large, and forms about two-thirds of a complete sphere, 
standing out prominently from the surrounding structures towards 
the ventral, or flexor, surface of the arm. The ulnar condyle (co.u.) 
is much smaller, and is approximately semi-elliptical in shape, while 
between the two condyles is a shallow trochlear groove (fs.tr.). Both 
condyles and groove are entirely cartilaginous. The fossa cubitalis 
centralis (fs.cu.v.)(Gaupp), into which the end of the radius fits when 
the arm is flexed, is very deep and well marked, but the olecranon 
fossa (fs.ol.) is shallow and indefinite. 

(b) The Fore-Arm (figs. 25-7 and 39-41). The radius and ulna 
are completely separate, and of approximately equal length and 
average size. The proximal end of the radius^ or pre-axial element 
(figs. 39—41), is more or less circular in section, and the bone tapers 
fairly evenly to the middle of the shaft where the smallest diameter is 
reached; there being only a very small crest on the ventro-mesial 
surface. The epiphysial cartilage of the proximal end is slight and 
saucer-shaped; its concavity is shallow. The distal end of the radius 
expands in an axial direction so that, when seen from the extensor 
or flexor surfaces, it is roughly triangular in shape. The cross-section 
of this end is approximately rectangular, and the cartilaginous 
epiphysis is more extensive on the pre-axial side than on the post- 
axial. It articulates with the pars intermedia of the fused ulnare 
+ intermedium and also with the radiale. 

The ulna (figs. 25-7), in contrast to the radius, is larger at its 
proximal end and smaller distally. The club-shaped proximal end 
is trapezoid in section, the longest side being mesial. The epiphysis 



42 THE SALAMANDER 

which articulates with the ulnar condyle of the humerus and bears 
an olecranon process is entirely cartilaginous. The central portion 
of the shaft is very slender, and is roughly triangular in transverse 
section, with a slight bony crest running along the flexor side. The 
distal end is but slightly expanded, and articulates only with. the 
ulnare + intermedium. 

To summarize: the radius and ulna are entirely separate. The 
greatest expansion of the radius is distal, and in an axial direction, 
while the ulna expands proximally, and in a vertical plane. Both 
elements articulate with the humerus as well as with the carpus. 

(c) The Carpus (fig. 28) of the adult Salamander consists of 
seven elements, namely, the radiale^ a fused ulnare -\-iniermedium, a 
centrale^ and four basalia. Much has been written concerning the 
homologies of these basal elements and of the four digits which they 
bear. The older authors, e.g. Gegenbaur, Wiedersheim, and Hoff- 
mann, supposed that the hand lacked the pre-axial digit, or pollex — 
apparently by analogy with the normal reduction sequence in the 
mammalian carpus. When, however, the carpus of the Salamander, 
or of any other four-fingered Urodele, is compared with the tarsus, 
which bears five toes, a strong similarity is to be observed in the pre- 
axial sides of the two limbs. Further, in some forms, e.g. Triton and 
Diemyctylus, there are evident traces of a reduction occurring in 
the post-axial region of the tarsus (cf. Steiner, 1921). Both Perrin 
and Sieglbaur have concluded, from a comparative study of the 
muscles of the hand and foot of Salamandra, that the pre-axial digits 
of each are serially homologous, and that therefore it is the post- 
axial, or fifth finger, which is missing from the hand, and this is the 
view adopted here. It should be mentioned that both Ribbing and 
Eisler, in spite of having investigated the muscles and nerves in 
detail, apparently retain the older view, although they do not directly 
discuss the question. 

Turning now to the carpalia themselves, the most pre-axial of the 
basal carpals (c.pr.p.) was considered by Gegenbaur, Wiedersheim, 
and the earlier authors to be the basal of the second digit (they con- 
sidered the first digit to be missing entirely) pushed a little out of its 
place. Strasser (1879), however, by studying the development in 
Triton and Salamandra, showed that primitively this cartilage has 
no connexion with the pre-axial digit, and that the next basal (his 
Basale commune) has a double origin and is always related, in the 
larva as well as in the adult, to each of the first two digits. He therefore 
concludes that this element represents basalia i and 2 fused together. 
In 1886 Kehrer suggested that the preaxial cartilage above referred 



THE SKELETON 43 

to is really the basal oi 2. pre-pollex, oi -which, digit no further trace has 
remained. This view was supported by Baur (1888), Rabl (1901), 
Steiner (1921), and others. 

So far as the present investigation of Salamandra is concerned the 
description of the anatomical relations given by these authors has 
been entirely confirmed, i.e. that the first two digits articulate with 
a common basal, and that the first basal is not at all closely associated 
with the first digit, and lies rather more proximally than the other 
basal elements. Further, in a specimen in which the bones had been 
stained with alizarin, and the soft tissues cleared with potassium 
hydroxide and glycerine, it was apparent that the basal common to 
the first two digits had ossified from two centres, in fact it presented 
an appearance comparable with that of the fused ulnare + inter- 
medium. Finally, the fact that the short flexor muscles for the first 
digit take their origin from this element and not from the most pre- 
axial cartilage seems to add conclusive evidence in favour of 
Strasser's view. Various other authors have made contributions 
bearing on the subject, and their views may be summarized, accord- 
ing to whether they support or oppose the conclusions set forth 
above, as follows. 

In support: Strasser (1879); Kehrer (1886); Baur (1888); 
Emery (1895); Zwick (1898); Rabl (1901); Sewertzoff (1907); 
Steiner (1921). 

Opposed : Gegenbaur ( 1 8 64) ; Goette (1879); Braus ( 1 906). 

It must be pointed out that Sewertzoff and Zwick do not regard 
the cart, pre-pollicis as having such a status, but rather suppose it 
to be a supernumerary centrale. 

In the carpus of Salamandra we have therefore (if the above inter- 
pretation be accepted) in the distal row of carpalia, starting from the 
pre-axial border, a cartilago pre-pollicis (c.pr.p.) which remains carti- 
laginous throughout life; a fused basale 1+2 (o.b.c. 1+2) {Easale 
commune^ Strasser), with an osseous centre in the adult, serving the 
first two digits ; a basale 3 for the third finger, and a basale 4 for the 
fourth, both of which also become ossified in the centre. There is a 
bony centrale (o.c.) which articulates with all the other elements of 
the carpus with the exception of the pars ulnaris (o.u'+i.) of the 
ulnare + intermedium. The radiale (c.rd.) remains entirely cartila- 
ginous. It is relatively small and articulates proximally with the 
radius, mesially with the pars intermedia (o.u.-j-i'.) of the ulnare + 
intermedium, and distally with the centrale and cart, pre-pollicis. 
As already indicated the ulnare and intermedium fuse in the adult 
to form a common element, of which the major portion ossifies. The 



44 THE SALAMANDER 

line of fusion between them is indicated by a foramen, the /or. per- 
jorans carpi (fo.per.c), through which the A. perforans carpi passes 
from the ventral to the dorsal side. It is by far the largest element in 
the carpus, and articulates with the ulna, radius, radiale, centrale, and 
basale 4. 

{d) The Metacarpals and Phalanges are all dumb-bell-shaped 
bones with cartilaginous epiphyses. The digital formula is 1.2.3.2. 
The homology of the fingers is fully discussed above. 

11. THE PELVIC GIRDLE AND HIND-LIMB (PL V). 
I. The Pelvic Girdle (Figs. 34 and 35). 

This girdle is a much firmer structure than the pectoral. It is 
firmly attached dorsally to the vertebral column, while ventrally the 
two halves meet one another in a median symphysis. The consti- 
tuents of the girdle are dorsally a club-shaped member, the ilium^ 
and a ventral irregularly shaped plate, the composite pubo-ischium. 
Anterior to this latter portion is a median-forked cartilage, the 
cartilago ypsiloidea^ which is only doubtfully to be regarded as a part 
of the pelvic girdle proper. 

The ilium (oaX.) (os ischii. Funk) projects dorsally from the aceta- 
bulum with a slight backward inclination, and is firmly attached to 
the rib of the sacral vertebra by fibrous tissue. It is a club-shaped 
structure with the 'head' directed dorsalwards. The dorsal extremity 
remains cartilaginous (= the epileon of Sabatier), but the remainder 
ossifies with the exception of a small area at the junction with the 
pubo-ischium. 

Th.Q pubo-ischium (ilio-pubis. Funk) is an irregularly shaped plate, 
approximately quadrangular, forming an inverted arch in the trans- 
verse plane. The posterior or ischial part of the plate is always 
ossified — except at the symphysis — while the anterior or pubic por- 
tion is usually pure cartilage, although it may show a varying amount 
of ossification (cf. Wiedersheim, 1892). In the latter case it is im- 
possible to distinguish the boundary between the pubis and ischium 
in the adult. The pubis is perforated by a small obturator foramen 
(fo.o) for N. obturatorius. Anteriorly the lateral part of the pubis is 
prolonged to a short pre-pubic process (pr.p-p.) while the lateral 
portion of the ischium projects posteriorly in a similar manner 
forming an ischial spine. 

Both portions of the ventral plate as well as the ilium contribute 
to the acetabulum (ac.) (cavite cotyloide, Perrin), which is large and 
roomy, though the rim is not complete at the antero- and postero- 



THE SKELETON 45 

dorsal margins. There are three prominences around its periphery, 
one dorsal arising from the ilium, one antero-ventral formed from 
the pubis, and one postero-ventral projecting from the ischium. 
The head of the femur is firmly attached to the two former points 
by a pair of cotyloid ligaments. 

The Cartilago Tpsiloidea (fig. 36) (Townson, 1795). 

Cart, marsupial ....... Duges (1834). 

Abdominal sternum ..... Sabatier (1880). 

Cart, epipubes . . Wiedersheim, Hoffmann (1876), and others. 

Cart, ypsiloides . . . Hoffmann (1873-8); Whipple (1906). 

As its name implies this is a Y-shaped cartilage lying in the middle 
line, anterior to the pubo-ischium with which it articulates. It lies 
dorsal to the M. rectus abdominis and is closely associated with it. 
As indicated by the synonyms, this cartilage has been homologized 
with various structures. Whipple (1906) investigated its develop- 
ment, on a limited supply of material, and concludes that it arises in 
the linea alba as the result of muscle strain, and has nothing to do 
with the pelvic girdle. Against the view held by Wiedersheim 
and Hoffmann that this cartilage is homologous with the epipubis 
of reptiles, she advances the argument that the cart, ypsiloidea 
develops and remains entirely dorsal to the rectus muscle, whereas 
the marsupial bone is ventral to it. 

From observations and experiments on aquatic Salamanders 
(Diemyctylus) Whipple arrived at the conclusion that, in these 
forms, the ypsiloid apparatus is definitely correlated with the hydro- 
static function of the lungs. Thus, by raising the ypsiloid cartilage 
the animal is able to compress the posterior end of the body cavity 
and so force the air in the lungs forwards to the anterior end, thus 
causing the animal's head to rise in the water, while by lowering the 
cartilage the air is brought back to the posterior end and the head 
of the animal sinks so that it is able to swim to the bottom. This 
explanation is of course irrelevant as far as Salamandra is concerned, 
since this animal is purely terrestrial in the adult stage. Nevertheless 
it is not unreasonable to suppose that it may be of assistance in 
defaecation and parturition by virtue of its capacity to compress the 
posterior abdomen. 

2. The Hind-Limb (Figs. 29-33). 

The form of the hind-limb corresponds with that of the fore-limb 
except that it bears five digits instead of four. For descriptive pur- 
poses it may be divided into three regions : {a) the thigh^ containing 



46 THE SALAMANDER 

the femur as its skeletal element; {b) the leg^ with the tibia z.rvAfibula\ 
and {c) the /<2rjz/j and digits supported by the tarsals^ meta-tarsals^ and 
-phalanges. As in the fore-limb all epiphyses and articular surfaces 
remain cartilaginous. 

{a) The Femur (Figs. 3 1-3). The head of the femur is rounded 
but not hemispherical, since there is a trochlear groove on its 
postero-ventral surface which articulates with the ischial process on 
the rim of the acetabulum, while the dorsal and antero-ventral sur- 
faces are somewhat flattened and possess well-marked foveal de- 
pressions (fv.c.fm.) wherein are inserted the ligaments attaching the 
bone to the acetabulum (l.pb. and l.il.). In transverse section, there- 
fore, the proximal end of the bone is roughly triangular, with the 
sides facing dorsal, antero-ventral, and postero-ventral. There is 
no neck, and the articular surface is terminal. There is a strong 
hook-shaped trochanter (pr.tch.) directed antero-mesially on the 
ventral surface, towards which, from either side of the trochlear 
groove, run two trochanteric crests^ of which the posterior is the 
less developed. The cross-section of the bone beyond the hook is 
approximately circular but it soon expands into a rectangle of which 
the long sides are dorsal and ventral . There are two articular condyles 
distally, the pre-axial or tibial being the larger, although neither 
is so pronounced as the corresponding surfaces of the fore limb. 
The longitudinal axis of the bone is approximately straight when 
seen from above, but shows a double curvature if viewed in an 
axial direction. 

{b) The Tibia (o.tb.) is a straight simple bone whose section is 
more or less rectangular at the ends and circular at its narrowest 
part, which is at about one-third of its length from the distal end. 
The long sides of the proximal end are on the flexor and extensor 
surfaces, but at the distal end the long axis of the rectangle is at right 
angles to these. A prominent /z^zW <:r(?j-/ (cr.tb. fig. 29) arises from 
the dorso-mesial side of the extensor surface which finally becomes 
independent of the surface of the bone so as to give it a thorn-like 
appearance. It receives the insertion tendon of the M. extensor 
ilio-tibialis. Distally the tibia articulates with the tibiale and inter- 
medium. 

(c) The Fibula (o.f.) is a rather more slender bone than the tibia 
but of about the same length. It is roughly triangular in section with 
the base directed mesially. Its distal end is very oblique and articu- 
lates with the fibulare and intermedium. 

{d) The Tarsus (Fig. 29 and 30). The fibulare (o.fb.) is the 
largest of the proximal row of tarsal elements, and is not fused with the 



THE SKELETON 47 

intermedium like the ulnare in the carpus. The intermedium (o.i.) 
is a wedge-shaped bone of which the thinner edge projects between 
the tibia and fibula, thus separating the distal ends of these two bones. 
Distally it articulates with the centrale, fibulare, and tibiale. Its 
post-axial border is slightly notched to allow the A. perforans tarsi to 
pass from the ventral to the dorsal side of the ankle. The tibiale 
(c.t.) is the smallest element of the proximal row. It is helmet- 
shaped and articulates with the tibia, intermedium, centrale, and 
the cartilago pre-hallucis. Normally it remains cartilaginous. The 
centrale (o.c.) is very large and articulates with all the other elements 
of the tarsus except the basal tarsal of the fifth digit. The basalia of 
the tarsus are strictly comparable with those of the carpus. The pre- 
axial represents the basal of 2. pre-hallux (c.pr.h.), the next is a basale 
commune (o.b.t. i +2) bearing the first two digits, while the remaining 
three are smaller, each supporting one digit only. All become more 
or less ossified except the basal of the pre-hallux. The meta-tarsals 
2ind. phalanges are all dumb-bell-shaped bones and the digital formula 
is 1.2.3.3.2. 

BIBLIOGRAPHY OF SECTION III 

8,9, 11,24,33,63,71,81,82,84, 109, 148, 163, 167, 170, 171, 172, 196, 
202, 203, 204, 215, 216, 221, 224, 225, 227, 229, 230, 231, 232, 251, 257, 
258, 260, 267, 329, 330, 344, 345, 365, 367, 370, 371, 374, 41 1, 426, 427, 
432, 456, 460, 488, 490, 491, 494, 495, 499, 506, 519, 535, 557, 561, 562, 
569, 571, 590, 591, 607, 713, 714, 716, 717, 718, 721, 729, 740, 757, 764, 
766, 767, 783, 787, 788, 790, 791, 792, 793, 8i I, 812, 813, 814, 815, 816, 
817,818,840. 



IV 
THE MUSCLES 

General Literature. 
Most of our modern knowledge of the muscles of the Salamander 
is due to the work of specialists who, taking some small part of the 
body, have endeavoured to work out the comparative morphology 
of the muscles concerned throughout the vertebrate series. Never- 
theless a few of the earlier workers have described the chief muscles 
in a general sort of way, naming them according to their position, or 
to their apparent resemblance to a muscle in the human body. Thus 
Funk (1827) gives quite a passable account of all the principal 
muscles, and a year later Carus (1828) also described and figured 
them very well indeed. Duges (1834) names and figures a large 
number, but his text is so scanty that it is frequently impossible to be 
sure of the identity of the muscle concerned. In 1852 Rymer Jones, 
in his article on 'Reptilia' in Todd's Cyclopaedia of Anatomy and 
Physiology^ gives two figures of the muscles of Salamandra (taken 
from Carus without acknowledgement) and describes the principal 
muscles. Some idea of the extent of the work of these authors may 
be obtained from the lists of synonyms preceding the account of 
each muscle described below. The endeavour has been made to get 
these lists as complete as possible, and in addition to the names 
actually applied to the muscles of Salamandra itself those given to 
the homologous muscles of Cryptobranchus by Humphry, Schmidt, 
Goddard and van de Hoeven, and by Osawa, have also been 
appended for comparative purposes. The figures reproduced in 
Owen's text-book, although credited to Funk by Owen, do not 
occur in Funk's monograph but are due to Carus (1828). 

A review of the more specialized literature precedes the anatomical 
description in each of the subsequent sections. 

MUSCLES OF THE HEAD 
I. Historical. 

Rusconi (i 854) gives a very good account of the visceral muscles 
and of the changes which they undergo during metamorphosis. His 
work is general rather than detailed, as would be expected from its 
date. In 1887 Walter also gives a good account of the visceral 



THE MUSCLES 49 

skeleton and its related muscles in Urodeles, Anura, and Reptiles, 
but it was Wilder (1892 and 1896) who first gave an exact descrip- 
tion of the larynx and its muscles. He endeavoured to homologize 
the latter with those of the visceral arches. A brief resume of his 
hypothesis and of those of subsequent workers in this field is given 
on p. 272. Further contributions to the discussion of the homologies 
of the laryngeal muscles were made by Goppert in 1894 and 1898. 
In 1902 Burkard undertook a comparative study of the muscles 
surrounding the orbit in the various classes of vertebrates, and 
included a description of the M. levator bulbi of Salamandra. A 
very detailed and exact description of the muscles supplied by the 
Vllth, IXth, Xth, and hypoglossal nerves in Salamandra and Triton 
was published by Driiner in 1901. He also describes the nerves 
themselves and the hyobranchial skeleton. Two years later (1903), 
and again in 1904, Driiner supplemented this account by describing 
the same region in other Urodeles, and included some theoretical 
considerations on the homology of the chorda tympani, and of the 
larynx. His whole work is a classic and very accurate. A compara- 
tive study of the jaw-muscles of vertebrates was published by 
Lubosch in 1913 and 19 14, and while he does not deal with 
Salamandra itself, his work is important from a comparative point 
of view. In 19 14 Luther described the muscles supplied by the 
Vth nerve, which include the M. levator mandibulae and the M. 
levator bulbi. He deals with Salamandra. Lastly Edgeworth (1920 
et seq.) has published many papers dealing with the development 
and homologies of the muscles of the head in Amphibia and other 
vertebrates, and was the first to show that the larynx and its muscles 
have no phylogenetic relationship with the visceral arches. It is 
Edgeworth's nomenclature which has been adopted here. 

2. Muscles of the Eye (PI. XIII, fig. 57). 

These are eight in number, namely, four recti, two oblique, 
together with the MM. levator and retractor bulbi. Of these, 
seven are described below, while the eighth — the M. levator 
bulbi — belongs to the muscles of the Vth nerve and is described on 
P-55- 
M. rectus superior (m.r.s.). 

This muscle arises from the dorso-posterior angle of the orbit and 
runs antero-laterally, broadening somewhat at its insertion on the 
dorsal section of the ambitus of the bulbus oculi. Its origin is 
tendinous. 

4038 Tj 



50 THE SALAMANDER 

Innervation: By the superior ramus of N. oculomotorius (third 
cranial nerve). 

Function: See below. 

M. rectus posterior (m.r.p.). 

The abducent muscle arises as a fine tendon from the orbito- 
sphenoid at about a third of its length from the posterior end, just 
behind the M. rectus inferior. It passes obliquely postero-laterally, 
ventral to the M. retractor bulbi, and is inserted in the postero- 
lateral portion of the eyeball. Like all the rectus muscles it broadens 
considerably towards its insertion. 

Innervation: From the abducent nerve (sixth cranial). 

Function: See below. 

M. rectus inferior (m.r.i.). 

The inferior rectus muscle also has a tendinous origin just anterior 
to that of the posterior rectus and posterior to the anterior rectus. It 
is considerably shorter than the muscle just described, and some- 
what broader. It runs laterally, its fibres lying almost at right angles 
to the axis of the skull. Its insertion occupies the ventral section of 
the ambitus of the bulbus. 

Innervation: From the inferior ramus of the N. oculomotorius 
(third cranial nerve). 

Function: See below. 

M. rectus anterior (m.r.a.). 

As mentioned above, the tendon of this muscle leaves the orbito- 
sphenoid just anterior to that of the inferior rectus. Its fibres take 
a direction practically at right angles to those of the former muscle 
and lie close along the side of the skull. They are inserted on the 
antero-mesial surface of the bulbus oculi. It is the largest and 
strongest of the rectus muscles. 

Innervation: From R. inferior N. oculomotorius. 

Function : The function of all the rectus muscles is obviously to 
rotate the eyeball in the horizontal and vertical planes at right angles 
to its own optical axis. They are all relatively weak. 

M. obliquus superior (m.o.s.). 

Arises as a fine tendon from about the centre of the cartilago 
antorbitalis of the nasal capsule. Its fibres run posteriorly and are 
inclined slightly mesialwards. They are inserted on the antero- 
dorsal portion of the eyeball just anterior to those of the M. rectus 
superior. The muscle lies dorsal to the main portion of the 



THE MUSCLES 51 

N. ophthalmicus profundus (Vj) although an important cutaneous 
branch of this nerve crosses its dorsal side (see p. 139). 

Innervation: By N. trochlearis (fourth cranial nerve). 

Function : See below. 

M. obliquus inferior (m.o.i.). 

Like its superior counterpart this muscle arises from about the 
middle of the cart, antorbitalis. It passes in a postero-ventral direc- 
tion and is inserted on the ventral side of the eyeball. It has rather 
a long insertion on a line approximately at right angles to the optical 
axis, so that the lateral fibres are much shorter than the mesial ones. 

Innervation : From the R. inferior N. oculomotorius. 

Function: The oblique muscles serve to rotate the eyeball about 
the optical axis. Both are, however, very weak. 
M. retractor bulbi {or ocult) (m.rt.b.). 

This is by far the most powerful of all the eye-muscles. It takes 
its origin in part from the lateral process of the parasphenoid that 
lies under the otic capsule, and in part from the orbito-sphenoid at 
the inner ventral angle of the orbit. Its fibres spread out to form a 
cone around the optic peduncle and are inserted on the mesial surface 
of the bulbus oculi within the ring formed by the rectus muscles. The 
cone is, however, not complete, since the edges of the muscle are not 
quite in contact mesially in the region of the anterior rectus muscle. 

Attached to the M. retractor bulbi is a strong tendon connecting 
it with the eyelids. This tendon forms a broad sheet posteriorly 
but bifurcates anteriorly to allow for the passage of the anterior 
rectus muscle. When this tendon is pulled the edges of the eyelids 
are brought together and the eye closed, and although there appear 
to be smooth muscle elements in the upper lid rendering it capable 
of independent movement, nevertheless the M. retractor bulbi and 
the tendon must be regarded as the chief agents effecting the closing 
of the eye, which is thus made to occur simultaneously with the 
retraction of the bulbus oculi. 

Innervation : From the N. abducens (sixth cranial nerve). 

Function: To retract the bulbus and to close the eyelids as de- 
scribed above. 

3. Muscles of the Fifth Nerve. 

M. levator mandibulae as a whole. 

Elevateurs de la machoire inferieure . . . Meckel (18 18). 

Temporalis .... Cuvier (1835); Owen (1866). 

Adductor mandibulae ..... . Driiner (1901). 

Masseter Wilder (189 1). 



Owen ( 1 866). 

Plessen & Rabinowicz (1891). 

Lubosch (19 14). 

. Luther (1914). 



52 THE SALAMANDER 

This muscle Is divided by all modern authors Into three main 
sections which are named according to their position relative to the 
mandibular branch of the Vth nerve. 

M. levator mandihulae anterior (Edgeworth). 

Temporalis v. Siebold (1828); Cuvier (1835); Fischer (1843); 

Schmidt, Goddard, and van d. Hoeven (1864); 

Mivart (1869); Druner (1901); Coghill (1901-6); 

Osawa (1902) 
Post-orbito-coronoidien et Atlanto-corono'idien . . Duges (1834). 

Pterygoidien Rusconi (1854). 

Atlanto-mandibularis 
Pterygo-temporalis 
Cranio-mandibularis (partim) . 
Adductor mandibulae internus 

This muscle lies wholly mesial to the mandibular branch of the 
trigeminal nerve (V3). It is divisible into two distinct portions, 
and most modern authors have thus distinguished a superficial and 
a deep layer. 

M. levator mandihulae anterior {superficial portion) (Edgeworth). 

Temporalis (3rd portion) Funk (1827); Cuvier (1835); Fischer (1843); 
Lubosch (19 1 4). 

Crotaphite Rusconi (1854). 

Pseudo-temporalis ...... Luther (19 14). 

This is a long, narrow, conical muscle which arises partly from the 
lateral border of the parietal^ and partly from the dorsal fascia that 
extends from the skull to the neural spine of the first vertebra, but 
mainly from a strong raphe which passes from the neural spine of the 
first vertebra through the centre of the muscle, the fibres arising 
from either side of it, forming a pinnate structure. The muscle 
crosses dorsally over the skull in the otic region, and then turns 
downwards in front of the ear capsule almost at right angles, and 
passes into a strong tendon which is inserted in the coronoid process 
of the pre-articular element of the lower jaw. 

M. levator mandibulae anterior {deep portion) (Edgeworth). 

Frontales Funk (1827). 

Pterygoideus Cuvier (1835); Fischer (1843); Owen (1866); Druner 

(1901); Lubosch (1914); Luther (1914). 

Pterygoidien Rusconi (1854). 

Pterygo-maxillaris ...... Hoffmann (1873-8). 

A short, fan-shaped, almost vertical muscle. Its fibres arise from the 
lateral borders of the frontal and parietal elements of the skull and 



THE MUSCLES 53 

converge to a long band-like tendon which is attached to the dorsal 
margin of the pre-articular, immediately anterior to the articulation 
of the jaw. It is largely covered by the superficial portion, the fibres 
of which tend to mingle with it near its insertion. 

Innervation: Both portions of the muscle are innervated by a 
special branch of the trigeminus nerve which leaves it either just 
before, or just after it emerges from the antrum petrosum laterale. 

Function: Both portions assist in closing the mouth. 

M. levator mandibulae posterior (Edgeworth). 

Cranio-mandibularis (partim) .... Lubosch (19 14). 

Adductor mandibulae posterior .... Luther (19 14). 

A somewhat triangular-shaped muscle arising from the free anterior 
edge of the squamosal^ the anterior face of the quadrate^ and the 
■pterygoid. It is traversed by the mandibular nerve (V3) so that, while 
some of its fibres lie mesial, the greater portion of the muscle lies 
posterior to that nerve. A portion of the muscle has a fleshy inser- 
tion on the articular portion of Meckel's cartilage, while a tendon 
passes forwards from the anterior fibres to a point immediately in 
front of the tendon of the M. levator mandibulae anterior (superficial 
portion), some of the mesial fibres being actually inserted on the 
tendon of this muscle. The general direction of the fibres is, there- 
fore, antero-ventral, the most anterior fibres being of moderate 
length and the most posterior extremely short. 

Innervation: By fine twigs from the mandibular nerve. 

Function: To assist in closing the jaws. 

M. levator mandibulae externus (Edgeworth). 

Masseter Funk (1827); v. Siebold (1828); Fischer (1843); Rusconi 

(1854); Schmidt, Goddard and van d. Hoeven (1864); 
Owen (1866); Mivart (1869); v. Plessen & Rabinowicz 
(1891); Druner (1901); Coghill (1901-6); Osawa (1902). 

Temporal, portion posterieur courte . . . Cuvier (1835). 

Pre-temporo-maxillaire ...... Duges (1834). 

Petro-tympano-maxillaris .... Hoffmann (1873-8). 

Mandibularis externus . ..... Lubosch (19 14). 

Adductor mandibulae externus .... Luther (1914). 

This muscle lies anterior to the one just described and entirely 
lateral to the mandibular nerve, the direction of its fibres being 
parallel with those of the M. lev. mand. post. It takes its origin 
from the antero-lateral edge of the squamosal and from the anterior 
wall of the ear capsule, and is inserted on the posterior end of 
the dentary and the lateral face of the coronoid process of the 



54 THE SALAMANDER 

pre-articular. The skin at the corner of the mouth is strongly 
reflexed dorsalwards and is firmly bound to the anterior portion of 
this muscle — in fact some of the fibres seem actually to be inserted 
on it. Like that of the previous muscle the insertion is fleshy 
posteriorly with a tendency to become tendinous anteriorly. 

Innervation : By fine twigs from the mandibular nerve (V3). 

Function: To assist in closing the mouth. The M. levator mandi- 
bulae in general is opposed by the M. depressor mandibulae, a 
muscle supplied by the Vllth nerve. 

M. intermandihularis (Edgeworth) (PI. VI, fig. 37). 

Mylohyoideus . v.Siebold (1828); Carus(i828)j Stannius (1854); 

Owen (1866); Coghill (190 1-6). 
„ (et interhyoideus) .... Walter (1887). 

„ (et interhyoideus et interhyoideus 

posterior) ..... Funk (1827). 

As its name implies this is a sheet of muscular tissue stretching 
across the floor of the mouth between the rami of the lower jaw. It 
lies immediately under the skin. In the larva it consists of two dis- 
tinct parts, a small anterior portion lying in the angle of the chin, 
and a larger posterior portion. It is the latter only which is found in 
the adult after metamorphosis, but the synonyms of the anterior 
portion are given below for the sake of completeness. 

M. intermandihularis anterior. Driiner (1901); Luther (19 14); 
Edgeworth (1920 et seq.). 

Submentalis . . Fischer (1843); Mivart (1869); Osawa (1902). 

Genio-glossus . Schmidt, Goddard and van d. Hoeven (1864). 

Submaxillaris ...... Hoffmann (1873-8). 

Camv (antr.) Ruge (1897). 

A purely larval muscle, aborting at metamorphosis. 

M. intermandihularis -posterior (m.i.m.). Driiner (1901); Luther 
(19 14); Edgeworth (1920 et seq.). 

Mylohyoideus anterior Fischer (1843); Mivart (1869); Osawa (1902). 
Partie anterieure du mylohyoidien . . Rusconi (1854) 

Mylohyoideus . Schmidt, Goddard and van d. Hoeven (1864). 

Intermaxillaris anterior . . Hoffman (1873-8); v. Plessen and 

Rabinowicz (1891). 
C2mv(post) Ruge (1897). 

From the above list of synonyms it will be seen that certain authors 
have called this the anterior part of the muscle, owing to the fact that 
they have mistaken the M. interhyoideus for the posterior part. 



THE MUSCLES 55 

The M. interhyoideus is a Vllth nerve muscle, however, and its 
connexion with the mandible is secondary. 

The M. intermandibularis posterior arises from the anterior two- 
thirds of the mesial upper edge of each ramus of the mandible except 
for a very short distance near the middle line anteriorly, which latter 
gives rise to the M. genio-glossus. The fibres run transversely across 
the space between the two rami of the lowerjaw, but are interrupted in 
the middle line by a wide aponeurosis of tough transparent connec- 
tive tissue, the linea alba. In a fresh specimen the MM. genio- 
hyoideus and genio-glossus are seen through this aponeurosis and 
the M. intermandibularis looks as though it were inserted into their 
edges. 

Innervation : From N. mandibularis (R. intermandibularis). 

Function: The contraction of this muscle serves to elevate the 
mouth floor, and it is therefore of service in breathing and degluti- 
tion. 

M. levator bulbi (^\. IX, fig. ^2)- Hoffmann (1873-8); Driiner 

(1901); Burkard (1902); Luther (1914); Edgeworth (1920 et seq.). 

Fronto-pterygoidien ...... Duges (1834). 

An excellent description of this muscle has been given by Luther 
(i 9 14) and his nomenclature has been adopted here. His anatomical 
results, as far as the adult is concerned, are accurate, but his conclu- 
sions regarding the phylogenetic history of the muscle do not agree 
with those of Edgeworth. 

It is by no means easy to determine the direction of the fibres 
in the several portions of this muscle, but the application of a little 
dilute saffranin is of material assistance. 

The muscle consists of a thin sheet forming an elastic floor to the 
orbit and lies between the bulbus oculi and the roof of the mouth, 
ventral to the MM. rectus and obliquus inferior. Luther distin- 
guishes the following three sections. 

{a) ThQ pars principalis (m.l.b.p.) is the most dorsal portion, lying 
nearest to the bulbus oculi. It arises as a thin, tough, tendinous sheet 
from the lateral borders of the parietal^ frontal^ and pre-frontal ele- 
ments of the skull as far as the cartilago antorbitalis of the nasal capsule. 
The fibres run mainly in a postero-lateral direction and converge so 
as to form a more or less scimitar-shaped plate. They are inserted 
on the ventral side of the posterior end of the lower eyelid (mem- 
brana nictitans). 

(J?) The pars sagittalis arises from the palato-quadrate at the pos- 
terior wall of the orbit, and may be further subdivided into two heads. 



56 THE SALAMANDER 

(i) The fibres of the caput laterale (m.l.b.s'.) pass obliquely antero- 
laterally and are inserted along the lower eyelid from a point immedi- 
ately anterior to the insertion of the pars principalis to about the 
centre of the eyelid. 

(ii) The fibres of the caput me di ale (m.l.b.s".) pass antero-mesially 
and are attached to the side wall of the skull immediately ventral 
to the origin of the pars principalis. This portion loops across the 
origin of the M. retractor bulbi. 

(<:) The pars transversalis (m.l.b.t.) takes its origin from the side 
wall of the skull — mainly from the lower edges of the frontal and 
parietal — anterior to the insertion of the caput mediale^ pars sagittalis^ 
and ventral to the origin oi\h^ pars principalis. Its fibres are the most 
ventral of all, and pass across the anterior portion of the floor of the 
orbit forming an arch which is convex anteriorly. They are inserted 
on the lower eyelid anterior to the caput laterale^ pars sagittalis^ and 
on the maxilla. 

Innervation : All sections are innervated by the second branch of 
the trigeminus nerve (Vg) in the Salamander. 

Function'. The muscle as a whole appears to have at least two 
principal functions : (i) as levator bulbi, (ii) as accessory respiratory 
muscle by enlarging the mouth cavity. It may also serve as an 
adductor of the lower lid, but the movement thereby caused must 
be very slight, even if it exists at all (cf. Johnson, 1926). 

4. Muscles of the Seventh Nerve. 

As already mentioned, an accurate description of these muscles in 
both larva and adult, illustrated by excellent figures, was published 
by Driiner in 1901. He divides the muscles supplied by the Vllth 
nerve into two groups, according to whether they are supplied by the 
nerve before., or after^ it has received the R. communicans IX+X ad 
VII, on the supposition that this ramus contains motor fibres. By 
comparison with the condition in other Urodeles this conjecture 
appears unwarranted, and it seems more likely that the R. communi- 
cans contains only general cutaneous and communis fibres, and there- 
fore has no influence on the muscular supply. Driiner's classification 
is accordingly unnecessary. 

M. depressor mandihulae (Edgeworth). 

Depressor maxillae inferioris .... v. Siebold (1828). 

Temporo-angulaire ...... Duges (1834). 

Digastricus Fischer (1843); Stannius (1854-6); Osawa (1902}. 

Digastrique .... Cuvier (1835); Rusconi (1854). 



THE MUSCLES 57 

Occipito-mandibularis s. Digastricus . . . Owen (1866). 

Digastric Mivart (1869). 

Digastric et Depressor mandibulae , . , Humphry (1872). 

Cephalo-dorso-maxillaris s. Digastricus-maxillae Hoffmann (1873-8). 

Camd Ruge (1897). 

Cephalo-dorso-mandibularis ..... Driiner (1901). 
A thick, powerful muscle originating from the posterior edge of 
the squamosal, the 'crista muscularis' of the ear capsule, and 
the anterior portion of the fascia cephalodorsalis (Driiner). This 
superficial dorsal fascia gives origin to a number of muscles. It 
is firmly united to the skin and extends backwards to behind the 
shoulder. Edgeworth (i 93 1) has shown that the M. depressor man- 
dibulae is derived from the M. levator hyoidei of the Dipnoi. 

Innervation: From R. jugularis VII by a fine branch which separ- 
ates from the main nerve immediately distal to the point at which the 
R. communicans IX+X ad VII is received. It enters the muscle 
mesially. 

Function: As its name implies, it is the opening muscle of the lower 
jaw and opposes the levator mandibulae. 

The M. interhyoideus (PI. VI, fig. 37) is the next muscle to be 
considered. It undergoes a rather considerable change during meta- 
morphosis, and becomes divided into two portions. The anterior 
portion retains more or less the original position and therefore 
retains also the original name — M. interhyoideus, while the posterior 
portion shifts backwards and hence becomes the M. interhyoideus 
posterior. Most of the earlier authors regarded the whole muscle as 
the posterior part of the 'Mylohyoideus'. Both portions together are 
spoken of as 'Duo-cutis musculi oriuntur in maxillae inferioris unco' 
by Carus (1828). 
M. interhyoideus (Edgeworth) (m.i.hy.). 

Constrictor pharyngis internus . . . v. Siebold (1828). 

Mylohyoidien, portion moyenne .... Rusconi (1854). 

Cghv Ruge (1897). 

Inter-ossa-quadrata ...... Driiner (1901). 

The origin of this muscle is by means of a short thin tendon from 
the postero-mesial edge of the quadrate, just ventral to the point 
of insertion of the ligamentum hyoquadratum. According to Driiner 
some fibres may occasionally arise from the ligament itself. The 
fibres spread out fan-wise over the hinder part of the mouth-floor, 
and are inserted into a posterior continuation of the median apo- 
neurosis of the M. intermandibularis posterior. The direction of the 
most posterior fibres is thus almost directly transverse and that of 



58 THE SALAMANDER 

the anterior ones obliquely antero-mesial. The mesial ends of the 
anterior fibres lie dorsal to the M. intermandibularis posterior. 

While this portion of the muscle retains its original position, its 
origin has nevertheless shifted, and has migrated from the posterior 
end of the hyoid arch, passing up the lig. hyoquadratum to the 
quadrate. 

Innervation'. By the terminal branches of R. jugularis VII which 
spread over its ventral side. 

Function'. It constricts the hyobranchial skeleton and posterior 
part of the mouth, and thus assists both in respiration and deglu- 
tition. 



M. interhyoideus -posterior (Edgeworth) (m.i.hy.p.). 

Constrictor pharyngis externus 
Mylo-hyoidien, partie posterieure 
Mylosternoideus 



V. Siebold (1828). 
. Rusconi (1854) 
. Walter (1887}. 
Ruge(i897). 
Quadrato-pectoralis ...... Driiner (1901). 

The posterior portion of the interhyoid muscle arises by means of 
a strong tendon from the ventro-lateral surface of the quadrate and 
the distal end of the squamosal. Its fibres spread out fan-wise across 
the throat ventral to those of the preceding muscle. The muscle 
becomes joined to the capsule surrounding the articulation of the 
jaw as it passes laterally across it. The direction of the anterior fibres 
is almost transverse, and they are inserted in backward continuation 
of the same aponeurosis as receives the MM. intermandibularis and 
interhyoideus. The most posterior fibres, forming the bulk of the 
muscle, turn more postero-mesially and are inserted into the skin at 
the gular fold. This fold of skin surrounds the throat on the ventral 
side, and represents the remains of the gill operculum of the larva 
fused with the skin of the pectoral region. To the same fold, posteriorly, 
is attached the base of a triangular aponeurosis — th.e. fascia pectoralis 
(Driiner) — the apex of which is directed backwards and is attached 
to the most anterior fibres of the M. pectoralis, so that by this means 
the two muscles, the M. interhyoideus posterior and the M. pecto- 
ralis are to some extent associated. The M. interhyoideus posterior 
lies ventral to the MM. procoraco-humeralis and supracoracoideus, 
which arise from the pectoral girdle, and is loosely attached to them 
by connective tissue. 

This muscle is peculiarly an adult structure, and Druner was 
somewhat doubtful as to its derivation, but thought that it might 
have arisen from the M. interbranchialis i. Edgeworth, however. 



THE MUSCLES 59 

holds that it is derived from the posterior fibres of the M. inter- 
hyoideus which have shifted their origin in a similar manner to the 
main portion, and have also migrated backwards and obtained a 
secondary attachment to the skin of the gular fold. 

Innervation-. From R.jugularisVII, which bends over the posterior 
edge of the muscle at its origin and then enters it from the dorsal 
surface. 

Function : The muscle must obviously act in more than one way. 
In so far as it acts as a constrictor of the pharynx — the anterior fibres 
in particular — it must be of importance in swallowing food and in 
breathing, while, by virtue of their attachment to the skin and to the 
fascia pectoralis^ the simultaneous contraction of the posterior fibres 
on both sides would tend to depress the head, and the contraction of 
one side at a time would cause the head to incline sideways. 

The three muscles just described are all superficial, and are seen 
directly the skin is removed. In order to expose the remainder of 
the hyobranchial muscles it is necessary to cut along the linea alba, 
and turn back the MM. intermandibularis and interhyoidei. 

M. subhyoideus (Driiner, in Ellipsoglossa) (m.s.hy.). 

Cerato-glossi extern! ...... Funk (1827). 

Os hyoides protrahens (14) ..... Carus (1828). 

Geniohyoideus anticus et posticus . . . v. Siebold (1828). 

Genio-hyoideus lateralis ..... Druner (in Sal. 1901). 

This is the last of the Vllth nerve muscles. It arises from the 
posterior end of the cerato-hyal^ which it encloses in a sort of muscular 
cup. It is narrow and ribbon-like in form, and its fibres run approxi- 
mately parallel with and ventral to the cartilage from which it arises. 
The muscle widens a little at its insertion, which is on the dorsal side 
of the aponeurosis of the M. intermandibularis. 

The homology of this muscle is in some doubt. Druner regarded 
it as the anterior portion of the larval interhyoideus, and its position 
and innervation are certainly in agreement with this view. On the 
other hand, the work of Miss Louise Smith (1920) on the develop- 
ment of the hyobranchial muscles of Spelerpes seem to be against 
it. Miss Smith describes muscle which she calls 'genio-hyoideus 
lateralis' which appears to be homologous with the muscle just 
described. Nevertheless, in Spelerpes — according to Miss Smith — 
it is proliferated not from the anlage of the interhyoideus but 
from that of the genio-hyoideus. Now the M. genio-hyoideus 
is a muscle belonging to the hypoglossal (first spinal) nerve, and 
all muscles derived from the same anlage should be innervated 



6o THE SALAMANDER 

by the same nerve, whereas in Salamandra the M. subhyoldeus is 
innervated by the seventh cranial nerve. To add to the confusion, 
Elizabeth Szamoylenko (1904) in describing the hyobranchial 
muscles of the adult Spelerpes states that the M. genio-hyoideus 
lateralis is innervated by the N. glossopharyngeus in both Spelerpes 
and Salamandra — which is wrong for the latter animal at least — 
with the addition in the former of fibres from NN. spinales i, 2, 
and 3. 

In the face of this conflicting evidence it is not possible to give 
a decisive verdict without a complete re-investigation of the develop- 
ment of the muscles concerned in both Salamandra and Spelerpes — 
an undertaking beyond the scope of the present work. In the mean- 
time a non-committal name (which has been used by Druner for what 
appears to be the homologous muscle in Ellipsoglossa) has been 
adopted, and it may be suggested that, in Salamandra, the muscle 
is most probably a true interhyoideus, and is not homologous with the 
muscle occupying a similar topographical position in Spelerpes. 

Innervation'. From R. jugularis VII by means of very fine fibres 
which enter the muscle on its ventral aspect after penetrating the 
M. interhyoideus. 

Function: The contraction of the muscle pulls the dorsal end of the 
cerato-hyal antero-ventrally, thus causing the cartilage to tip, and so 
force the anterior end thereof, with its overlying tongue, to move 
antero-dorsally. It is thus the elevator of the tongue. 

5. Muscles supplied by the Ninth and Tenth Nerves. 

M. subarcualis rectus i (Edgeworth) (m.s.r. i). 

Cerato-glossi-interni ...... Funk (1827). 

Alter (M. OS hyoides protrahens) (15) . . . Cams (1828). 

Ceratoglossus v. Siebold (1828). 

Pre-stylo-prebranchial Duges (1834). 

Cerato-hyoideus intemus Fischer ( 1 843) ; Mivart ( 1 869) ; Hoffmann 

(1873-8); Walter (1887); Druner (1901). 

Protracteur de la corne hyoVdienne posterieure . . Rusconi (1854). 

Cerato-hyoideus Stannius (1854-6). 

Cerato-hyoideus externus Osawa (1902). 

This muscle is very similar in appearance to the M. subhyoideus, 
since it bears the same relation to the cerato-branchial i as does that 
muscle to the cerato-hyal. It arises from the dorsal side of the 
posterior end of the first cerato-branchial cartilage by means of a 
pinnate raphe, from which the fibres radiate around the end of the 
cartilage so as to enclose it in a muscular cup. This cup is more 



THE MUSCLES 6i 

complete, and larger, than that formed at the origin of the M. sub- 
hyoideus. The fibres run more or less parallel with the cartilage on 
its ventral side, and are inserted on the ventro-anterior border of the 
expanded anterior end of the cerato-hyal. It thus forms a muscular 
link connecting the opposite ends of the two cartilages. 

Innervation: By several fine twigs from the N. glossopharyngeus 
(IXth) which passes along its dorso-mesial border, and also from 
a nerve formed by the fusion of twigs from the recurrent branches 
of the nerves of the visceral arches (Xth). 

Function : The obvious result of the contraction of this muscle is 
to advance the branchial arches, and their associated copula, relative 
to the cerato-hyal, and, since the movement of this latter element in 
a posterior direction is limited by the lig. hyoquadratum, the muscle 
is able to have a positive effect on the branchial apparatus relative 
to the jaws as well. It also serves to transmit to the branchial arches 
the movement occasioned by the M. subhyoideus. 

Driiner (1901, p. 526) gives an admirable account of the action 
of these two muscles in extruding the tongue to secure food, based 
on experiments carried out by him on pithed animals, consisting of 
applying an electrical stimulus to the nerves concerned, and also on 
observations made on the living animal. The substance of his account 
is given below. 

As already explained, the action of the M. subhyoideus is to ad- 
vance the cerato-hyal, first in an antero-dorsal direction, and later 
to tip the anterior end of the cartilage more dorsalwards. Now if the 
jaw is simultaneously opened wide, as it is in snapping at prey, the 
effect is to cause the dorsal surface of the tongue to face directly 
forwards, the action of the M. subhyoideus alone being about suffi- 
cient to bring the surface of the tongue to the level of the snout. The 
cerato-hyal being held in this position by the M. subhyoideus, its 
anterior end becomes Tipunctumfixum for the M. subarcualis rectus i, 
which, by its rapid contraction, forces the branchial arches and the 
copula, together with the tongue, still farther forwards. The muscle 
also pulls the posterior end of the cerato-branchial i ventralwards, 
causing the arches to twist about the copula and thus to force it still 
farther forwards. The combined result of these operations is to 
project the tongue to a surprising distance in front of the mouth so 
as to secure the prey on its sticky dorsal surface. The tongue, with 
its adhering food, is returned to the mouth by the action of muscles 
subsequently to be described — MM. hyoglossus and rectus cervicis. 
When the mouth has been shut the MM. subhyoideus and sub- 
arcualis rectus I again come into play so as to force the tongue 



62 THE SALAMANDER 

against the pre-vomerine teeth, and thus assist in working the food 

backwards to the pharynx so that it can be swallowed. 

The whole operation is performed with lightning-like rapidity, 
but the action is easily observed in the living animal by putting it 
in a glass receptacle, and allowing a small worm or slug to crawl 
down the side. In snapping at it the Salamander not infrequently 
gets its jaws momentarily wedged open against the glass with the 
tongue protruded, so that its action may be seen. 

It will have been noticed that the Salamander uses a very different 
method in securing its food from that of the Frog. In the latter 
animal the tongue is freely mobile, and it is this organ alone which 
is extruded, the skeleton of the hyoid taking but little part in the 
operation. In the Salamander, on the other hand, the tongue is 
securely fixed to the floor of the mouth, so that its movement carries 
with it the whole mouth floor and the hyobranchial skeleton as well, 

M. cephalo-dorso-subpharyngeus (mihi)^ (m.c.ph.). 

Dorso-trachealis .... Fischer (1843); Osawa (1902). 

Constrictor pharyngis Mivart (1869). 

Cephalo-dorso-pharyngeus ..... Druner (1901). 

This is a compound muscle formed from the M. transversalis ven- 
tralis iv and the MM. levatores arcuum branchiarum iii and iv of the 
larva (PI. XXIII, fig. 77). 

It arises dorsally by two heads, the anterior arising — together with 
the anterior part of the M. cucuUaris — from the crista muscularis^ a 
ridge formed by the squamosal above the ear capsule. The posterior 
head arises from t\\c fascia dorsalis together with the posterior portion 
of the M. cucullaris and the M. depressor mandibulae, and lies deep 
to the latter muscle, between it and the M. cucullaris. The direction 
of the anterior fibres is obliquely postero-ventral, while the posterior 
ones pass ventralwards practically at right angles to their origin, so 
that the whole muscle forms roughly a right-angled triangle of which 
the origin is the base and the anterior edge the hypotenuse. The 
ultimate insertion of the whole muscle is a very strong fibrous tendon, 
firmly attached in the middle line dorsally to the pharynx, just in 
front of the larynx, and ventrally to the truncus arteriosus. A short 
distance on either side of this median tendon is another tendinous 
inscription dividing the muscle into two portions, namely, a -pars 
subpharyngea (pars ventralis, Druner) and a pars dorsalis (Druner), 
the former lying mesial and the latter lateral to the inscription. A 
comparison with the larva shows that the pars subpharyngea is the 
' This name has been adopted in consultation with Dr. F. H. Edgeworth. 



THE MUSCLES 63 

transformed larval M. transversus ventralis iv, while the pars dor- 
salis represents the larval MM.levatores arcuum branchiarum iiiand 
iv, and the inscription itself takes the place of the atrophied cerato- 
branchial cartilages iii and iv. The relation of the two parts of the 
pars dorsalis is also interesting. They are almost always separated 
by the 'third' aortic arch, or its vestigial remains, so that the portion 
of the muscle representing the M, lev. arc. branch, iii of the larva 
passes between the systemic arch and the 'third', while that repre- 
senting the M. lev. arc. branch, iv passes between the 'third' arch 
and the pulmonary. The only exception so far recorded is one case 
reported by Driiner in which the whole muscle passed posterior to 
the 'third' arch, between it and the pulmonary. 

Innervation: Pars subpharyngea from the R. recurrens n. intesti- 
nalis X, Pars dorsalis^ anterior portion from the nerve of the fifth 
visceral (third branchial) arch, and th.^ posterior portion from the R. 
intestino-accessorius X. These nerves are exceedingly fine and enter 
the muscle from its mesial aspect. They are best seen from a dorso- 
lateral view, by separating the muscle from its origin and carefully 
turning it out laterally, when the nerves may be seen running across 
the space against the darker blood-vessels beneath. 

Function : One obvious function of the muscle is to constrict the 
pharynx, and thus it is probably of use in deglutition and respiration ; 
but, as Driiner rightly points out, the intimate connexion between 
the insertion of the muscle and the truncus arteriosus may cause it 
to have some effect on that organ. Again, the distal part of the 
'third' arch and the ductus Botalli between the pulmonary and sys- 
temic arches both pass mesial to the pars dorsalis^ so that the contrac- 
tion of the muscle would tend to compress these vessels against the 
wall of the throat and may thus have some influence in separating 
the arterial from the pulmonary blood. It is not possible to estimate 
its exact function, if any, in this respect without physiological experi- 
ments on the living animal. 

M. dilatator laryngis (Edgeworth) (m.d.l.). 

Dilatator aditus laryngis . ..... Henle (1839}. 

Dorso-laryngeus et Dorso-trachealis . . . Fischer (1843). 

Dorso-laryngeus et Dorso-trachealis, s. Dorso-branchialis v. 

Wilder (1892-6). 
Dorso-pharyngeus of Dorso-laryngeus et Dorso-trachealis 

Goppert (1894-8). 
Dorso-laryngeus .... Driiner (1901); Osawa (1902). 

This muscle is narrow and ribbon-like and arises from the dorsal 



64 THE SALAMANDER 

fascia lateral to the M. cucullaris and immediately posterior to the 
muscle just described. It passes ventralwards round the pharynx 
mesial to the thymus gland, and posterior to the pulmonary arch. 
It is inserted by means of a strong thread-like tendon — the ligamen- 
tum dorso-laryngeum (Driiner) — on \}plQ. processus muscularis at the side of 
the cartilago lateralis of the laryngeal skeleton. The muscle narrows 
considerably at its insertion and becomes somewhat spindle-shaped. 

The muscle has every appearance of being a serial homologue of 
the M. cephalo-dorso-subpharyngeus, that is to say, it looks 2iS though 
it were descended from a levator arcuum branchiarum posterior to 
the M. lev. arc. branch, iv, and indeed several notable authors have 
attempted to prove it to be such a muscle. Edgeworth (i 920), how- 
ever, has shown definitely that this is not the case, and that the re- 
semblance is apparent rather than real. A more detailed discussion 
of the various theories concerning the homology of the larynx is 
given on p. 272. 

Innervation: From the N. intestino-accessorius X by fine twigs 
which enter the mesial aspect of the muscle, and also by a branch from 
the R. recurrens intestinalis X which enters the muscle ventrally. 

Function: As its name implies it dilates the larynx and opens the 
glottis. 

M. constrictor laryngis (Edgeworth) (m.c.l.). 

Constrictor aditus laryngis Henle (1839); Fischer (1843); Druner 

(1901). 
Ring of Periarytenoideus dorsalis et ventralis . Wilder (1892-6) 

Sphincter laryngis ...... Goppert (1894-8). 

A small but powerful muscle which surrounds the larynx immediately 
ventral to the glottis. The shape of the whole muscle is much like 
a napkin ring. The fibres do not pass right round the circle, how- 
ever, but are interrupted in the middle line, both dorsally and ven- 
trally, by strong tendinous inscriptions. Most of the muscle fibres 
arise from, and are inserted into, these inscriptions, but a few of the 
inner ones have both their origin and insertion confined to the/>^rj 
laryngea of the cartilago lateralis which the muscle surrounds. The 
normal direction of the fibres is somewhat oblique so that their ven- 
tral ends are anterior to their dorsal ends, but not infrequently a 
small bundle may be found which reverses this order, the ventral 
ends of its fibres being posterior to their dorsal ends, so that the 
fibres of this bundle cross those of the main mass. Driiner calls them 
Fihrae cruciatae. The M. constrictor laryngis is always relatively a 
more powerful muscle in the adult than in the larva, and is normally 



THE MUSCLES 65 

the only one of the MM. laryngei of the larva which persists to the 
adult stage. The M. laryngeus ventralis may, however, be repre- 
sented by a few fibres passing from the lateral end of the lig. dorso- 
laryngeum to the ventral inscription of the M. constrictor laryngis 
(Driiner, 1901, p. 532). 

Innervation: From the R. recurrens n. intestinalis X. 

Function: It constricts the larynx, closes the glottis, and is the 
antagonist of the M. dilatator laryngis. 

6. Muscles of the Head supplied by Spinal Nerves (PI. VI, figs. 
37 and 38). 

M. genio-hyoideus (m.g.hy.), Carus (1828); Fischer (1843); Mivart 
(1869); Osawa (1902). 

Rectus lingualis ....... Funk (1827}. 

Levator maxillae inferioris s. Geniothyroideus . v. Siebold (1828). 

Genio-sous-hyoidien ...... Duges (1834). 

Genio-hyoidien .... Cuvier (1835); Rusconi (1854). 

Genio-hyoidei-mandibularis .... Stannius (1854-6). 

Levator maxillae inferioris longus Schmidt, Goddard, and van d, 

Hoeven (1864). 

Genio-branchial ...... Humphry (1872). 

Maxillo-hyoideus (genio-hyoideus) . . Hoffmann (1873-8). 

Genio-hyoideus s. rectus superficialis hypobranchialis anterior 

Driiner (1901). 

After removing the skin from the floor of the mouth and throat, and 
turning back the MM. intermandibularis and interhyoideus a pair 
of strong longitudinal muscles are exposed — they are the MM. 
genio-hyoidei. They may be regarded as arising from the inner edge 
of the lower jaw immediately lateral to the origin of the M. genio- 
glossus, and then, passing backwards and mesialwards, the fibres 
from either side become very closely approximated in the middle 
line as they continue their course backwards. The right and left 
halves of the muscle may remain distinct, but there is often some 
crossing over and mingling of the fibres. In the larva the muscle is 
inserted on the posterior end of the uro-branchial cartilage, and in 
the adult the bulk of the muscle remains attached to the vestige of 
this cartilage, the os triangulares but some fibres spread to a tendinous 
inscription which extends in a transverse plane from the lateral 
extremities of this bone. In addition a small bunch of the most 
lateral fibres are inserted on the capsule of the thyroid gland. There 
is, however, some considerable variation in the exact relations of the 
muscle at its insertion. 

4038 F 



66 THE SALAMANDER 

Innervation : The hypoglossal nerve traverses the muscle on its way 
to the tongue, and, in doing so, supplies it with numerous fine twigs. 

Function: The muscle may act in several different ways. It may 
depress the lower jaw, or the whole head, or it may pull forwards the 
OS triangulare with its attached truncus arteriosus and heart according 
to which other muscles are acting in conjunction, or in opposition. 

M. genio-hyoideus tertius (Druner). 

Driiner (1901), p. c^'t^'}^^ describes what he calls an abnormality of 
the M. genio-hyoideus, in which a few of the most lateral fibres of 
the muscle on the right side, instead of passing from the jaw to the 
OS triangulares pass from the jaw to the posterior end of the cerato- 
branchial i, and lateral to the M. subarcualis rectus i (ceratohyoideus 
internus of Druner), which in this case was weaker than the corre- 
sponding muscle on the left. He illustrates the muscle in Fig. 24 and 
there labels it geniohyoideus tertius. The innervation is by fine fibres 
from the hypoglossal nerve. 

Another case very similar to Driiner's has been observed in which 
a muscle, apparently rather smaller than Driiner's, passed from the 
posterior cornu of the hyoid to the anterior part of the lower jaw. It 
was also innervated from the N. hypoglossus. 

It is interesting in this connexion to refer to the discussion given 
on p. 59 regarding the homology of the M. subhyoideus, since it 
seems not unreasonable to suggest that the abnormal muscle just 
described may represent the muscle occurring normally in Spelerpes, 
and described by Miss Louise Smith as the M. genio-hyoideus 
lateralis. 

M. genio-glossus (m.g.gl.) Siebold (1828); Fischer (1843); Mivart 
(1869); Druner (1901). 

Genio-hyoidei . . , . . . . Funk (1827). 

A fairly strong and somewhat extensive muscle in the Salamander, 
having a fleshy origin from the mandibular symphysis. After re- 
moving the M. intermandibularis the genio-glossus muscle may be 
seen as two triangular blocks filling the angle of the 'chin', but it is 
necessary to remove both the M. genio-hyoideus and the cerato-hyal 
cartilages before the whole muscle is revealed. It is then seen to 
consist of two portions : (i) a mesial portion — which is the genio- 
glossus in the true sense — consisting of more or less parallel fibres 
passing on either side of the middle line to become inserted in the 
root of the tongue and in a tough ligament connecting the tongue 
with the copula — the aponeurosis lingualis (Druner) — and with the 



THE MUSCLES 67 

insertion of the M. rectus cervicis profundus; (ii) a second portion 
in which the fibres spread out, more or less fan-shaped, over the 
floor of the mouth, into which they are inserted at the sides of the 
tongue. 

Innervation : By the terminal twigs of the N. hypoglossus. 

Function: The contraction of this muscle must necessarily produce 
several results according to circumstances. For example, if the 
tongue were at rest then the muscle would tend to draw it towards 
the front of the mouth and to arch the dorsal surface, but if the tongue 
were fully extruded then the contraction of the muscle would tend 
to retract it towards the mouth. Probably its main function, how- 
ever, is to compress the glandular tissue of the tongue. This it would 
do by drawing the tongue towards the front of the mouth under the 
circumstances first mentioned, and the muscle would thus act as 
a preliminary for the extrusion of the tongue by causing it to be 
coated with a sticky mucus to which the prey may adhere. 

M. hyoglossus (m.hy.gl.) Fischer (1843); Druner (1901). 
In Salamandra this is a small and comparatively unimportant muscle. 
Its fibres pass from the dorsal side of the anterior extremity of the 
copula in a posterior direction, on either side of the cartilage. It lies 
dorsal to the anterior radials (hypohyals) and some fibres also pass 
dorsal to the posterior radials to be inserted in the tongue, but the 
more ventral fibres are attached to these cartilages. Driiner distin- 
guishes the ventral fibres as the M. basiradialis. The whole muscle 
lies deep to the M. rectus cervicis profundus, which must therefore 
be removed to expose it. According to Driiner the muscle does not 
exist in the larva. 

Innervation : By terminal twigs from the N. hypoglossus. 

Function: Beyond attaching the hyobranchial apparatus securely 
to the tongue, and possibly assisting to compress the glands of the 
latter organ, it is difficult to see what useful purpose the muscle can 
serve. The basi-radial portion would tend to draw the posterior 
radials antero-dorsally, and thus cause the tongue to stand up from 
the floor of the mouth as well as to compress the mucus glands. 

M. rectus cervicis (Edgeworth). 

Pubio-hyoidien s. Pubio-glosse (partim) . , , Cuvier (1800). 

Hyoideo-ypsiloideus ...... Funk (1827). 

Sternohyoidien .... Duges (1834); Rusconi (1854). 

Sternohyoideus Fischer (1843); Owen(i866); Mivart(i869); Hum- 

phry (1872); Furbringer (1873); Walter (1887); 
Osawa (1902}. 



68 THE SALAMANDER 

Levator maxillae inferioris brevis Schmidt, Goddard, and van d. 

Hoeven (1864). 
Thoracico-hyoideus .... Hoffmann (1873-8). 

Retracteur de I'hyoide ...... Perrin (1892). 

This muscle represents the direct forward continuation of the M. 
rectus abdominis, as Cuvier's name indicates, and, like that muscle, 
it is divisible into two strata, one superficial and the other deep. 
These two layers will be dealt with separately. 
M. rectus cervicis superficialis (Edgeworth) (m.r.c.'). 

Rectus V. Siebold (1828). 

Sterno-hyoideus superficialis .... Fiirbringer (1873). 

Rectus superficialis hypobranchialis s. Sterno-hyoideus Druner (1901). 
With M. rectus abdominis superficialis = 

Abdominis musculus rectus (8) .... Carus (1828). 

The superficial stratum of the M. rectus cervicis may be regarded 
as arising from the sternum and from the transverse inscription con- 
tiguous to it. It passes anteriorly, dorsal to the coracoids and ventral 
to the pericardium, as a broad thin sheet of muscle. It has at least 
three insertions: (i) at the angle formed by the hypobranchial i 
with the copula; (ii) on the tendon of insertion of the profundus 
portion of the muscle by a small deep bundle. The above-mentioned 
insertions apply only to the more lateral portions of the muscle; the 
mesial fibres are attached (iii) to the os triangulare and to the in- 
scription from which the M. genio-hyoideus arises. Between the 
sternum and the hyobranchial skeleton there are two transverse ten- 
dinous inscriptions, and at the anterior of these the M. pectori- 
scapularis is inserted. It is very closely attached to the pericardium 
(see also p. 270). 

Innervation : The muscle is supplied by twigs from each of the 
first three spinal nerves. 

Function: The mesial fibres support the M. genio-hyoideus, while the 
lateral fibres assist the rectus cervicis profundus in retracting the 
tongue. 

M. rectus cervicis profundus (Edgeworth) (m.r.c"). 

Hebosteoglossus ...... v. Siebold (1828). 

Sterno-hyoideus profundus .... Fiirbringer (1873). 

Rectus hypobranchialis profundus s. Abdomino-hyoideus Druner (1901). 
With M. rectus abdominis profundus = 

M. epischio-hyoideus (7) ..... Carus (1828). 

As already indicated, this muscle forms the direct forward continua- 
tion of the M. rectus abdominis profundus, so that it is a moot point 
to speak of its 'origin'. By analogy with the superficial portion it is 



THE MUSCLES 69 

perhaps best to regard it as arising at about the level of the sternum. 
The muscle passes forwards around the lateral parietes of the peri- 
cardium, mesial to the thyroid gland, and between the hypobranchial 
cartilages i and 2, passing ventral to the latter and dorsal to the 
former. It is inserted on the dorsal side of the apex of the copula by 
means of a strong tendon. The section of the muscle alongside the 
pericardium passes through a sort of 'sleeve' of connective tissue, 
so that its contraction does not affect the surrounding structures. 
Within this 'sleeve', a short distance posterior to the os triangulare, 
a small bundle of fibres separates from the main mass and loops 
round mesialwards so to become attached to the antero-dorsal aspect 
of the OS triangulare. This bundle was seen by v. Siebold as early as 
1828, and was named by him M. hebosteoypsiloideus^ while it may 
represent the muscle referred to by Carus as the oblique epischio- 
hyoideum (9). 

Innervation : As in the superficial portion ; see above. 

Function : To retract the tongue or to depress the head, according 
to circumstances. 

M. pectori-scapularis (Edgeworth) (m.p.s.). 

Schulterzungenbeinmuskel (omohyoideus) . . . Meckel (1828). 

Omo-hyoideus v. Siebold (1828); Rusconi (1854); Mivart (1869}; 

Humphry (1872); Hoffmann (1873-8); Osawa 

(1902). 

Scapulo-post-hyoidien ...... Duges (1834). 

Pectori-scapularis internus .... Fiirbringer (1873). 

Pectori-scapularis s. omo-hyoideus . Walter (1887); Druner (1901). 
A small, relatively weak, spatulate muscle arising from the mesial 
aspect of the ventral end of the scapula and inserted, as mentioned 
above, at the lateral edge of the superficial portion of the M. rectus 
cervicis at the level of the anterior inscriptio tendinea. 

Innervation: By a fine twig arising from the R. communicans 
between the first and second spinal nerves. 

Function: The muscle acts as a brace to the M. rectus cervicis 
superficialis. 

MUSCLES OF THE FORE-LIMB AND SHOULDER 
I. Historical. 

The earliest author to make a special study of this region of the 
body was Riidinger (1868), whose classic account of the comparative 
anatomy of the muscles of the neck, shoulder, and fore-limb in Am- 
phibia, Reptiles, Birds, and Mammals is quite well known. This 



70 THE SALAMANDER 

account was followed some five years later by the more extensive 
and even better known work of Fiirbringer (1873). Although 
Fiirbringer's work is more extensive as regards the range of types 
studied, it is more limited than Riidinger's in respect of the area 
investigated, since Fiirbringer does not deal with the muscles of the 
fore-arm and hand. Nevertheless his work shows a considerable 
advance over that of previous authors in that he deals with the nerves 
as well as the muscles, and it represents the real starting-point of all 
subsequent investigations. Both his text and his figures are reproduced 
by Hoffmann (1873-8). In 1895 Eisler published a long treatise on 
the homology of the muscles of the tetrapod extremities. His work is 
not of great importance so far as the Salamander itself is concerned, 
but is fundamental with regard to the wider question of muscle mor- 
phology. The next worker of note is Perrin (189 3-9), who has made 
a detailed study of the muscles of both fore- and hind-limbs in the 
Amphibia, both Anura and Urodela, Salamandra being taken as 
the main type of the latter. He makes a useful comparison between 
the muscles of the anterior and posterior limbs, but his work suffers 
from the rather serious defect that he has not studied the nerves in 
conjunction with the muscles, and hence his groupings are some- 
times a little unfortunate. The work of McMurrich ( 1 903), although 
not of specific importance so far as Salamandra is concerned,- is 
nevertheless relevant from a phylogenetic point of view. His con- 
clusions are discussed in the text. A comparative account of the 
extensor muscles and their nerves in both fore- and hind-limbs of 
Urodeles was given by Sieglbaur in 1 904, but no mention is made 
of the condition on the flexor side of the limbs. The muscles and 
nerves of the distal portions of the limbs of Amphibia, Reptiles, and 
Mammals have been adequately dealt with by Ribbing (1907—9). 
His account is clear and critical, and both his terminology and 
grouping have been adopted here. More recently (1924) Hellen 
Rylkoff has studied the development of the shoulder-muscles of 
Salamandra and finds that they are divisible into two main groups, 
namely, the primary shoulder muscles which develop/ro;^ the humerus 
towards the shoulder-girdle, and the secondary shoulder muscles which 
develop centrijugally towards the limb. The former group is again 
divisible into a dorsal and a ventral group. The dorsal primary 
group comprises the M. anconaeus, the M. dorso-humeralis,^ the 
M. dorsalis scapulae, and the dorsal part of the M. procoraco-humer- 
alis, while the ventral primary group consists of the M. humero- 

' The names here given are those used in this work; RylkofF's synonyms may be 
found by consulting the lists preceding the description of each muscle. 



THE MUSCLES 71 

antibrachialis inferior, the M. pectoralis, the MM. coraco-brachialis 
longus and brevis, the M. supra-coracoideus, and the ventral part of 
the M. procoraco-humeralis. The dorsal and ventral groups may- 
be further subdivided by a vertical plane into anterior and posterior 
elements. The secondary muscles of the shoulder-girdle are the 
MM. thoraci-scapularis, developing from the trunk myotomes 2—5, 
the M. opercularis, which develops from the trunk myotomes 1—2, 
and the M. cucullaris which arises from the visceral myotomes. 

2. Shoulder-muscles (PI. VII, fig. 43). 

M. opercularis (m.o.). Gaupp (1898); Kingsbury and Reed (1909). 
Levator anguli scapulae . . . Funk (1827); Rudinger (1868) 

Mivart (1869). 
Levator scapulae Carus (1828); Schmidt, Goddard, and van d 

Hoeven (1864); Humphry (1871); Osawa (1902) 
Rylkoff (1924). 
Sous-occipito-adscapulaire (32). .... Duges (1834) 

Fasciculus of protractor scapulae .... Ow^en (1866) 
Basi-scapularis (levator scapulae) . . . Fiirbringer (1873) 

Basi-scapularis (levator scapularis) . . , Hoffmann (1873-8) 

Protracteur du scapulum (98) ..... Perrin (1899) 
Levator scapuli ....... Driiner (1901) 

This peculiar muscle has a fleshy origin from the cartilaginous oper- 
culum of the ear capsule. It passes directly backwards alongside the 
dorsal trunk-muscles of the neck region, mesial to the M. cucullaris 
and to the ninth and tenth cranial nerves, and is inserted along the 
anterior border of the cartilaginous suprascapula. It is practically 
circular in section at its origin, but spreads out fan-shaped at its 
insertion. It is a derivative of the M. levator scapulae, and was first 
named M. opercularis by Gaupp. It seems better to retain Gaupp's 
name for the muscle in the Salamander owing to its peculiar origin 
and function. 

Innervation : By one or two fine twigs from the first spinal nerve — 
N. hypoglossus. 

Function : The function of this muscle is more fully discussed in the 
section dealing with the auditory organ (p. 295), but, in addition to 
acting as a track for auditory impulses, a tension on the muscle would 
tend to 'damp out' the effect which would be produced by the inertia 
of the operculum at every sudden movement of the head. For these 
functions alone the muscle seems surprisingly powerful, but in view 
of the lack of any direct experimental evidence it is unsafe to specu- 
late further. 



72 THE SALAMANDER 

M. cucullaris (Edgeworth) (m.c). Rylkoff (1924). 

Omo-mastoideus Funk (1827). 

Vorwartszieher der Schulter Meckel (1828). 

Levator scapulae (/)tfr/z;w) (20} .... Cams (1828). 

Spini-sus-scapulaire, portion du trapeze, masto-sus-acromial ou sterno- 
mastoidien, ex-occipito-sus-scapulaire (28, 29, 30, 31) Duges (1834). 
Fasciculus of Protractor scapulae .... Owen (1866). 
Cucullaris et Sterno-cleido-mastoideus . . Rudinger (1868). 

Trapezius . . Mivart (1869); Humphry (1872); Druner (1901); 

Osawa (1902). 
Capiti-dorso-scapularis s. Cucullaris Furbringer (1873); Hoffmann 

(1873-8). 
Petroso-dorso-scapularis (96-7} .... Perrm (1899). 

A thin triangular sheet of muscle which arises by two heads. The 
posterior, or cucullaris minor^ arises from the, fascia cephalodorsalis^ 
and is consequently firmly attached to the skin. Its fibres run directly 
ventralwards and it passes mesial to the anterior portion towards its 
insertion, which is on the lateral border of the pro-coracoid and 
scapula. The fibres are nearly parallel, converging slightly towards 
their insertion. The anterior portion, or cucullaris major ^ arises partly 
from the fascia cephalodorsalis^ but chiefly from the postero-dorsal 
surface of the skull and from a raphe which occurs between this 
portion of the muscle and the M. depressor mandibulae. It has a 
fleshy insertion on the lateral face of the pro-coracoid near the 
shoulder-joint, while a few of the posterior fibres are attached to the 
anterior edge of the ventral end of the scapula. The cucullaris major 
is triangular in shape, the anterior fibres being very oblique, while 
the posterior fibres are nearly vertical. The antero-ventral edge of 
the muscle is considerably thicker than the postero-dorsal. Although 
the M. cucullaris is essentially a superficial muscle it lies deep to the 
M. depressor mandibulae anteriorly, as well as to the MM. cephalo- 
dorso-subpharyngeus and dilatator laryngis, it being necessary to re- 
move, or turn back, these muscles in order to expose it fully. Its 
insertion is on the adjoining parts of the pro-coracoid and scapula 
between the M. dorsalis scapulae and the M. procoraco-humeralis. 
Carus (1828) labels a muscle M. scapulare-cutaneus which is appar- 
ently identical with the cucullaris minor. 

Innervation-. From N. accessorius X which enters its mesial sur- 
face. The cucullaris minor also receives a twig from the N. thora- 
cicus superior 2 (second spinal nerve). 

Function: The muscle must be mainly responsible for turning or 
depressing the head according to circumstances. If the head were 



THE MUSCLES 73 

held firm by the dorsal trunk muscles then it would protract the 
pectoral girdle. 

M. dorsalis scapulae (m.d.s.). Humphry (1872); Riidinger (1868); 
Fiirbringer (1873); Hoffmann (1873-8); Osawa (1902). 

Sur-epineux et Sous-epineux conjoints . . . Cuvier (1800). 

Scapularis ........ Funk (1827). 

Auswartszieher oder oberer ausserer Schulterblattmuskel Meckel (1828). 
(?) Externus (supra — et infra — spinatus) et internus scapulae (subscapu- 

laris) (22) Carus(i828). 

Adscapulo-humeral, sous-epineux .... Duges (1834). 
Suprascapularis . . . Stannius (1854-6); Rylkoff (1924). 

Subscapularis . Schmidt, Goddard, and van d. Hoeven (1864). 

Supra- and infra-spinatus, and perhaps also Deltoideus Owen (1866). 

Deltoid Mivart (1869). 

filevateur scapulaire du bras (90) .... Perrin (1899). 

This muscle is fan-shaped and arises from the dorso-lateral surface 
of the cartilaginous suprascapula, and is partly covered by the M. 
dorso-humeralis. Its fibres run ventralwards and converge to a very 
strong tendon which is inserted on the side of the crista ventralis 
humeri. This insertion lies between those of the M. procoraco- 
humeralis and the M. dorso-humeralis, but is entirely separate from 
both. 

Innervation: By the N. dorsalis scapulae from the third spinal 
nerve. 

Function: Its function is obviously to raise the arm in a vertical 
plane passing through the shoulder-joint. 

M. dorso-humeralis (m.d.h.). 

Grand dorsal ....... Cuvier (1800). 

Oberer Riickw^artszieher oder breiter Riickenmuskel . Meckel (1828). 
Latissimus dorsi . Funk (1827); Carus (1828); Stannius (1854-6); 

Schmidt, Goddard, and van d. Hoeven (1864); 

Owen (1866); Rudinger (1868); Mivart (1869); 

Humphry (1872); Osawa (1902); Rylkoff (1924). 
Vertebro-costo-humeral, grand dorsal . . . Duges (1834). 

M. dorso-humeralis (latissimus dorsi) Furbringer (1873); Hoffmann 

(1873-8). 
Elevateur dorsal du bras (89) ..... Perrin (1899). 

A large, flat, triangular muscle plate, situated behind the shoulder, 
which takes its origin from the dorsal fascia continuous with that 
from which the M. cucullaris and M. depressor mandibulae, &c., 



74 THE SALAMANDER 

arise — the fascia cephalodorsalis. The fibres are closely connected 
with the skin dorsally, and with the tendinous inscriptions of the 
body-muscles mesially. The origin of the muscle extends backwards 
over about three to four vertebrae, while its anterior border partially 
overlies the M. dors. scap. The fibres converge very strongly, 
the anterior ones being nearly vertical and the posterior ones very 
oblique. At the shoulder-joint they pass into a very strong, flat 
tendon which has two insertions. The first is at the postero-lateral 
edge of the shoulder-joint, where it merges into the tendon of origin 
of caput a (scapularis medialis) of the M. anconaeus. The second 
insertion is on the antero-lateral border of the humerus, just behind 
that of the M. dors. scap. The tendon of this latter insertion is very 
long and strong, and passes mesial to the M. dors, scap., forming 
a sort of tendinous bridge over the articulation. 

Innervation: By the N. dorso-humeralis which emerges from an 
anastomosis of the third and fourth spinal nerves. It enters the 
anterior border of the muscle. 

Function: It is the retractor of the humerus, and opposes the M. 
procoraco-humeralis. 

The three muscles just described are all dorsal shoulder-muscles 
and are lateral to the scapula. The one now to be considered also lies 
dorsal to the shoulder-joint but mesial to the scapula. 

M. thoraci-scapularis [serratus magnus). Fiirbringer (1873); Hoff- 
mann (1873-8). 

Depressor anguli scapulae inferioris .... Funk (1827) 
Serratus anticus magnus (21} ..... Carus (1828) 
Grosser Riickwartszieher der Schulter . . . Meckel (1828) 

Costo-sous-scapulai re, ou grand dentele . . . Duges (1834) 

Levator scapulae et serratus anticus major . . Owen (1866} 

Serratus anticus magnus . .... Riidinger (1868) 

Serratus magnus . . . Mivart (1869); Humphry (1872). 

Retracteur du scapulum (99, ioo)+Suspenseur scapulaire (loi) 

Perrin (1899), 
Thoraci-superiores-serrati ..... Rylkoff(i924) 

This muscle consists of a series of bundles arising from the first five 
ribs, which are inserted on the mesial face of the scapula. Furbringer 
divides it into two sections, an 'Obere Partie' and an 'Untere Partie', 
while Perrin distinguishes two distinct muscles (see above), the 
former of which he again subdivides into two sections. Perrin's 
grouping has been retained here, but it seems preferable to regard 
his 'M. retracteur du scapulum' and 'M. suspenseur scapulaire' as 



THE MUSCLES 75 

'oblique' and 'straight' portions respectively, of a common muscle, 
since the innervation is virtually the same in each case. 

The straight portion consists of two flat sheets of muscle arising 
from the lateral faces of the expanded ends of the first two ribs. The 
fibres pass directly dorsalwards and are inserted on the dorso-mesial 
border of the suprascapula. 

Innervation: From the NN. thoracici superiores of the second, 
third, and fourth spinal nerves. 

Function : The muscles serve to depress the scapula and thus cause 
the ventral portion of the pectoral girdle to expand. 

The oblique -portion again consists of two sections. The anterior 
of these arises from the lateral face of the second rib, just ventral to 
the straight portion. It passes anteriorly, inclined slightly dorsal- 
wards, and is inserted on the antero-mesial border of the supra- 
scapula. The second portion arises by three heads from the third, 
fourth, and fifth ribs. These unite into a single bundle passing antero- 
dorsally to be inserted on the postero-dorsal angle of the suprascapula. 
This portion of the muscle lies deep to the M. dorso-humeralis. 

Innervation: As above. 

Function : It serves to retract the scapula, while the general func- 
tion of the M. thoraci-scapularis as a whole is to attach the pectoral 
girdle to the axial skeleton, thus forming an elastic suspension for 
the anterior part of the body. 

The muscles associated with the ventral half of the pectoral girdle 
are all inserted in the limb, and are therefore considered in the 
sections which follow (PI. VI I, figs. 43 and 45). 

3. Muscles of the Arm.^ 

M. pectoralis (m.p.). Schmidt, Goddard, and van d. Hoeven (i 864) ; 
Owen (1866); Mivart (1869); Humphry (1872); Furbringer 
(1873); Hoffmann (1873-8); Rylkoff (1924). 

Grand pectoral ....... Cuvier (1800). 

Portio inferior m. pectoralis majoris . . . . Funk (1827). 

Grosser Brustmuskel Meckel (1828). 

Abdomino-coraco-humeral, portion du grand pectoral Duges (1834), 

Pectoralis major . . . Stannius (1854-6); Rudinger (1868). 

Pectoralis sternalis +pectoralis abdominalis . . Eisler (1895), 

Tete sternale; flechisseur du bras (86) . . . Perrin (1899). 

Together with MM. supracoracoideus and procoraco-humeralis = 

Pectoralis magnus + Pectoralis magnus, pars anterior . Carus (1828). 

' As in the case of the skeleton the proximal joint of the limb is referred to as the 
'arm', and the distal joint as the 'fore-arm'. 



76 THE SALAMANDER 

The M. pectoralis is a large, superficial, fan-shaped plate of muscle 
covering the posterior region of the breast. Its anterior fibres are 
approximately transverse to the vertebral axis and gradually become 
more and more oblique until eventually the posterior ones are almost 
longitudinal. The muscle has a varied origin. Anteriorly the fibres 
lying ventral to the coracoid arise from a median aponeurosis separat- 
ing the muscles of the right and left sides, next a few fibres arise 
from the sternal plate, then follows a section which takes its origin 
from the wide aponeurosis of the M. obliquus externus superficialis, 
and finally the most posterior fibres of all are continuous with 
those of the M. rectus abdominis superficialis. All the fibres con- 
verge to a strong tendon, which is inserted, together with that of the 
M. supra-coracoideus, on the posterior aspect of the crista ventralis 
humeri. 

Innervation : By the N. pectoralis which arises from the fourth (or 
fourth and fifth) spinal nerve and enters the posterior portion of the 
muscle on its mesial aspect. 

Function: To draw the arm inwards towards the body and pos- 
teriorly, or if the hand is resting on the ground, to advance the body. 

MM. supracoracoideus and coraco-radialisproprius. Furbringer (1873); 
Hoffmann (1873-8); Osawa(i902). 

These two muscle slips are very difficult to separate, and in fact their 
separation must be regarded as largely artificial, while they are most 
clearly distinguished by their insertions. 

(a) M. supracoracoideus (m.s.c). Osawa (1902); Rylkoff (1924). 

Portio media m. pectoralis majoris .... Funk (1827). 

Einwartszieher, Theil des grossen Brustmuskels . Meckel (1828). 

Clavi-humeral, portion du grand pectoral . . . Duges (1834). 

Pectoralis secundus ..... Stannius (1854-6). 

Pectoralis minor . Schmidt, Goddard, and van d. Hoeven (1864). 

Part of pectoralis ....... Owen (1866). 

Coraco-brachialis proprius .... Riidinger (1868). 

First part of coraco-brachialis ..... Mivart (1869). 

Epicoraco-humeral. ..... Humphry (1872). 

The more superficial of the two slips arises from the ventral surface 
of the cartilaginous portion of the coracoid, with the exception of the 
posterior third. Its fibres converge to a flat tendon, which is inserted, 
together with that of the M. pectoralis, on the posterior face of the 
crista ventralis humeri. The posterior half of the muscle is covered 
by the M. pectoralis. 



THE MUSCLES 77 

(J?) M. coraco-radialis proprius, Stannius (1854-6). 

Biceps brachii Rudinger (1868). 

Part of biceps Mivart (1869). 

Coraco-radialis or biceps .... Humphry (1872). 

Long flechisseur de I'avant-bras (85) . . . Perrin (1899). 

Coraco-radialis ....... Osawa (1902). 

This section of the muscle also arises from the ventral surface of the 
coracoid, deep to the foregoing slip. It is a very thin, fan-like muscle 
and converges, at the glenoid cavity, to a strong two-lipped tendon. 
The shorter lip is inserted close to that of the former slip on the 
flexor side of the head of the humerus, while the longer tendon 
(m.c.r'.) passes right down the arm, lying parallel with the humerus, 
and between the M. coraco-brachialis longus and the M. humero- 
antibrachialis. It is inserted on the mesial aspect of the proximal end 
of the radius. 

Innervation : Both slips are innervated by the N. supracoracoideus, 
from the second and third spinal nerves. 

Function : Both muscles serve to draw the arm towards the body^ 
while the tendon of the latter portion also assists in flexing the elbow. 
In general it is the opposer of the M. dors. scap. 

M. procoraco-humeraUs (m.p-h.). Fiirbringer (1873); Hoffmann 
(1873-8); Osawa (1902); Rylkoff (1924). 

Portio superior m. pectoralis majoris . . . Funk (1827). 

Vorwartszieher oder Heber des Oberarms, dreieckiger Muskel 

Meckel (1828). 

Acromio-humeral, Deltoide ..... Duges (1834). 

Deltoides .... Stannius (1854-6); Rudinger (1868). 

Subclavius Mivart (1869). 

Precoraco-brachial Humphry (1872). 

Adducteur inferieur du bras (92) .... Perrin (1899). 

This muscle arises from the pro-coracoid portion of the pectoral 
girdle, and passes posteriorly to be inserted on the head of the 
humerus. The tendon of insertion is attached to the humerus a little 
antero-mesial to the insertion of the M. dorsalis scapulae, while a 
few fibres may be inserted between this muscle and the anterior 
insertion of the M. dorso-humeralis. The shape of the muscle is 
that of a narrow fan. 

Innervation-. The muscle has a dual innervation. At its lateral 
border it receives a branch from the N. dorsalis scapulae, and at its 
mesial border it is supplied by the N. supracoracoideus. 

Function : If the foot is raised from the ground the contraction of 



78 THE SALAMANDER 

the muscle tends to draw the Hmb forwards and at the same time to 
turn the humerus so as to depress the pre-axial side of the limb. If, 
however, the foot is resting on the ground this tendency to twist the 
humerus would assist in propelling the body forwards. 

MM. coraco-hrachialis longus et brevis. Humphry (1872); Furbringer 
(1873); Hoffmann (1873-8); Osawa (1902); Rylkoff (1924). 

Hakenarmmuskeln Meckel (1828). 

Coraco-humeral ....... Duges (1834). 

Coraco-brachialis . Stannius (1854-6); Rudinger (1868); Schmidt, 
Goddard, and van d. Hoeven (1864). 

Second part of coraco-brachialis .... Mivart (1869). 

Coraco-brachialis longus, superficialis, and brevis . Eisler (1895). 

(supracoracoideus +coraco-brachialis brevis) = 

Tete coracoidienne; flechisseur du bras (87) . . Perrin (1899). 

(coraco-brachialis longus) = 

Deducteur du bras (88) ...... Perrin (1899). 

Tht caput longum (m.Q.hA.) IS a muscle of medium power covering the 
posterior face of the humerus. It arises from the postero-lateral 
border of the coracoid a little towards its dorsal side. Its fibres run 
parallel with one another and with the humerus, and have a fleshy 
insertion along the posterior surface of the distal half of this bone. 
The caput breve is a short but powerful triangular muscle, separ- 
ated from the former head by the N. brachialis. It arises from the 
posterior third of the ventral surface of the coracoid, i.e. posterior 
to the M. supracoracoideus, and also from the postero-lateral edge — 
lateral to the head just described — and encroaches a little on the 
dorsal and mesial sides of the coracoid. It is inserted on the posterior 
face of the proximal half of the humerus, and immediately proximal 
to the caput longum. 

Innervation: By the NN. coraco-brachiales from the R. super- 
ficialis, N. brachialis. 

Function : Both heads serve to draw the arm caudalwards, or con- 
versely, if the foot is resting on the ground to propel the body 
forwards. 

M. subscapularis. Funk (1827); Stannius (1854—6); Rudinger 
(1868); Mivart (1869); Osawa (1902). 

Sous-scapulo-humeral, sous-scapulaire . . . Duges (1834). 

Coraco-brachialis quartus (subscapularis) . . Humphry (1872). 

Subcoracoscapularis . Furbringer (1873); Hoffmann (1873-8); 
Rylkoff (1924). 

Subscapularis (dorsal portion) Subcoracoscapularis . Eisler (1895). 

Adducteur superieur du bras (93 and 94) . . . Perrin (1899). 



THE MUSCLES 79 

A muscle of medium strength arising from the dorsal surface of the 
pro-coracoid. The fibres pass caudalwards to the posterior margin 
of the coracoid where they turn sharply round lateralwards, and are 
inserted on the crista dorsalis humeri. The M. anconaeus coracoideus, 
and the MM. coraco-brachialis longus et brevis lie on the ventral 
side, and the M. anconaeus scapularis medius is on the dorsal side 
of this muscle. 

Innervation : By the N. subscapulars from the third spinal nerve. 

Function : The contraction of the muscle draws the arm backwards 
and thus, if the foot is resting on the ground, it serves to advance the 
body. 

M. anconaeus. Furbringer (1873); Hoffmann (1873-8); Rylkoff 
(1924); Osawa (1902). 

Anconaeus internus et externus .... Funk (1827) 

Triceps brachii extensor (24) ..... Carus (1828). 

Strecker des Vorderarms . . . . . Meckel (1828) 

Scapulo-humero-olecranien, triceps .... Duges (1834) 

Streckmuskelmasse ..... Stannius (1854-6) 

Triceps brachii s. anconaeus . Riidinger (1868); Mivart (1869) 

Triceps and coraco-olecranalis . . . Humphry (1872) 

Extenseur de I'avant-bras (81-3) .... Perrln (1899) 

The M. anconaeus arises from the shoulder-girdle and humerus by 

four heads which unite distally, to a greater or lesser extent, to form 

a powerful muscle which is inserted on the olecranon process of the 

ulna. In general it is innervated by the extensor nerves, 2ind functions 

as extensor of the fore-arm. The details of its several components 

may be considered separately. 

Caput (a). M. anconaeus scapularis medialis (m.a.s.m.). Furbringer 
(1873); Hoffmann (1873-8); Rylkoff (1924). 

Anconaeus longus ....... Eisler (1895). 

Tete anterieur (81) ...... Perrln (1899). 

Anconaeus caput scapulare ..... Osawa (1902). 

This head arises from the scapula at the posterior margin of the 
glenoid cavity. The main tendon arises a little towards the mesia/ side 
of the scapula, but retains a strong fibrous connexion with the whole 
posterior region of the capsule surrounding the joint. The muscle is 
thin and flat, and is separated from the M. anconaeus coracoideus 
by the entry of the extensor nerves and the V. brachialis medialis into 
the arm, as well as by the insertion of the M. subscapularis. 

Innervation: By twigs from theN.extensorius caudalis(cf. p. 166). 
Function: See above. 



8o THE SALAMANDER 

Caput (b). M. anconaeus coracoideus (m.ac.c). Fiirbringer (1873); 
Hoffman (1873-8); Eisler (1895); Rylkoff (1924). 

Coraco-olecranonalis . . . . . Humphry (1872). 

Tete posterieur (82) ...... Perrin (1899). 

Anconaeus caput CO racoideum . .... Osawa (1902). 

This portion of the anconaeus muscle arises by a long, tough tendon 
from the mesial surface of the bony portion of the coracoid. It is 
relatively small and slight, and joins the former muscle at about the 
middle of the humerus. As already mentioned it is separated from 
the head just described by the extensor nerves and by the insertion 
of the M. subscapularis. 

Innervation : As caput {a). See above. 

Function : See above. 

Caput {c). M. anconaeus humeralis lateralis (m.a.h.l.). Fiirbringer 
(1873); Hoffmann (1873-8); Rylkoff (1924). 

Anconaeus caput humerale mediale . . . Osawa (1902). 

With caput ^/ = Tete profonde (83) . . . Perrin (1899). 

Situated on the dorso-lateral surface of the arm the M. anconaeus 
humeralis lateralis is considerably larger than the two portions of the 
anconaeus muscle already described. It arises from the proximal 
third of the antero-lateral surface of the humerus, just posterior to 
the insertions of the MM. dorso-humeralis and dorsalis scapulae. 
The direction of its fibres is at first approximately parallel with the 
humerus, but distally they become oblique in an antero-posterior 
direction, and the whole muscle passes over to join the other heads 
on the extensor surface of the arm. It is separated from the head 
next to be described by the extensor nerves. 

Innervation : By twigs from both extensor nerves. 

Function: See above. 

Caput (d). M. anconaeus humeralis medialis (m.a.h.m.). Fiirbringer 
(1873); Hoffmann (1873-8); Rylkoff (1924). 

Anconaeus caput humerale laterale . . . Osawa (1902). 

With caput c = Tete profonde (83) . . . Perrin (1899). 

The fourth head of the anconaeus takes its origin from the whole 
extensor surface of the humerus. It lies mesial to the scapularis and 
coracoideus portions of the muscle and joins them distally near the 
common insertion. 

Innervation: From the N. extensorius cranialis. 

Function : See general account of whole muscle. 



THE MUSCLES 
M. humero-antibrachialis (m.h.a.). Rylkoff (1924). 
Brachialis medius ...... 

Flexor brachii (biceps cum brachial! interno) (23) 
Oberer Beuger des Vorderarms 
Humero-radial s. biceps . 
Humero-radialis 



Biceps . 

Biceps and brachialis 
Brachialis internus 
Part of biceps 
Brachialis anticus . 



Funk (1827). 

Carus(i828). 

Meckel (1828). 

• Duges(i834). 

Stannius (1854-6); Osawa (1902). 



Schmidt, Goddard, and van d. Hoeven (1864). 
internus ..... Owen (1866). 

Rudinger(i868}. 

Mivart (1869). 

. Humphry (1872); Eisler (1895) 
Humero-antibrachialis inferior (Brachialis inferior) 

Fiirbringer (1873); Hoffmann (1873-8). 
Court flechisseur de I'avant bras (84) . . . Perrin (1899). 

This muscle arises from the flexor side of the humerus, distal to the 
crista ventralis humeri^ and runs parallel with the bone, converging 
slightly towards its distal end. Its insertion on the proximal end of 
the radius is inclined to become tendinous. 

Innervation-. It is supplied by one or two branches from the R. 
superficialis N. brachialis. 

Function-. It is the true flexor muscle of the elbow. 

4. Extensor Muscles of the Fore-arm and Hand (PI. VII, figs. 43 
and 44). 
These should be dissected from the dorsal side. 

M. extensor digitorum communis (m.e.d.). Rymer Jones (1852); 
Schmidt, Goddard, and van d. Hoeven (1864); Riidinger (1868); 
Ribbing (1907). 



Extensor digitorum (30) . 
Gemeinschaftlicher Strecker 
Epicondylo-digital . 
Extensor longus 
Extensor digitorum sublimis 
Humero-digiti II-V dorsalis 
Extensor digitorum communis longi 
Extenseur de la main (63) 
Extensor digitorum longus 
Humero-metacarpalis 
This is the most superficial muscle on 



. Carus(i828). 

. Meckel (1828). 

. Duges(i834). 

. Mivart (1869). 

Humphry (1872); Rylkoff (1924). 

Hoffmann (1873-8). 

Eisler (1895). 

Perrin (1899). 

Osaw^a (1902). 

Sieglbaur (1904). 

the extensor surface of the 



fore-arm. It arises by means of a thin, broad tendon from the epicon- 
dylus lateralis humeri., superficial to the MM. extensor antibrachii et 
carpi ulnaris and radialis, and spreads out as a flat plate of muscle 
covering the extensor surface of the fore-arm and wrist. Distally it 

4038 G 



82 THE SALAMANDER 

separates into six tendons, which are inserted on either side of the 
bases of the metacarpals of each of the three ulnar fingers. Occasion- 
ally a seventh tendon may be present which is then inserted on the 
post-axial side of the base of the first metacarpal. 

Innervation : From the N. dorsalis manus ulnaris, by twigs passing 
between the extensor muscles of the ulnar and radial sides of the 
fore-arm. 

Function : As its name implies it is the common extensor of the 
hand. 

M. extensor antihrachii et carpi radialis (m.e.a.c.r.). Eisler (1895); 
Ribbing (1907). 

Extensor carpi radialis (28) Carus (1828). 

Gemeinschaftlicher Heber Meckel (1828). 

Epicondylo-carpien; Sus-epicondylo- radial . . Duges (1834}. 

Extensor carpi radialis . Rymer Jones (1852); Schmidt, Goddard, and 

van d. Hoeven (1864); Osawa (1902). 
Abductor ....... Stannius (1854-6). 

Supinator longus and brevis .... Rudinger (1868). 

Supinator Mivart (1869). 

Radial section of the superficial stratum representing the Supinator longus 

and brevis, and the Extensor carpi radialis . Humphry (1872). 

Humero-radialis dorsalis . .... Hoffmann (1873-8). 

filevateur carpien interne (66); filevateur du radius (67) Perrin (1899). 

A very powerful muscle, arising from almost the whole surface of the 
epicondylus lateralis humeri and from the strong raphe which occurs 
between its proximal end and that of the M. ext. ant. et carp, ulnaris. 
It has a fleshy insertion along the whole length of the lateral surface 
of the radius with the exception of about a millimetre at the proximal 
end (= M. ext. ant. rad.), and on the radiale (= M. ext. carp. rad.). 

Innervation : Mainly from the N. extensorius cranialis with a few 
fibres from the N. extensorius caudalis. 

Function: In conjunction with the M. ext. ant. et carp, ulnaris and 
the M. ext. dig. comm. it would act as an extensor muscle. Alone it 
would tend to turn the hand into the supine position. 

M. extensor antihrachii et carpi ulnaris (m.e.a.c.u.). Eisler (1895); 
Ribbing (1907). 

Extensor carpi ulnaris . Carus(i828); Rymer Jones(i852);Schmidt, 
Goddard, and van d. Hoeven (1864); 
Humphry (1872); Osawa (1902). 

Eigener Heber Meckel (1828). 

Epicondylo-cubital ...... Duges (1834). 



THE MUSCLES 83 

Extensor carpi ...... Stannius (1854-6). 

Extensor carpi s. metacarpi radialis und ulnaris . Riidinger (1868). 

Humero-ulnaris dorsalis . . Hoffmann (1873-8); Rylkoff (1924). 

filevateur cubito-carpien (64-5) .... Perrin (1899). 

Ulnaris externus ...... Sieglbaur (1904). 

This muscle forms the ulnar counterpart of the foregoing. It arises 
from the extensor border of the epicondylus lat. hum., and from the 
raphe mentioned above as occurring between this and the previous 
muscle. It is inserted along the central portion of the lateral face of 
the ulna (= M. ext. antibr. ulnaris) and on the ulnare+intermedium 
(= M. ext. carp, ulnaris). 

Innervation: From the N. extensorius caudalis. 

Function : It is the true extensor of the fore-arm. 

All three of the foregoing muscles are closely associated at their 
origin. The separation of the two latter muscles into 'antibrachii' and 
'carpi' portions is to be regarded as in an incipient state in the Sala- 
mander, and varies in different individuals. They are usually separ- 
able distally, since the 'antibrachii' portions have fleshy insertions on 
the radius or ulna, while the 'carpi' portions are gathered into strong 
tendons before their insertions. 

M. abductor and extensor digiti primi (m.a.e. i .). Sieglbaur (i 904). 



(?) Hallici proprius 
Cubito-sus-pollicien 
Extensores und Adductores 
Extensor digiti I 
Extensor brevis 



. Carus(i828). 

. Duges(i834). 

Stannius (1854-6). 

Rudinger(i868). 

Mivart (1869). 



Uppermost part of the deep stratum s. Supinator manus 

Humphry (1872). 
Ulnari-phalanx II dorsalis .... Hoffmann (1873-8). 
Abductor metacarpi II -|-extensor digiti II brevis superficialis 

Eisler (1895). 

Court adducteur du premier metacarpien (69)+Court extenseur du 

premier doigt (70 et 71) ..... Perrin (1899). 

Abductor digiti secundi-j-Extensor brevis digiti II . Osawa (1902). 

This muscle group arises mainly from the combined ulnare+inter- 
medium^ while a few fibres (possibly representing the M. extensor 
brevis digiti I profundus ^) take their origin from the os centrale. It 
passes obliquely across the wrist and is inserted on the outer edge of 
the proximal end of the first metacarpal (= M. abductor digiti I), 
and on the distal end of the same bone (= M. extensor digiti I). 

Innervation-. From the N. dorsalis manus intermedius. 

Function: It is the extensor muscle of the first digit. It would 



84 THE SALAMANDER 

appear that again this muscle is to be regarded as a case of the inci- 
pient separation of two muscles primitively united. The 'abductor' 
portion probably assists in supination (cf. Ribbing). 

MM. extensores breves digitorum (m.e.b.). Ribbing (1907). 

Extensor brevis Cams (1828). 

Extensores et adductores .... Stannius (1854-6). 

Extensor digitorum brevis s. profundus . . Humphry (1872). 

Carpo-phalangei ...... Hoffmann (1873-8). 

Extensor brevis superficialis digitorum +Extensor brevis profundus 
digitorum ........ Eisler (1895). 

Courts extenseurs des doigts (72-7) .... Perrin (1899). 

Extensores digitorum breves s. profundus III-IV . Osawa (1902). 

Extensores digitorum II, III, et IV. . . Sieglbaur (1904). 

These small muscles arise in two layers. The more superficial and 
thicker stratum takes its origin from the ulnareAr intermedium and the 
OS centrale (for the second digit), while the deeper and thinner layer 
arises from the basal carpalia, each muscle springing from the basal 
of the corresponding digit. The two strata of the muscle are separated 
by the branches of the A. perforans carpi. They have a tendency to 
unite distally into long thin tendons which pass along the extensor 
surfaces of the fingers and are inserted on the proximal end of each 
phalanx. 

Innervation: From the NN. dorsales manus inter, and ulnaris. 
Function : Extensors of the fingers. 

M. extensor lateralis digiti quarti (m.e.l.d.4.) (mihi). 

Court deducteur du quatrieme metacarpien (80) . Perrin (1899) 

First described by Perrin, this very small muscle arises from the distal 
external angle of the ulnare + intermedium and is inserted laterally 
on the proximal angle of the fourth metacarpal. Its fibres are accord- 
ingly very short. 

Innervation : From N. dorsalis manus ulnaris. 

Function-. To assist in extending the hand by drawing the fourth 
digit laterally. Opposer of the MM. intermetacarpals. 

5. Flexor muscles of the Fore-arm and Hand (PI. VII, figs. 45 

and 46). 

These should be dissected from the ventral side. They are more 
numerous than the extensor muscles. 

M. flexor primordialis communis (m.f.p.). Ribbing (1807). 

Flexor digitorum (29) ...... Carus (1828). 

Palmaris superficialis . , Eisler (1895); McMurrich (1903). 



THE MUSCLES 85 

Together with MM. flexores breves superficiales and flexores accessorii 
lateraHs and medialis = 

Flexor digitorum communis profundus et sublimis Funk (1827). 

Langer gemeinschaftlichcr Beuger .... Meckel (1828). 

Epitrochlo-digital ....... Duges (1834). 

Flexor digitorum communis Stannius (1854-6); Rymer Jones (1852); 
Schmidt, Goddard, and van d. Hoeven 
(1864); Riidinger (1868). 

Flexor longus Mivart (1869). 

Humero-phalangei volares II-V . . . Hoffmann (1873-8). 

Flechisseur commun des doigts (14-28) . . . Perrin (1899). 

Together with MM. flexores breves superficiales, and flexor accessorius 
lateralis = 

Flexor digitorum sublimis .... Humphry (1872). 

Together with MM. flexores accessorii lateralis et medialis = 

Flexor digitorum longus sublimis .... Osawa (1902). 

The flexor primordialis communis is a thin, flat sheet of muscle arising 
from the epicondylus medialis humeri between the MM. flexor anti- 
brachialis et carpi radialis and ulnaris and from aponeuroses which 
occur between it and these muscles. It is very difficult to separate 
proximally from the flexor carpi ulnaris; indeed, there is no natural 
division between them other than the common aponeurosis from 
which their fibres originate. In the region of the carpus the muscle 
passes into a thin, flat tendon which occupies the whole of the palm 
of the hand, and then divides into four narrow tendons which pass 
along the fingers and are inserted on the terminal phalanges, while 
an auxiliary tendon passes to the second phalanx of the third finger. 
The tendon is fixed to the skin at the carpus by tough fibrous con- 
nective tissue and is also related to various other muscles. See below. 

Innervation-. From N. ulnaris. 

Function : As its name indicates, it is the common flexor of the hand 
and fingers. 

To observe the following muscles the M. flex. prim. comm. should 
be severed at its origin and turned backwards over the hand, care 
being taken not to damage the short flexor muscles of the wrist which 
are inserted on the palmar tendon. 

M. flexor antihrachii et carpi radialis (m.f.a.c.r.). Eisler (1895); 
McMurrich (1903); Ribbing (1907). 

Pronator carpi radialis (?) ..... Funk (1827). 

Flexor carpi radialis (25) . . Carus (1828); Osawa (1902). 

Abzieher oder Beuger der Hand .... Meckel (1828). 



86 THE SALAMANDER 

Sous-epitrochlo-radial Duges (1834). 

Adductor Stannius (1854-6). 

Pronator teres Schmidt, Goddard, and van d. Hoeven (1864); Riidinger 

(1868); Mivart(i869). 
Radial sector of the superficial stratum representing the Pronator teres 
and the flexor carpi radialis . . . Humphry (1872). 

Humero-radialis volaris Hoffmann (1875-8). 

Abaisseur radio-carpien interne (39-40) . . . Perrin (1899). 

This muscle arises from the epicondylus medialis humeri together with 
the M. flex. prim, comm., from which it is very difficult to separate 
at the proximal end. It has a fleshy insertion all along the external 
face of the radius (= flexor antibrachii radialis), while the most ex- 
ternal fibres are attached to the radiale (= flexor carpi radialis). The 
separation of these two strata is not a natural one in Salamandra, 
since in this animal the muscle — although of medium strength — is 
relatively weak when compared with other Urodeles (cf. Ribbing). 

Innervation : From the N. ulnaris, and from the R. superficialis N. 
brachialis before it anastomoses with the R. profundus — for details 
see these nerves, p. 168. 

Function : It is a flexor of the fore-arm. 

The muscle of the ulnar side of the arm corresponding to the fore- 
going radial muscle is comparatively easily separated into its com- 
ponents, which may therefore be considered separately. 

M. flexor carpi ulnaris (m.f.a.c.u.). Ribbing (1907). 

Abductor digiti minimi . . Schmidt, Goddard, and van d. Hoeven 

(1864); Humphry (1872). 
Flexor digiti minimi proprius .... Riidinger (1868). 

Humero-metacarpus V volaris . . . Hoffmann (1873-8). 

Ulnaris internus ....... Eisler (1895). 

Abaisseur carpien externe (37) .... Perrin (1899). 

Together vv^ith the following muscle = 

Flexor carpi ulnaris . Carus(i828); Rymer Jones (1852); Humphry 
(1872); Osawa (1902). 

The carpal portion is the more superficial of the two strata. It arises 
partly from the epicondylus medialis humeri, but mainly from either 
side of a strong raphe, from which also some of the fibres of the M. 
flex. prim. comm. originate. It runs down the ulnar side of the fore- 
arm and, at the level of the wrist, converges into a strong tendon 
which is inserted on the lateral face of the ulnare+intermedium, just 
proximal to the origin of the M. flex, brevis profundus IV. 
Innervation : By N. ulnaris. 



THE MUSCLES 



87 



Function: In addition to depressing the foot it has a sHght flexor 
action on the fore-arm, especially when the limb is well extended. 

M. flexor antihrachit ulnaris (m.f.a.c.u.). Eisler (1895); Ribbing 
(1907). 

Epitrochlo-cubital ...... Duges (1834) 

Flexor carpi ...... Stannius (1854-6) 

Ulnaris ........ Mivart (1869) 

Humero-ulnaris volaris .... Hoffmann (1873-8) 

Abaisseur du cubitus (38) ..... Perrin (1899) 

Together with preceding muscle = 

Flexor carpi ulnaris . Carus(i828); Rymer Jones (1852); Humphry 
(1872); Osawa (1902). 

The second portion of the ulnar flexor muscle lies deep to the former. 
It is much smaller and forms a triangular block of muscle extending 
from its origin on the epkondylus medialis humeri to the proximal half 
of the ulna. It has a fleshy insertion on the outer edge of this bone. 

Innervation-. Like the former slip it is innervated by the N. ulnaris. 

Function : The contraction of the muscle draws the fore-arm back- 
wards — relative to the body — so that if the foot is resting on the 
ground it assists in advancing the body (cf. Perrin). 



The following three muscles are accessory to the M. flexor prim- 
ordialis communis. They are inserted on the dorsal side of its 
palmar tendon. 

M.. flexor accessorius lateralis (m.f.a.l.). Ribbing (1907). 

Palmaris profundus III . . Eisler (1895); McMurrich (1903). 

Caput ulnare m. flex. dig. long, sublimis . . . Osawa (1902). 

(For other homologies see M. flex. prim, comm.) 

The M. flexor accessorius is a small muscle arising from the distal 
end of the ulna, from the ulnare+intermedium, and from basale 4. 
It passes obliquely radialwards and is inserted, as already stated, on 
the dorsal side of the palmar tendon of the M. flex, prim. comm. 

Innervation: By N. ulnaris. 

Function'. It acts as an accessory flexor of the hand. 

M. flexor accessorius medialis (m.f.a.m.). Ribbing (1907). 

Palmaris profundus II . . Eisler (1895); McMurrich (1903). 

Caput ulno-carpale dig. long, sublimis . . . Osawa (1902). 

With M. pronator profundus = 

Pronator manus ...... Humphry (1872). 

(For other homologies see M. flex. prim, comm.) 



88 THE SALAMANDER 

This muscle arises from the mesial side of the distal third of the 
ulna, from the mesial side of the ulnare+intermedium, and from the 
ulnar carpal. It runs parallel with the former muscle and is separated 
from it by the M. caput longum musculorum contrahentium (see 
below). Its insertion is likewise on the palmar tendon, anterior to 
the M. flex. ace. lat., and extending a little more proximally than 
does that muscle. 

So far as Salamandra itself is concerned it seems a little pedantic 
to separate this muscle from the M. pronator profundus, and thus 
Humphry's term, Pronator manus, applied to the combined muscle, 
would appear to be the better term to use. Ribbing's arrangement 
has been adhered to, however, since his judgement is based on the 
study of an extensive series. It is impossible, nevertheless, to draw 
a sharp distinction between the two muscles. 

Innervation : From N. ulnaris, R. medialis. 

Function : As the former muscle. 

MM. flexores breves superficiales (m.f.b.s.). Eisler (1895); McMur- 
rich (1903); Ribbing (1907). 

Flexor breves ....... Carus (1828). 

Flexores digitorum sublimes breves .... Osawa (1902). 
(For other homologies see M. flex. prim, comm.) 
These are small weak muscles which are really auxiliary to the 
M. flexor prim, comm., and are grouped with it by most authors. 
They arise from the dorsal surface of its palmar tendon, and from 
the edges of its digital tendons. They are inserted on the edges of 
the metacarpals, at the bases of the proximal phalanges, and, in the 
third finger, on the above mentioned (p. 85) auxiliary tendon 
which passes to the second phalanx. 

M. caput longum musculorum contrahentium (m.c.l.c). Ribbing (i 907) 
Flexor metacarpi IV profundus longus . . . Eisler (1895) 

Flexor profundus longus ..... Osawa (1902) 

Ulno-carpalis McMurrich (1903) 

Together with MM. contrahentes digitorum = 

Kurzer beuger Meckel (1828) 

Flexor profundus digitorum . Rudinger (1868); Humphry (1872) 
Flexor brevis ....... Mivart (1869) 

Ulnari-phalangei volares II-V . . . Hoffmann (1873-8) 

Flechisseur accessoire des doigts (30 and 34) . . Perrin (1899) 

This is a long slip of muscle arising from the mesial border of the 
proximal end of the ulna. It is inserted, by means of a fan-shaped 
tendon, on the several carpalia. It lies deep to the M. flex. prim. 



THE MUSCLES 89 

comm., and its distal end separates the M. flex. ace. lat. from the M. 
flex. ace. med. Since the MM. contrahentes digitorum (see below) 
arise partly from the tendon of insertion of this muscle, as well as 
from the carpalia, Ribbing concludes that they originally formed a 
single muscle, and that the connexion with the carpalia is secondary. 

Innervation : From the R. medialis N. ulnaris. 

Function : It is the deep flexor muscle of the wrist. 

MM. contrahentes digitorum (m.c.d.). Ribbing (1907). 
Flexor breves medii s. contrahentes digitorum 

Eisler (1895); McMurrich (1903). 
Flexores digitorum profundi breves .... Osawa (1902). 
(See also M. cap. long. muse, contra, for other homologies.) 

These form a series of small flexor muscles which lie deep to the 
MM. flex, breves super., and superficial to the MM. flex. brev. prof. 
They arise partly from the tendon of insertion of the M. cap. long, 
muse, contra., and partly from the carpalia as follows : 
The 4th digit. From the tendon and from the fourth and third 

carpalia. 
The 3rd digit. From the tendon and mainly from the third carpal, 

but a few fibres arise from the second carpal also. 
The 2nd digit. From the tendon and from the combined first and 

second carpal. 
The ist digit. From the tendon and from the combined first and 

second carpal, not from the carpal of the prepollex (see also 

p. 42). 

All are inserted on the proximal ends of the corresponding proxi- 
mal phalanges. Ribbing's result in not being able to find any con- 
nexion between these muscles and the tendon of the M. flex. prim, 
comm., described by McMurrich, is confirmed. Ribbing regards 
the muscles as being derived from the M. cap long. muse, contra, 
(see above). 

Innervation : From the palmar branches of the NN. ulnaris and 
interosseus. 

Function : They serve both to flex the fingers and, to a less extent, 
to approximate them towards one another. 

M. pronator prof undus (m.p.p.). Ribbing (1907). 

Flexor indicis digit! proprius .... Riidingcr (1868). 

Ulnari-metacarpi volares II-III . . . Hoffmann (1873-8). 

Palmaris profundus I . Eisler (1895); McMurrich (1903). 

Rotateur direct de la main (61) .... Perrin (il 



90 THE SALAMANDER 

With M. flexor accessorius medialis = 

Pronator manus ...... Humphry (1872). 

With M. interosseus antebrachii = 

Pronator radii ....... Osawa (1902). 

The deep pronator muscle arises from nearly the whole of the mesial 
side of the ulna, from the ulnare+intermedium, from the centrale 
and the third (and sometimes the fourth) carpal. It converges 
towards the radial side of the hand, where it is Inserted on the 
radiale, on the carpal of the pre-pollex, on the combined first and 
second carpal, and, by means of a tendon, on the bases of the first 
and second metacarpals. 

Innervation : From the N. interosseus which passes dorsal to it. 

Function : To rotate the hand so as to bring the first digit nearer to 
the ground — i.e. pronation. 

M. interosseus antibrachii. Eisler (1895); Ribbing (1907). 
Cubito-digital . . . . . . . Duges (1834) 

Pronator quadratus . . Mivart (1869); McMurrich (1903) 

Pronator radii quadratus .... Humphry (1872) 

Radio-ulnaris ...... Hoffmann (1873-8) 

Interosseux de I'avant bras (62) .... Perrin (1899) 

Pronator ....... Sieglbaur (1904) 

With the M. pronator profundus = 

Pronator radii ....... Osawa (1902). 

This muscle is very closely associated with the foregoing and arises 
from almost the whole of the mesial surface of the ulna. It is inserted 
on the distal three-quarters of the mesial surface of the radial, the 
fibres passing obliquely across the interosseal space between the two 
bones of the fore-arm. It may be exposed from the ventral side by 
by removing the M. pronator profundus. 

Innervation: From the N. interosseus which crosses its dorsal 
surface. 

Function'. It forms an elastic ligament between the two bones of 
the fore-arm. 

MM. flexores breves prof undi (m.f.b.p.). Eisler (1895); McMurrich 
(1903); Ribbing (1907). 

Carpo-metacarpales Humphry (1872); Hoffmann (1873-8); Osawa 

(1902). 
Flechisseurs des metacarpiens (49-52) . . . Perrin (1899). 

These are four short, deep, flexor muscles which may be seen by 
removing the MM. contra, dig. They are separated from these 
muscles by the palmar branches of the flexor nerves, and arise 



THE MUSCLES 91 

from the distal angles of the carpalia, while the most posterior muscle 
arises also from the ulnare+intermedium. They are inserted on the 
respective metacarpalia. It is noteworthy that the M. flex. brev. 
prof, of the first digit arises from the combined first and second 
carpal and not from the carpal of the pre-pollex. It is a very slight, 
narrow muscle, much smaller than the others. 

Innervation : From the palmar branches of the NN. ulnaris and 
interosseus. 

Function : They are the flexors of the metacarpalia. 

MM. flexores digitorum minimi. Ribbing (1907). 

Metacarpo-phalangei . Humphry (1872); Hoffmann (1873-8). 

Flechisseurs primitifs des phalanges (45-8) . . Perrin (1899). 

Metacarpo-phalangeales ..... Osawa (1902). 

These are very slight, narrow muscles which arise from about the 
middle of the ventral side of the metacarpals of the second, third, and 
fourth fingers, and are inserted on the proximal ends of the proximal 
phalanges of the corresponding digits. There is no representative of 
this series of muscles in the first digit. 

Innervation : From the palmar branches of the NN. ulnaris and 
interosseus. 

Function : Flexors of the digits. 

M. interphalangeus digiti III (m.i.ph.3) (mihi). 

Phalangeus ....... Humphry (1872). 

Interphalangeus digiti IV Hoffmann (1873-8); Eisler(i895); Ribbing 

(1907)- 
Flechisseur primitif de la troisieme phalangine (43) . Perrin (1899). 
Interphalangeahs . . . Osawa (1902); McMurrich (1903). 

A very small, weak muscle arising from the ventral surface of the 
proximal phalanx of the third finger. It is inserted on the proximal 
end of the second phalanx of the same finger. 

Innervation : From the N. interosseus. 

Function : Flexor of the third digit. 

MM. intermetacarpaies (m.i.mc). McMurrich (1903). 

Interossei dorsales ..... Riidinger (1868). 

Interossei Humphry (1872); Hoffmann (1873-8); Eisler (1895); 

Osawa (1902); Ribbing (1907). 
Intermetacarpiens (53-5) ..... Perrin (1899), 

These are short triangular muscles lying between the digits con- 
necting each finger with its neighbour. They can be fully seen 
from the ventral side only after removing the MM. flex. brev. prof. 



92 THE SALAMANDER 

They are attached to the metacarpals, and, although it is difficult to 
distinguish between 'origin' and 'insertion', nevertheless, since they 
occupy rather less of the radial side of a given metacarpal, they may 
perhaps be correctly regarded as arising from this side, and as being 
inserted on the whole of the ulnar side of the metacarpal of the 
adjoining digit. 

Innervation: From NN. ulnaris and interossei. 

Function; They are the adductors of the fingers, and serve to 
draw the digits towards one another. 

MUSCLES OF THE TRUNK AND TAIL 

(PL XVI, %. 6z and PI. XIX, fig. 71) 

1. Historical. 

The muscles of the trunk and tail regions have not been the 
recipients of much attention ; in fact it is only necessary to cite two 
authors in order to cover the special literature relevant to the sub- 
ject. The first of these is Maurer (1892), who made a careful study 
of the ventral body muscles of Urodeles and compared them with 
the corresponding muscles of fish. His work is quite well known. 
The dorsal trunk muscles of Urodeles have been investigated by 
Nishi (19 1 6), who deals with Cryptobranchus and Necturus. The 
contributions of other workers are incidental rather than specific 
additions to our knowledge of these muscles. 

2. Dorsal Trunk Muscles. 

The dorsal body musculature remains very primitive and fish-like, 
and by most authors is treated generally as the dorsal muscle mass 
extending from the head to the tip of the tail. Nishi, however, dis- 
tinguishes three portions. 

M. dorsalis trunci (m.d.tr.). Nishi (19 16). 

Lateralis magnus (5) ...... Carus (1828). 

Gemeinschaftlicher Riickgrats- und Kopfstrecker . Meckel (1828). 

Dorsal muscle mass ...... Most authors. 

The muscle forms the bulk of the dorsal muscle mass. It is com- 
pletely segmented, being interrupted at each vertebra by a myo- 
septum which is attached to the neural spine and transverse processes 
of the vertebra. The myoseptum is not directly transverse but has 
a strong convexity caudalwards. Its edge is attached mesially to the 
strong connective tissue surrounding the median dorsal venom 
glands. Dorso-laterally it is attached to the skin. The dorsal edge of 



THE MUSCLES 93 

the septum, that is, the free edge after the skin and glands have been 
removed, follows a peculiar course (see fig. 78). Commencing 
from the neural spine, at the caudal end of the vertebra, it passes 
very obliquely anterodorsally, and slightly laterally, to the level of 
the articulation with the vertebra next in front, where it reaches 
a point on the dorsal surface of the muscle immediately lateral to 
the median dorsal venom glands. It now turns sharply backwards 
at a very acute angle and passes in a postero-lateral direction to some 
distance behind the transverse process of the vertebra to which it 
belongs, and then curves round again so as to lie directly along the rib. 
It is continuous with the inscriptio tendinea oithQ ventral musculature. 

The fibres of the M. dorsalis trunci run between the successive 
myosepta in an approximately sagittal direction. The whole muscle 
is very solid and powerful. In transverse section it is 'comma' shaped. 

A fairly well-defined differentiation of this muscle mass is dis- 
cernible on the dorsal side of the vertebrae, where a number of 
fibres arise from the postero-dorsal edge of the post-zygapophysis 
of one vertebra and are inserted along the dorsal surface of the neural 
arch of the vertebra next behind, thus filling the concavity which 
exists in this region on either side of the neural spine. This portion 
may be distinguished as the M. interspinalis (Nishi) (m.i.s.) (figs. 7 1 
and 78), while that portion of the M. dorsalis trunci which passes 
between the transverse processes of the vertebrae may be dis- 
tinguished by the name M. intertransversarius. This latter portion 
is distinguishable from the main mass only by the fact that it arises 
from, and is inserted on, the bone instead of the myoseptum. 

The other region where some differentiation occurs is at the head. 
The condition is not essentially different from that in the body 
region, as described above, but the several portions are more easily 
distinguishable. Arising from the dorsal side of the transverse pro- 
cesses of the second and third vertebrae a strong muscle mass spreads 
out laterally, and is inserted over the postero-lateral aspect of the ear 
capsule. It extends anteriorly to immediately behind the dorsal end of 
the squamosal, and lies lateral to the superficial portion of the M. lev. 
mandib. ant. It is the M. intertransversarius capitis superior (Gaupp, 
in the Frog), or M. longissimus capitis (Nishi). As in Necturus it is 
traversed by three myosepta. 

The M. rectus capitis posterior (^isKi)^ or M. occipitalis (Funk), is 
the mesial section of the spino-occipital musculature. It lies deep to 
the foregoing and to the M. lev. mandib. ant., and arises from the 
neural spine and neural arch of the first vertebra. It spreads over 
the dorsal surface of the occipital region of the skull on to which it 



94 THE SALAMANDER 

is inserted. It is quite distinct from the M. intertransversarius 

capitis superior. 

The M. intertransversarius capitis inferior (Gaupp, in the Frog) is 
really the direct continuation of the subvertebral musculature and 
arises from the transverse process of the second vertebra. It is 
inserted over the ventral surface of the occipital region of the skull, 
and is separated from the superior portion of the intertransversarius 
muscle by the first spinal nerve, and is necessarily antagonistic to it. 

Innervation: In general the dorsal trunk muscles are innervated 
from the dorsal rami of the spinal nerves. 

Function : Their general function is to produce a lateral flexion of 
the spine by alternate contraction on either side. The portions of the 
muscle attached to the skull similarly turn the head from side to side. 

3. Ventral Trunk Muscles. 

The muscles on the ventral side of the sulcus lateralis comprise two 
distinct series, the one — the rectus group — with fibres running 
parallel with the body axis, and the other — the oblique muscles — 
whose fibres are at an angle with this direction. 

It is not easy to give synonyms for these muscles, since there 
is some variation among the different genera of Urodeles in which 
they have been investigated. The reader should, therefore, refer to 
Maurer's paper for a detailed comparison, but the following may be 
quoted as applying to Salamandra. 

Funk (1827) calls the whole rectus muscle, including the M. 
rectus cervicis, the M. hyoideo-ypsiloideus. Carus (1828) dis- 
tinguishes the superficial from the deep portion and calls the former 
abdominis musculus rectus (8), and the latter M. epischio-hyoideus (7). 
He does not distinguish the 'cervical' from the 'abdominal' portions. 
Rymer Jones (1852) refers to the superficial portion as the Rectus 
abdominis and to the profundus portion as the M. pubo-hyoideus. 
He also does not separate the rectus cervicis and rectus abdominis 
muscles. Owen (1866) also calls the deep portion the M. pubo- 
hyoideus, while Hoffmann (1873-8) describes a muscle as the M. 
pubo-thoracicus (Rectus abdominis) which appears to represent the 
superficial portion of the muscle only, the profundus portion being 
undescribed. Hoffmann further distinguishes the M. ypsiloideus 
posterior as the M. pyramidalis. 

M. rectus abdominis superficialis (m.r.ab'.). Maurer (i 892). 

This portion of the rectus muscle forms a large flat sheet covering 

the ventral parietes of the abdomen, the right and left halves being 



THE MUSCLES 95 

separated by a wide median aponeurosis, or linea alba. The muscle 
may be regarded as extending from the anterior edge of the pubis 
to the level of the sternal cartilage, and is interrupted by ten tn- 
scriptiones tendineae — not counting the terminal inscription lateral 
to the sternum. It is, therefore, difficult to speak of 'origin' and 'in- 
sertion', since each inscription is firmly attached to the overlying 
skin. The direction of the fibres is longitudinal. The muscle is 
fairly easily separable into two portions, namely, a thicker mesial 
and a thinner lateral section, while the epigastric arteries run along 
the line of junction between them. Anteriorly the fibres take several 
courses, the most lateral merge into the M. pectoralis, the most 
mesial fibres pass dorsal to this muscle and are inserted on the 
postero-lateral edge of the sternal plate, and on the pericardium, 
while the remainder of the muscle continues directly forwards, 
dorsal to the coracoids, as the M. rectus cervicis superficialis (q.v.), 
and is inserted on the hyo-branchial apparatus. In addition to its 
attachment to the pubis a few of the posterior fibres are inserted on 
the lateral processes of the ypsiloid cartilage. 

M, rectus abdominis prof undus (m.r.ab."). Maurer (i 892). 
This muscle also extends from the pelvis to the hyoid, but unlike the 
superficial portion it lies more or less free throughout nearly its whole 
length. It arises from the anterior edge of the pubes deep to the 
superficial stratum, and is also interrupted by tendinous inscriptions 
of which only one or two of the most caudal are attached to those of 
the M. rect. abd. super., the remainder being entirely free. Anteriorly 
the muscle becomes rather more lateral in position and the inscrip- 
tions fail to correspond exactly with those of the other muscles, and 
also tend to become less distinct. This is doubtless to be explained 
by supposing that the freedom of the muscle has allowed a certain 
amount of stretching of the myomeres to occur. 

The portion of the muscle anterior to the sternum forms the M. 
rectus cervicis profundus (q.v.) 

Innervation: Both sections are innervated by the terminal 
branches of the spinal nerves 5-15 inclusive. 

Function : In general they serve to flex the vertebral column, while 
they also act on the hyo-branchial apparatus and tend to compress 
the viscera. 

M. ypsiloideus anterior. Whipple (1906). 

Consisting of only a few fibres which take their origin from the 

anterior edge of the lateral processes of the ypsiloid cartilage^ and 



96 THE SALAMANDER 

which have their insertion in the inscription next anterior, this 
muscle is scarcely to be distinguished from the remainder of the M. 
rectus abdominis superficialis, from which it is derived. 

Innervation: By the terminal branches of N. spinalis 14. 

Function: It is the elevator of the cartilago ypsiloidea. 

M, ypsiloideus posterior (m.y.p.). Whipple (i 906). 

M. pyramidalis .... Most other authors. 

This is a larger, more powerful, and better defined muscle, although 
its derivation from the rectus abdominis is still apparent. It arises 
from the antero-dorsal edge of the pubis deep to the M. rect. abd. 
prof, and spreads in a more or less fan-like manner to be inserted on 
the ypsiloid cartilage — on the lateral edges of the shaft, and the 
posterior edges of the lateral processes. The muscle is thicker at its 
lateral edges than mesially, and has considerable power. 

Innervation: By twigs from the combined nerve formed from 
NN. spinales 14 and 15 (.f* N. ileo-hypogastricus). 

Function : It is the depressor of the ypsiloid cartilage (see also p. 45). 

The M. ohliquus externus forms the most superficial layer of the 
lateral body-wall muscles. Like the rectus muscle it is divisible into 
superficial and deep strata. With regard to synonyms. Funk (1827) 
refers to the muscles of the lateral body-wall collectively as MM. 
intercostales, and while later authors dealing with the body muscles 
of Urodeles (mainly Cryptobranchus), e.g. Humphry, Mivart, 
Hoffmann, &c., make some distinction between the several layers, 
their views are not always identical, and there is also consider- 
able variation in the relative development of the various strata in 
different genera. It is therefore better that the reader should refer 
to Maurer's paper for a comparative discussion, than to attempt to 
correlate these authors' synonyms here. 

M. ohliquus externus superficialis (m.o.s.'). Maurer (i 892). 
The fibres of this portion arise dorsally from the ribs immediately 
ventral to the sulcus lateralis^ and also from the several inscriptiones 
tendineae which are continuous with their lateral extremities. The 
insertion is mainly on the ensuing inscription, but the most ventral 
fibres terminate along a line joining the shoulder-joint with the hip, 
and are inserted into a tendinous sheet which passes over the rectus 
muscles, ventrally, to the opposite side. This tendon, as well as the 
intermuscular inscription, is closely applied to the skin, and is very 
difficult to separate therefrom. The muscle extends from the rib of 



THE MUSCLES 97 

the third vertebra to the sacrum. The fibres run in an oblique 
direction from antero-dorsal to postero-ventral. 

M. ohliquus externus profundus. Maurer (1892). 
The deep stratum is an even thinner sheet of muscle than that just 
described. To expose this layer it is better to commence removing 
the superficial stratum from the ventral edge, as it is more easily 
separable there than at the dorsal edge. It is entirely similar to the 
superficial layer in origin and insertion, but the direction of its 
fibres tends to become more longitudinal. 

Innervation: Both parts are innervated from the ventral rami of 
NN. spinales 5-15. 

Function: In general both layers serve to contract the body wall 
and consequently to compress the body cavit)^, but the specific 
results of its action vary according to whether it is antagonistic or 
complementary to its neighbouring muscles. 

The most internal layer of the ventral trunk muscles is, according 
to Maurer, composed of three parts which, in higher forms, separate 
into distinct muscles, but which, in Salamandra, are not clearly 
separable one from another. 

The most dorsal and mesial section, lying next to the vertebral 
column, is the pars suhvertehralis (m.s.). It passes from vertebra to 
vertebra and is attached to the lateral part of each centrum and to the 
ventral face of each transverse process. Its lateral edge may be 
roughly defined as the point at which the spinal nerves emerge, but 
apart from the separation thus caused it merges quite imperceptibly 
into the pars transversalis. 

The pars transversalis is similarly attached to the ventral surfaces 
of the ribs, and, at their lateral extremities, merges in turn into the 
pars ohliqua interna. 

The pars ohliqua interna is attached to the myosepta which are 
continuous with the ribs. 

The direction of the fibres of the several portions varies. The 
subvertebral portion is longitudinal; the pars transversalis becomes 
rather more oblique, while the pars obliqua interna is very 
markedly so. The fibres of this latter portion pass from antero- 
ventral to postero-dorsal and cross those of the M. obliquus exter- 
nus almost at a right angle. The pars obliqua interna extends from 
the shoulder to the pelvis. The anterior fibres are inserted around 
the bases of the lungs and on the pericardium and help to form the 
diaphragm (see also p. 270). The other two portions are virtually 



98 THE SALAMANDER 

continuous from the head to the tip of the tail, although the sub- 
vertebral portion is somewhat deficient in the sacral region. The 
anterior fibres are inserted on the postero-ventral aspect of the 
occipital region of the skull. 

Innervation: From the ventral rami of the spinal nerves. 

Function-. The -pars suhvertehralis flexes the spine. The pars 
transversalis also tends to flex the spine if both sides contract simul- 
taneously, but if they contract alternately then a lateral flexion of the 
spine would occur. The function of the pars obliqua interna is 
similar to that of the obliqua externa (q.v.). 

4. Caudal Muscles. 

The caudal muscle mass is dorsally very similar to that of the body. 
The anterior part forms the M. ilio-caudalis (m.il.cd.), the fibres of 
which arise from the first two or three caudal vertebrae and are 
inserted on the ilium. The dorsal and ventral muscles of the tail are 
continuous laterally, forming strong C-shaped masses of muscle on 
either side of the vertebrae. In the mid-ventral line the muscles of 
the two sides are separated by a tough ligament which attaches the 
spines of the haemapophyses to the skin, while dorsally there is a 
deep median groove between the muscles wherein lie the cutaneous 
glands. 

The ventral muscle of the tail shows some slight tendency to 
differentiation in so far as some of the fibres pass from one haemapo- 
physis to the next instead of lying between myosepta, but the dis- 
tinction is not clear enough to justify their separation as an individual 
muscle. 

Innervation: From both dorsal and ventral rami of the spinal 
nerves. 

Function: They obviously serve as the motors of the tail. 

MUSCLES OF THE HIND-LIMB AND PELVIS 
(Pis. VIII and XVI) 

I. Historical. 

In general the hind-limb does not seem to have received so much 
attention as the fore-limb. One of the earliest workers to make a 
special study of its muscles is Perrin (1892). His work is very care- 
ful and detailed, but, as has already been noted in discussing his 
work on the arm muscles, it suffers from the absence of correlation 
with the nerves. Eisler's work (1895), already mentioned, covers 



THE MUSCLES 99 

the hind-limb as well as the arm, as also does that of Sieglbaur ( 1 904). 
McMurrich (1904) made a study of the phylogeny of the crural 
flexors and their nerves, and, while he does not deal expressly with 
Salamandra, his work is of sufficiently general nature to have a bear- 
ing on the problems involved. Most of his observations were made 
on serial sections, and this has caused him to draw one or two con- 
clusions which scarcely seem justified when viewed in the light of 
actual dissections. These conclusions are criticized by Ribbing 
(1909) and will be referred to again later. Ribbing's work is very 
sound and has the advantage that the results are correlated with those 
obtained for the fore-limb. More recently a phylogenetic study of 
the muscles and nerves of the post-axial region of the tetrapod 
thigh has been undertaken by Appleton (1928), who describes the 
condition found in the Salamander, while Low (1926—9) has studied 
the development of the pelvic girdle and its muscles in a number of 
Urodeles (but not Salamandra) and throws some light on the 
phylogeny of certain muscles. 

2. Muscles of the Pelvis and Thigh (PI. VIII, figs. 48 and 49). 

On removing the skin from the ventral surface of the pelvis and 
the base of the tail the following three muscles may be observed. 
They are grouped together by Appleton as the Caput ventrale^ M. 
flexor cruris. 

M. pubo-ischio-tibialis (m.p.i.t.). de Man (1873); Osawa (1902); 
Low (1926). 

Rectus ........ Funk (1827) 



Gracilis cruris flexor (35) ..... Carus (1828) 

Zweiter Beuger Meckel (1828) 

Sous-ischio-tibial ....... Duges (1834) 

Flexores abducentes ..... Stannius (1854-6) 

Depressor and Adductor of the pelvic limb . . Owen (1866) 

Gracilis Mivart (1869) 

Superficial stratum of Plantar muscles of Thigh (Gracilis +Semitendino- 
sus+Semimembranosus) .... Humphry (1872) 

Pubo-ischio-tibialis (Semitendinosus) . . Hoffmann (1873-8) 

Flechisseur du tibia (i lo-i i) ..... Perrin (1892) 
Rectus femoris ....... Eisler (1895) 

This is the most superficial muscle of the pelvis as seen from the 
ventral side. It arises just lateral to the pubo-ischiadic symphysis 
and covers the M. pubo-ischio-femoralis externus with the exception 
of a narrow strip anteriorly. It passes right down the ventral surface 
of the thigh and is inserted on the tibia. It is wide and thin at its 



loo THE SALAMANDER 

origin and converges rapidly while passing under the pelvic girdle to 
form a narrower, thicker muscle which tapers only slightly while 
passing down the thigh. Anteriorly it has a tough fibrous connexion 
with the M. pubo-tibialis, while at its posterior border it receives 
the M. caudali-pubo-ischio-tibialis which is inserted into it by 
means of a tendon at about the level of the proximal end of the femur. 
This is an excellent example of the rare phenomenon of one muscle 
being inserted into another, the fibres of the two muscles being 
approximately at right angles. 

Innervation: From the N. ischiadicus ventralis which passes 
between this muscle and the M. pub. isch. fem. ext. The fibres 
enter the muscle from the dorsal side. 

Function: It either serves to flex the knee, or, if the extensor 
muscles are antagonistic, the flexion would occur at the hip and it 
would depress the foot towards the ground. 

M. pubo-tibialis (m.p.t.). de Man (1873); Hoffmann (1873-8). 

Vastus Funk (1827). 



Carus(i82»j. 
. Meckel (1828). 
• Duges(i834}. 
Stannius (1854-6). 
. Owen (1866). 
Mivart (1869); Eisler (1895). 



Anticus cruris protractor (34) 

Grosster Beuger 

Pubo-sous-tibial 

Gracilis .... 

Protractor or flexor of the thigh 

Sartorius .... 

Gracilis part of the superficial stratum of the plantar muscles 

Humphry (1872). 
Adducteur du tibia (106-8) ..... Perrin (1892). 

This muscle has a tendinous origin from the antero-ventral edge of 
the pelvic girdle, i.e. from the pubic cartilage, in front of the obtura- 
tor foramen. It passes along the antero-ventral border of the thigh 
and is inserted on the anterior face of the proximal end of the tibia, 
close to the M. pub. isch. tib. It is a narrow muscle, more or less 
oval in cross-section, and of medium strength. 

Innervation : From N. ischiadicus ventralis. 

Function: It serves mainly to adduct the limb, i.e. to draw it 
towards the body, but it also has a slight flexor action on the knee- 
joint. 

M. ischio-flexorius (m.is.f.). de Man (1873); Eisler (1895); Osawa 
(1902); Low (1926). 

Plantae pedis (38) Carus (1828). 

Langer, starker Beuger ...... Meckel (1828). 

Flexor of the leg ...... Rymer Jones (1852). 



THE MUSCLES 



Flexores abducentes 



Stannius (1854-6). 



Mivart (1869). 

Humphry (1872). 

Hoffmann (1873-8). 

Perrin (1892). 



Semitendinosus .... 

Caudo-pedalis .... 

Ischio-flexorius (Semimembranosus) . 

Flechisseur externe de la jambe (115) 
Proximal portion = 

Ischio-flexorius ...... McMurrich (1904). 

Distal portion = 

Plantaris superficialis medialis .... McMurrich (1904). 

A Strap-like muscle, which arises from the posterior ventro-lateral 
angle of the os ischium, immediately behind the M. pub. isch. tib. 
During its course along the ventro-posterior border of the thigh it 
passes dorsal to the insertion of the M. caud. pub. isch. tib. into the 
M. pub. isch. tib. At the popliteal region it becomes more mesial 
and passes to about the middle of the flexor surface of the leg, and, 
at a point about midway between the knee and the ankle, it dis- 
appears into a tendon which mingles with the plantar aponeurosis of 
the M. flexor primordialis communis. 

This muscle had always been regarded as a single muscle until 
1904, when McMurrich, during his investigation of Amblystoma, 
noticed that its fibres were interrupted by a tendinous inscription at 
about the level of the knee, and hence he suggested that the muscle 
was really composed of two separate muscles fused end to end. The 
proximal portion he regarded as the true ischio-flexorius, but the 
distal portion he thought was the serial homologue of the superficial 
flexor muscle of the fore-arm, which arises from the humerus (M. 
flex. prim, comm.), and he named it accordingly M. plantaris super- 
ficialis medialis. McMurrich based his view largely upon the 
appearance of the cross-section of the leg just below the knee com- 
pared with that of the fore-arm below the elbow, and on the fact 
that otherwise a superficial flexor with parallel longitudinal fibres is 
wanting in the hind-limb {vide infra). 

In Salamandra the relations of the M. ischio-flexorius are very 
similar to those described by McMurrich for Amblystoma. That 
two muscles are involved in the composition of this adult 'ischio- 
flexorius' seems almost certain, since the innervation of the two 
portions is different. While it may be that McMurrich's inter- 
pretation is correct it is nevertheless open to doubt, and, until a 
thorough investigation of the ontogeny of the muscle is available, it 
seems preferable to retain the name 'ischio-flexorius' for the whole 
muscle and to apply the term pars propria to the proximal portion, 
2ind pars plantaris to the distal portion. 



I02 THE SALAMANDER 

Innervation: Pars propria is supplied from N. ischiadicus ventralis 
and pars plantaris from N. sciaticus. 

Function : It is an auxiliary flexor of the knee-joint, and, by virtue 
of its insertion into the plantar aponeurosis, of the toes also. 

M. caudali-puho-ischio-tihialis (m.c.p.i.t.). de Man (1873) ; Hoffmann 
(1873-8); Low (1926). 

Parvus cruris retractor (37) ..... Carus (1828) 

Subcaudalis .... Meckel (1828); Stannius (1854-6) 

Coccy-sous-femoral ...... Duges (1834) 

A long muscular slip ..... Rymer Jones (1852) 

Semimembranosus ....... Mivart (1869) 

Caudo-crural ...... Humphry (1872) 

Deducteur caudal inferieur de la cuisse (112) . . Perrin (1892) 

Caudali-pubo-ischiadicus ..... Osawa (1902) 

Together with M. caudali-femoralis = 

Caudofemoralis ...... Appleton (1928). 

This is the middle member of the three ventral caudal muscles which 
separate from the general subvertebral muscle mass of the tail at the 
level of the first haemal-arch (fourth caudal vertebra). They may in 
fact be regarded as arising from this bone, and from the transverse 
processes of the vertebra. The other two muscles of this series are 
the M. ischio-caudalis, on its mesial side, and the M. caudali- 
femoralis lateral to it. The unusual character of its insertion has 
already been mentioned, namely, by means of a flat tendon into the 
posterior edge of the M. pub. isch. tib. The direction of its fibres is 
almost at right angles to those of the latter muscle. 

Innervation-. From N. pudendus. 

Function: Its main function must be to flex the tail. Whether it has 
any action on the limb is doubtful. 

M. ischio-caudalis. Hofi^mann (1873-8); Osawa (1902); Low (1926). 
Caudae flexor (43) ...... Carus (1828). 

Flexor of the tail ...... Rymer Jones (1852). 

Ischio-caudal . . . Mivart (1869); Humphry (1872). 

Forming the most mesial of the three muscles arising from the fourth 
caudal vertebra, as above described, it passes directly forwards 
between the M. caud. pub. isch. tib. and the cloaca, and is inserted 
on the posterior border of the ischium. It is a flat, ribbon-like 
muscle, and, in the male, it passes between the dorsal and ventral 
portions of the cloacal gland. 

Innervation : From N. pudendus. 

Function: It is a flexor of the tail. Together with the former 



THE MUSCLES 103 

muscle it must have a special function in the male by compressing 
the glandular tissue around the cloaca. It is probably for this reason 
that the tail is strongly flexed during copulation. 

M. caudali-Jemoralis (m.cd.fm.). de Man (1873). 

Tricipitum, capita media ..... Funk (1827). 

Longus femoris reductor et attractor (32) . . Carus (1828). 

Pyriformis .... Meckel (1828); Stannius (1854-6). 

Coccy-sus- femoral ....... Duges (1834). 

Long extensor and Adductor of the thigh . . Rymer Jones (1852). 

Femoro-caudal ....... Mivart (1869). 

Caudo-femoralis . Humphry (1872); Osawa (1902}; Low (1926). 
Caudali-femoralis (pyriformis) . . . . Hoffmann (1873-8). 

Deducteur caudal superieur de la cuisse (116) , . Perrin (1892). 

Together with Caudali-pubo-ischio-tibialis = 

Caudo-femoralis Appleton (1928). 

Like the other two muscles, already described, which arise from the 
fourth and fifth caudal vertebrae, it is a narrow, strap-like muscle 
with an oval cross-section. It is the most powerful muscle of the 
three, and the most lateral. At what Appleton terms 'the posterior 
groin' it turns lateralwards to enter the leg just dorsal to the M. 
ischio-flexorius. It is inserted on the femur, at about a third of its 
length from the pelvic girdle, on a slight tuberosity which may 
probably be regarded as the external trochanter. 

Innervation'. From the N. pudendus. 

Function : It serves either to flex the tail in a lateral direction or to 
retract the femur, according to circumstances. 

The deeper flexors are two in number, and are exposed to view 
on removing the MM. pubo-ischio-tibialis and caudali-pubo-ischio- 
tibialis. 

M. pubo-ischio-femoralis externus (m.p.i.f.e.). de Man (1873); Osawa 
(1902); Low (1926). 

(?) Cruraeus amicus ...... Funk (1827). 

Crassus cruris flexor s. attractor et reflexor (36) . . Carus (1828). 

Nach aussen Zieher ...... Meckel (1828). 

Ex-pelvi-femoral ....... Duges (1834). 

Obturatorius et adductor ischiadicus . . Stannius (1854-6). 

Adductor Mivart (1869). 

Middle part of deeper stratum (adductors and external obturator) 

Humphry (1872). 
Pubo-ischio-femoralis-externus (pectineus) . Hoffmann (1873-8). 

Flechisseur du femur (1 19) ..... Perrin (1892). 
Obturator anterior and posterior . . . Appleton (1928). 



104 THE SALAMANDER 

This muscle arises from the ventral surface of the pelvic girdle, its 
posterior portion lying deep to the M. pub. isch. tib. It tapers 
sharply, so that it has a triangular shape when seen from the ventral 
side. It is inserted along the middle of the ventral surface of the 
femur. Appleton regards the muscle as being composed of two 
sections, and calls them respectively obturator anterior and posterior. 
There is considerable justification for this view, but, since in the 
adult they have fused indistinguishably into a single mass, the older 
name is adhered to. There is, however, an anterior slip of muscle 
arising from the pubis which is distinct, but which was probably 
included by the older authors as forming part of the muscle just 
described (see p. 1 06). 

Innervation : This muscle has a double innervation in accord with 
its double nature. The anterior fibres (Appleton 's obturator anterior) 
are supplied by the obturator nerve, while the posterior fibres 
(Appleton's obturator posterior) receive their supply from the N. 
ischiadicus ventralis. 

Function : It is the true flexor of the hip-joint. 

M. ischio-femoralis (m.is.fm.). Mivart (1869); de Man (1873); 
Osawa (1902). 

Gemellus Meckel (1828). 

„ s. quadratus femoris . . . Stannius (1854). 

Anterior abductor Rymer Jones (1852}. 

Hinder (ischio-femoral) part of deeper stratum of thigh 

Humphry (1872) 
Ischio-femoralis (quadratus femoris ) . . Hoffmann (1873-8). 

Rotateur inverse du femur (124) .... Perrin (1892). 

Ischio-femoralis a and jS Low (1926). 

Together with previous muscle = 

Flexor profundus Appleton (1928). 

This is a small, short muscle which arises from the posterior sector of 
the concave lateral border of the ischium, and passes in a latero-dorsal 
direction to its insertion on the posterior face of the head of the femur. 

Innervation-. From N. ischiadicus ventralis. 

Function: It tends both to depress and to retract the femur, or, 
conversely, to support and project the pelvis. 

With reference to the phylogeny of the ventral thigh muscles 
described above, the conclusions of Appleton (1928, p. 427) may 
be quoted. 

'It is concluded that some modification has occurred in the superficial ventral 
thigh muscles of Urodela to which the extensive distribution of N. ischiadicus 



THE MUSCLES 105 

ventralismaybe attributed. In them again, there is a rectus internus(Humphry). 
There is no reason to beheve that either this or the pubotibiahs (with metazonal 
innervation) were ever present in pro-ReptiHa. A study of modern Urodela, 
moreover, points to a progressive differentiation of M. pubotibiaHs, with the 
formation of an "adducteur du femur" (Perrin) perhaps better distinguished as 
a distinctive urodele formation by the term M. pubtfemoralis. . . . The 
femoro-fibular muscle (Humphry) of urodeles appears to be a caenotelic 
structure, being apparently unrepresented in Salientia and Amniota. There 
is, again, no definite evidence that the fore-runners of Amniota ever possessed 
the caudocrural muscle (of Humphry), which has probably been derived in 
urodela from a common caudofemoral mass along with the caudalifemoralis 
(of de Man). 

'The attachments of the caudo-crural and the course of the N. perinealis 
favour this interpretation rather than a derivation of the caudocrural from the 
ischiocaudal.' 

M. pubo-ischio-femoralis internus (m.p.i.f.i.).^ de Man (1873); Siegl- 
baur (1904); Low (1926). 

Crassis femoris tensor s. protractor (31) . . . Carus (1828). 

Pectineus . Meckel (1828); Stannius (1854-6); Appleton (1928). 

Intra-pelvi-femoral Duges (1834). 

Iliacus Mivart (1869). 

Anterior-suprapubic-portion of the deeper stratum of the plantar of thigh 

s. pectineus Humphry (1872). 

Pubo-ischio-femoralis internus (Adductor) . Hoffmann (1873-8). 

Extenseur du femur (121)2 Perrin (1892). 

Together with M. extensor iliotibialis = 

Extensor femoris biceps (Caput pubicum) . . Osawa (1902). 

A very large, powerful muscle arising from the whole internal face of 
the pubis, and from part of that of the ischium and ilium. It bends 
around the anterior edge of the pubis, and passes down the anterior 
aspect of the thigh. It is inserted along the whole anterior face of 
the femur with the exception of the extreme ends. As disclosed by 
its innervation it belongs to the dorsal, or extensor muscles. 

Innervation-. From N. femoralis, and a few twigs proximally 
from N. obturatorius. 

Function : It has only a relatively slight influence in extending the 
limb in a dorso-ventral plane, but has a very pronounced action in 
drawing the femur towards the body anteriorly. 

Criticism of Perrin. 

Hoffmann describes a muscle under the name 'Pubo-ischio- 
femoralis internus' which he says Von der oberen (Bauch-) Flache 

J See also M. pubo-extensorius, p. 106. ^ See 'Criticism of Perrin'. 



io6 THE SALAMANDER 

des Ospubo-ischii entspringt', while Perrin (i 892) in describing what 
he claims to be the homologous muscle (viz. his Adducteur du 
femur, 117) says: 'Son aponeurose d'insertion occupe a la face 
ventral du pubis, le bord anterieur depuis I'epine pubienne jusqu'au 
commencement du dernier tiers.' His figures confirm this origin 
from the ventral face of the pubis only. The same author, later on, 
describes and figures two muscles (Deducteur du femur, 120, and 
Extenseur du femur, 121) which he considers to be the homologues 
of the ileo-femoralis of Hoffmann. Of the latter muscle he says: 
'C'est un muscle puissant dont I'insertion superieure est tout entiere 
sur la face dorsale du bassin ou elle recouvre tout le pubis. Tangle 
antero-interne de I'ischion, et I'extremite anterieure de I'ilion.' 

Both his descriptions and his figures of these two muscles 
correspond exactly with Hoffmann's ilio-femoralis and pubo-ischio- 
femoralis internus respectively. Perrin himself seems to be aware 
of some discrepancy, since he remarks at the conclusion of his de- 
scription of the extenseur du femur that 'Hoffmann les fait sortir 
uniquement de I'ilion . . .' 

The muscle which Perrin homologizes with Hoffmann's pubo- 
ischio-femoralis internus (his Adducteur du femur) is most pro- 
bably included by Hoffmann, and the other authors, in the muscle 
Hoffmann calls M. pubo-ischio-femoralis externus. The validity 
of this arrangement is questionable since the innervation is different 
from that of the adjoining part of the M. pub. isch. fem. ext. Fol- 
lowing Appleton's suggestion it is therefore described here as a 
separate muscle, M. pubi-femoralis. 

M. pubi-femoralis (m.p.fm.). Appleton (1928). 

Adducteur du femur (117) . . . Perrin (1892). 

This muscle arises from the antero-ventral surface of the pubo- 
ischium, immediately anterior to the M. pub. isch. fem. ext., and 
deep to the M. pub. tib. Some fibres arise from a common raphe 
with this latter muscle. It is relatively weak, and passes obliquely 
lateralwards to be inserted along the ventral face of the femur, 
antero-distally from the M. pub. isch. fem. ext. and posterior to 
M. pub. isch. fem. int. 

Innervation: From N. ischiadicus ventralis. 

Function: It serves to draw the femur towards the body anteriorly. 

M. pubo-extensorius. Low (1926). 

This muscle is only described by Low. By other authors it is in- 
cluded in the M. pub. isch. fem. int., with which muscle it is very 



THE MUSCLES 107 

closely associated in the adult (although from his figure it would 
appear to correspond with Humphry's rectus femoris)^ nevertheless 
it has an insertion distinct from this muscle and is fairly easily 
separable from it, so that, in view of the fact that Low finds it more 
distinct in the larva (he has not investigated Salamandra), it seems 
justifiable to include it as a separate muscle. 

In Salamandra the muscle takes its origin from the antero-dorsal 
edge of the pubis and passes lateralwards along the antero-dorsal 
aspect of the thigh to a tendinous insertion on the knee capsule. 
Although very closely associated with the M. pub. isch. fem. inter- 
nus, it is distinguishable from this muscle distally by its insertion, 
and, starting from this end, it may be fairly easily peeled away from 
it proximally. 

Innervation : From N. femoralis. 

Function: It is an extensor of the thigh. 

The remaining three thigh muscles are quite superficial and are 
strictly dorsal in position (PI. VIII, fig. 47). 

M. extensor iliotibialis (m.e.il.t.). Appleton (1928). 
Cruraeus posticus ..... 

Longus cruris abductor (33) 

Starker Strecker ..... 

Long abductor of the leg ... 

Streckmuskelmasse des Unterschenkels 
Retractors or extensors of the thigh and leg 
Glutaeus maximus. Rectus fcmoris . 
Middle sector of the Ilio-crural s. gluteo-rectus 
Ilio-tibialis ...... 

Ileo-extensorius (extensor cruris) 
Extenseur superficielle du tibia; Tete externe inl 
superieur ...... 

Rectus femoris ..... 

Ileo-extensorius .... Sieglbaur (1904); Low (1926). 

Together with M. pubo-ischio-femoralis internus = 

Extensor femoris biceps (Caput iliacum) . . . Osawa (1902). 

This muscle arises by two separate heads from the dorso-lateral sur- 
face of the ilium and passes superficially along the dorsal, or extensor, 
surface of the thigh to the knee. The two heads usually unite, more 
or less, for a short distance proximal to the knee, although they may 
remain separate and appear as two discrete muscles. In either case 
their distal ends pass into a strong flat tendon which fuses with the 
connective tissue of the knee capsule, and finally attaches itself to the 



Funk (1827). 

Carus(i828). 

. Meckel (1828). 

Rymer Jones (1852). 

Stannius (1854-6). 

. Owen (1866). 

. Mivart (1869). 

Humphry (1872). 

. de Man (1873). 

Hoffmann (1873-8). 

erieur, et Tete externe 

Perrin (1892). 

Eisler (1895). 



io8 THE SALAMANDER 

tibial spine, passing between the MM. extensor digitorum com- 
munis and extensor cruris tibialis in order to do so. Both sections 
are narrow, oval in cross-section, and of medium strength. 

Authors have homologized this muscle in different ways. 
Humphry and Mivart, in Cryptobranchus, and Noble in the 
Salientia, regard it as composed of two distinct muscles. De Man, 
in Triton, regards any separation into two sections as artificial. 
Perrin, in Salamandra, describes it as one muscle with two heads, 
while Appleton — whose name is adopted here — in his list of muscle 
homologies (Part II, pp. 434-5) calls it M. extensor iliotibialis, and 
distinguishes the two heads as pars anterior and pars posterior^ while 
the same author in another place (Part I, p. 378, fig. 2) labels the 
former section Ilio-tibialis, and the latter Ilio-extensorius, thus 
following Noble. 

Innervation : From N. extensorius. 

Function : True extensor of the leg. 

M. ilio-fibularis (m.il.f.). de Man (1873); Sieglbaur (1904); Osawa 
(1902); Appleton (1928). 

Semimembranosus ....... Funk (1827) 



Wadenbeinbeuger ..... 
Ileo-peronien ..... 

Abductor fibularis primus 
Ileo-peroneal ..... 

Biceps flexor cruris (outer or ilio-fibular sector) 
Deducteur du fibula (i 13) 
Ilio-femoro-fibularis .... 



. Meckel (1828} 

. Duges (1834), 

Stannius (1854-6), 

. Mivart (1869) 

Humphry (1872) 

Perrin (1892) 

Low (1926) 



Together with M. femoro-fibularis = 

Ileo-femoro-fibularis (biceps) .... Hoffmann (1873-8). 

Most authors regard this muscle as the long head of a biceps 
femoris muscle. The validity of this is doubtful, and the question 
will be discussed in detail after the description of what is regarded 
by these authors as the short head, namely, the M. femoro- 
fibularis. 

The M. ilio-fibularis arises from the external face of the ilium, 
posterior to the M. extensor ilio-tibialis. It passes down the postero- 
dorsal border of the thigh to its tendinous insertion on the proximal 
end of the fibula. The N. peroneus passes over its dorsal edge at its 
insertion. It is a relatively weak muscle. 

Innervation : From N. extensorius. 

Function: In conjunction with the M. extensor ilio-tibialis it 
would tend to extend the leg, but acting with the M. pubo-tibialis 



THE MUSCLES 109 

it would tend to flex it. Probably the latter is its principal and 
stronger function. 

M.femoro-fibularis (m.fm.f.). Humphry (1872); Osawa (1902). 

Popliteal Cams (1828) 

Femoro-peronien ....... Duges (1834 

Abductor fibularis secundus .... Stannius (1854-6) 

Flechisseur primitifdu fibula (i 14) .... Perrin (1892) 

Ilio-femoro-fibularis ...... Low (1926) 

Together with M. ilio-fibularis = 

Ileo-femoro-fibularis (biceps) .... Hoffmann (1873-8). 

This muscle is considered here on account of its supposed associa- 
tion with the preceding one, although it hardly belongs to the thigh 
muscles proper. It is a short, narrow muscle band arising from the 
median ventral face of the femur, near the distal end, and, passing 
across the popliteal space behind the knee, is inserted on the postero- 
lateral face of the proximal end of the fibula. 

Innervation : From N. sciaticus. 

Function : It is the direct flexor of the knee. 

This muscle is regarded by Hoffmann and most of the earlier 
authors as the short head of a biceps muscle, of which the other 
longer head is the M. ilio-fibularis. Hoffmann states that the two 
muscles are joined at their insertion, but this is incorrect so far as 
Salamandra is concerned, for, although their insertions are always 
close together, they are easily and naturally separable. The strongest 
fact against regarding the two muscles as the two heads of a biceps is 
that their innervation is entirely diff"erent, the M. ilio-fibularis being 
innervated by the dorsal nerve — N. extensorius — and the M. 
femoro-fibularis by the ventral, or flexor nerve — N. sciaticus. It 
therefore seems more correct to regard them as two distinct and 
unrelated muscles, and this is the view held by the later writers, e.g. 
Perrin and Appleton. The latter author regards the M. femoro- 
fibularis as 'a caenotelic structure, being apparently unrepresented in 
Salientia and Amniota' (see also p. 105). 

M. ilio-femoralis. de Man (1873); Osawa (1902); Low (1926); 
Appleton (1928). 

Ileo-rotulien ....... Duges (1834). 

Thick flexor of the thigh .... Rymer Jones (1852). 

Gluteus ....... Stannius (1854-6). 

Gluteus medius and minimus ..... Mivart (1869). 

Ileo-femoral stratum of the dorsal muscles of the thigh 

Humphry (1872). 



no THE SALAMANDER 

Ileo-femoralis (Iliacus) Hoffmann (1873-8). 

Deducteur du femur (120)1 Perrin (1892). 

This is the deepest of the dorsal thigh muscles. It takes its origin 
from the postero-lateral face of the ilium, and to some extent from 
the dorsal (inner) face of the ischium also. It is inserted along the 
middle portion of the posterior face of the femur. Low (1926) dis- 
tinguishes the anterior from the posterior portion. 

Innervation : From N. extensorius. 

Function: It tends to draw the femur towards the tail and is there- 
fore the opposer of the M. pub. isch. fem. int. 

3. Extensor Muscles of the Leg and Foot (PL VIII, fig. 47). 
These should be dissected from the dorsal side. 

M. extensor digitorum communis (m.e.d.). Funk (1827); Ribbing 
(1909). 

Longus digitorum extensor (40) .... Carus (1828). 

Peroneo-sus-digital. ...... Duges (1834). 

Extensor longus digitorum pedis . . . Rymer Jones (1852). 

Extensor hallucis longus, et Extensor quatuor digitorum longus 

Schmidt, Goddard, and van d. Hoeven (1864}. 
Extensor longus digitorum . Mivart (1869); Humphry (1872). 

Femoro-digiti I-V ..... Hoffmann (1873-8). 

Extenseur des metatarsiens ..... Perrin (1892 a). 

Extenseur commun des doigts (76) .... Perrin (1892). 

Extensor digitorum longus ..... Osawa (1902). 

Femoro-metatarsalis ..... Sieglbaur (1904). 

Together with the following muscle = 

Extensor digitorum communis longus . . . Eisler (1895). 

This is the most superficial muscle on the extensor side of the leg. 
It arises from the epicondylus lateralis femoris^ between the MM. 
extensor cruris tibialis and fibularis, by means of a narrow tendon. 
The muscle spreads out rapidly and becomes thin and fan-shaped. 
At the bases of the digits it breaks up into nine small tendons, eight 
of which are inserted on either side of the bases of the metatarsals of 
the four post-axial digits, while the remaining one is inserted on the 
fibular side of the first metatarsal. Sieglbaur denies the existence of 
this ninth tendon, but it certainly exists in the majority of cases, if 
not in all. There is a slight aponeurosis between the proximal por- 
tion of this muscle and the M. extensor cruris fibularis. 

Innervation : By the NN. dorsales pedis, from N. peroneus. 

Function: It is the chief extensor of the foot. 

^ See also ' Criticism of Perrin', pp. 105-6. ■ 



THE MUSCLES in 

M. extensor tarsi tibialis (m.e.t.t.). Ribbing (i 909). 

Rotateur inverse du pied (92) ..... Perrin (1892). 

Extensor tarsi tibialis longus ..... Eisler (1895). 

Together with M. extensor cruris tibiahs == 

Tibialis anticus . Funk (1827); Schmidt, Goddard, and van d. Hoeven 

(1864); Humphry (1872). 
Abductor et flexor pedis +introflexor pedis (39). . Carus (1828). 

Femoro-tibialis ...... Hoffmann (1873-8). 

Tibialis .... Osawa (1902); Sieglbaur (1904). 

This is a small muscle which arises from the epicondylus lateralis 
jemoris very close to, or from a common tendon with, the M. ext. 
dig. comm. Although its origin is slight, the fibres soon swell into 
a spindle-shaped muscle of moderate size which passes alongside 
the M. extensor cruris tibialis to be inserted on the ventral surface 
of the tibiale and cart, pre-hallucis. It is easily separated from the 
neighbouring muscles, and in this respect differs from the corre- 
sponding muscle in the fore-arm. It is a fairly superficial muscle 
but is partly covered by the M. ext. dig. communis. 

Perrin homologizes this muscle with Hoffmann's M. fibulae 
metatarsum II and Humphry's M. supinator pedis, admitting the 
discrepancy between the origin and insertion of the muscle described 
by the German and English authors and of the one which he himself 
describes. There does not appear to be a muscle in the Salamander 
which exactly corresponds with Humphry's supinator pedis, perhaps 
the nearest being the M. abductor digiti I. This muscle admittedly arises 
from the intermedium, and not from the fibula, but it is inserted on 
the metatarsal and /j joined to the M. extensor digiti brevis superior I. 
Further, it is definitely a muscle of the deep stratum, while the 
rotateur inverse du pied of Perrin is rather to be associated with the 
superficial muscles, and in Cryptobranchus (Menopoma) Eisler 
figures the muscle as actually arising from the fibula as well as from 
the intermedium (Eisler 1895, ^'^^- ^^^' ^&- ^5' 7 ^)- ^^^^ absence 
of figures in Hoffmann's account renders identification difficult, 
but there can be no doubt that his description in general has been 
largely influenced by Humphry's, and that the identification of 
Hoffmann's M. fibulae-metatarsum II with Humphry's M. supinator 
pedis is sound. 

It seems, therefore, probable that Perrin's M. rotateur inverse du 
pied is included by Humphry in his 'Tibialis anticus passing to 
tarsus', and by Hoffmann in his M. femoro-tibialis. 

Innervation: From N. dorsalis pedis intermedius. 

Function : It tends to turn the foot into the supine position. 



112 THE SALAMANDER 

M. extensor cruris tibialis (m.e.c.t.). Ribbing (i 909). 

Extenseur tarsien interne +extenseur primitif du tibia Perrin (1892). 
Extensor tarsi tibialis brevis+femoro-tibialis superficialis+femoro- 

tibialis profundus Eisler (1895). 

Together with M. extensor tarsi tibialis = 

Tibialis anticus . Funk (1827); Schmidt, Goddard, and van d. Hoeven 

(1864); Humphry (1872}. 
Abductor et flexor pedis + Introflexor pedis (39). . Carus (1828). 

Femoro-tibialis Hoffmann (1873-8). 

Tibialis .... Osawa (1902); Sieglbaur (1904). 

Arising from the epicondylus lateralis jemoris the muscle is inserted 
along the whole lateral border of the tibia and extends also to the 
tihiale and cart, pre-hallucis, while some of the proximal fibres pass 
to the mesial side of the crista tibiae. It will be noticed that both 
Eisler and Perrin distinguish between the portion which is inserted 
on the tarsus and that which is inserted on the tibia, but any 
such differentiation must be regarded as largely artificial, or at most 
incipient, in the Salamander. 

Innervation: From N. dorsalis pedis intermedius and N. dors, 
ped. tibialis (N. saphenus). 

Function : Extensor of the tibia and leg. 

It should be observed that the separation of the pre-axial extensor 
muscle mass into two is more complete in the hind-limb than in the 
anterior limb. 

MM. extensores digitorum breves (m.e.b.). Eisler (1895); Osawa 
(1902). 

Extensor brevis ....... Carus (1828). 

Extensor hallucis brevis -j-extensor quatuor digitorum brevis 

Schmidt, Goddard, and van d. Hoeven (1864). 
Extensor brevis digitorum . . Mivart (1869); Humphry (1872). 

Tarsu-digiti I-V Hoffmann (1873-8). 

Extenseurs des doigts ...... Perrin (1892). 

Extensor digiti 1; extensores breves digitorum II-V . Sieglbaur (1904). 
Extensores breves superficiales et profundi . . Ribbing (1909). 

Two Strata of the digital extensor muscles are distinguishable and 
will be referred to as- 'superficialis' and 'profundus' respectively. 
They arise generally from the tarsal elements and are inserted by 
means of a tendon on the base of the terminal phalanx of each finger. 
Each tendon is also attached to the other phalanges by means of 
small lateral lips at the inter-phalangeal joints. Hoffmann and Eisler 



THE MUSCLES 113 

wrongly describe these tendons as being associated with the M. ext. 
dig. communis. They have nothing whatever to do with this muscle 
(cf. Sieglbaur and Perrin). 

The details of the several digits will be dealt with individually. 

MM. abductor et extensor digiti I (m.a.e. i). Sieglbaur. 

MM. abductor et extensor hallucis . . Osawa (1902). 

These muscles arise from the dorsal surfaces of the intermedium and 
centrale and pass obliquely across the tarsus. The abductor portion 
is inserted on the lateral face of the metatarsal, while the extensor 
portion is distinguishable therefrom by its tendinous insertion on the 
phalanx of the same digit. The abductor portion of this muscle is 
probably identical with Humphry's M. supinator -pedis and Hoff- 
mann's M. fibulae-metatarsum II (see also p. 1 1 1). 

The M. extensor digiti brevis superficialis II arises alongside the pre- 
vious muscle from the intermedium and centrale^ and passes across the 
tarsus to the level of the base of the proximal phalanx of the second 
digit. Here it passes into a tendon which is inserted on this phalanx, 
and on the base of the terminal one, in the manner already described. 

The M. extensor digiti brevis superficialis III arises by two heads : 
(i) from the intermedium together with I and II, and (ii) from the 
dorsal surface of the fibulare. The two heads pass into a common 
tendon which is inserted on the bases of the phalanges of the third 
digit. 

The M. extensor digiti brevis superficialis IV has a single origin from 
the fibulare and has the typical insertions on the phalanges of the 
fourth digit. 

The M. extensor digiti brevis superficialis V also arises from the fibu- 
lare and is inserted on the bases of the phalanges of the fifth digit. 
It is a relatively slight muscle. 

The MM. extensores digitorum breves profundi of the several digits 
arise from the respective basal elements of the tarsus. It is note- 
worthy, however, that the muscle to the first digit arises from the 
united basals of the first and second digits and not from the cart, pre- 
hallucis (cf. p. 42). They are all short feeble muscles and are 
inserted on the dorsal side of the tendons of the corresponding 
superficial portions at about the level of the distal ends of the 
metatarsi. 

Eisler describes and figures three layers of extensor digiti muscles 

4038 I 



114 THE SALAMANDER 

for Menopoma, namely, 'superficialis', 'medius', and 'profundus', 

but the 'medius' layer is indistinguishable in Salamandra. 

Innervation : From the NN. dorsalis pedis intermedius and fibu- 
laris. 

Function: To extend the digits. 

M. extensor cruris et tarsi fibularis (m.e.c.t.f.). Ribbing (i 909). 

Extensor and abductor of the tarsus . , . Rymer Jones (1852). 

Peroneus longus . Schmidt, Goddard, and van d. Hoeven (1864). 

Peroneus . Humphry (1872); Osawa (1902); Sieglbaur (1904); 

Appleton (1928). 
Femoro-tarsali-fibulare+femoro-fibularis . Hoffmann (1873-8). 

Extenseur tarsien externe (94), Extenseur primitif du fibula (96) 

Perrin (1892). 
Part of extensor digitorum communis longus +femorofibularis brevis 

Eisler (1895). 

The superficial and deep strata of the extensor muscle on the post- 
axial side of the limb are not so easily separable as on the tibial side. 
They arise together from the epicond. lat. fern, immediately poster- 
ior to the M. ext. dig. comm. as well as from an aponeurosis on the 
post-axial border of this muscle. The superficial fibres — represent- 
ing the muscle designated by the first name in the synonyms of 
Ribbing, Rymer Jones, Hoffmann, Perrin, and Eisler — are inserted 
on the fibularcy while the deeper fibres are inserted all along the 
lateral face of t\\Q fibula. 

Innervation: From N. peroneus. 

Function : Extensor and abductor of the fibula and post-axial side 
of the limb. 

4. Flexor Muscles of the Leg and Foot (PL VIII, figs. 48 and 49). 

These should be dissected from the ventral side. 
M. flexor primordialis communis (m.f.p.). Ribbing (1909). 
Together with M. flexor accessorius lateralis = 

Flechisseur externe des doigts ..... Perrin (1892}. 

Plantaris superficialis major and minor . . . Eisler (1895). 

Plantaris profundus III ..... McMurrich (1904). 

Together with MM. flexores accessorii and pronator profundus = 

Flexor digitorum sublimis ..... Osawa (1902). 

Together with MM. flexores breves superficiales = 

Superficial stratum of plantar muscles of leg and foot, s. flexor sublimis 
digitorum ....... Humphry (1872). 

Together with MM. flexores breves superficiales and flexores accessorii = 
Flexor digitorum communis ..... Funk (1827). 



THE MUSCLES 115 

Flexor longus (41) Carus (1828). 

Oberflachlicher Beuger Meckel (1828). 

Flexor longus digitorum pedis . , . . Rymer Jones (1852). 

Flexor quatuor digitorum communis longus, and flexor hallucis proprius 

Schmidt, Goddard, and van d. Hoeven (1864). 

Flexor digitorum Mivart (1869). 

Femoro-fibulae-digiti I-V .... Hoffmann (1873-8). 

This muscle differs somewhat from its counterpart in the anterior 
limb, since instead of taking its origin entirely from the lateral con- 
dyle of the proximal bone of the limb — the epicond. lat.jemoris — the 
greater portion of it arises from the distal joint of the limb, viz. from 
the lateral face of the fibula. The direction of the fibres is approxi- 
mately parallel with the axis of the limb. In the region of the tarsus 
the muscle passes into a strong, broad aponeurosis spreading over 
the sole of the foot — the fascia planiaris (fas.pl.) (Eisler). This fascia 
divides up into five strong tendons which pass along the flexor side 
of the digits and are inserted in the proximal end of the terminal 
phalanx of each toe. Where the toe possesses more than one phalanx 
the tendon is strapped down to the sides of the proximal end of each 
by means of small lateral slips. The. fascia plantaris is thus equivalent 
to the palmar fascia of the hand. It is joined on its ventral — super- 
ficial — side by the tendon of the M. ischio-fiexorius, while on the 
dorsal — deep — face the MM. flexorius accessorius medialis and late- 
ralis are inserted, and the MM. fiexores breves superficiales originate. 
It should be noticed that the fibres on the fibular side of the leg are 
necessarily very short, while those on the tibial side pass along the 
whole length of the leg. 

Both Eisler and McMurrich distinguish from the above de- 
scribed major portion of the muscle a small slip which they respec- 
tively call Plantaris superficialis minor and Profundus III minor. This 
portion is relatively quite small and arises from the ventro-mesial 
edge of the fibula immediately proximal to the origin of the M. 
caput longum musculorum contrahentium. It passes somewhat 
obliquely to fuse with the major portion on its tibial border. Ribbing, 
probably more correctly, regards it as a portion of the M. flexor 
accessorius medialis (see p. 116). 

Innervation: Both portions are supplied by the main branch of 
N. fibularis, and by its R. medianus. 

Function: It mainly serves to flex the digits and the tarsus, but, 
owing to its origin spreading along the whole length of the fibula and 
the inequality in the length of the fibres on the fibular and tibial 
sides of the limb, the contraction of the muscle must also tend to 



ii6 THE SALAMANDER 

rotate the leg so as to turn the pre-axial side of the limb inwards and 

caudalwards. 

M. flexor accessorius lateralis (m.f.a.l.). Ribbing (1909). 
This forms a small sheet of muscle arising mainly from the lateral 
edge of thtfibulare^ while a few fibres take their origin from the fifth 
basale and from the distal end of the fibula. The fibres pass obliquely 
tibialwards to about the middle of the tarsus where the muscle is in- 
serted on the/«ia<^/>/<^///^m of the M. flexor prim. comm. Indeed most 
authors regard it as forming a part of the latter muscle, but, as Ribbing 
correctly points out, they are clearly separated by the fibular nerve. 

Innervation : By the terminal twigs of R. medianus, N. fibularis. 

Function: It is accessory to the M. flex. prim, comm., with a dis- 
tinct pronating action. 

M. flexor accessorius medialis (m.f.a.m.). Ribbing (i 909). 

Flechisseur interne des doigts ..... Perrin (1892). 

Plantaris profundus I, II, et III .... Eisler (1895). 

Plantaris profundus II McMurrich (1904). 

Together with M. pronator profundus = 

Pronator pedis ...... Humphry (1872). 

Together with M. flexor primordialis communis and M. pronator pro- 
fundus = 

Flexor digitorum sublimis ..... Osawa (1902). 
Together with M. pronator profundus and MM. flexores breves superficiales 
may = 

Femoro-fibulae metatarsi I, II, III . . . Hoffmann (1873-8). 

This is a larger and more powerful muscle than the former whose 
fibres run parallel with it and are separated from it by the M. caput 
longum musculorum contrahentium. It arises from the distal two- 
thirds oit\i& fibula on the ventro-mesial aspect of this bone, from the 
fihulare^ intermedium^ and from the basalia III—V inclusive. It is 
inserted over the dorsal surface of th.Q fascia plantaris on the tibial side 
of the M. flex. ace. lat., while its insertion also extends rather more 
proximally than this muscle. 

Innervation : By terminal twigs from R. medianus, N. fibularis. 

Function: Like the former muscle it is accessory to the M. flex, 
prim. comm. with a distinct pronating action. 

MM. flexores breves superficiales (m.f.b.s.). Eisler (1895); McMur- 
rich (1904); Ribbing (1909). 

Flechisseur de la I, II, III, IV, V phalange; Flechisseur du I, II, III, 



THE MUSCLES 117 

IV, V metatarsien; Flechisseur de la III, et IV phalangine; Flechisseur 
de la IV phalanginette ..... Perrin (1892). 

Flexores digitorum sublimes breves .... Osawa (1902). 
Together with the M. flex. prim. comm. = 

Flexor digitorum sublimis .... Humphry (1872). 

Together with flexores accessorii and flex. prim. comm. = 

Femoro-fibulae-digiti I-V .... Hoffmann (1873-8). 

This series of small muscles correspond very closely with the similarly 
named muscles of the hand. They arise from the dorsal side of the 
fascia plantaris and from its digital tendons. The details of the 
several digits vary somewhat as follows : 

Digit I possesses one small muscle slip inserted along the tibial 
border and distal end of the metatarsal. 

Digit II has three small fascicles, two lateral inserted on either 
side of the metatarsal, and a median inserted on the base of the 
proximal phalanx. 

Digit III has five muscle slips recognizable as belonging to 
this series. These comprise two lateral pairs, one pair inserted on 
each side of the metatarsal, and the other pair, somewhat smaller, 
inserted on the base of the proximal phalanx. The remaining 
slip is a median one and is inserted on the base of the middle 
phalanx, but although it is a median muscle its insertion is double 
so as to allow for the passage of the insertion tendon of the M. inter- 
phalangeus. 

Digit IV IS exactly similar to digit III. 

Digit V has a very small median slip attached to the base of the 
proximal phalanx and a pair of lateral muscles inserted on the sides 
of the metatarsal. 

Innervation-. All the slips are innervated by twigs from the 
plantar nerve loop. 

Function : To flex the digits. 

M. caput longum musculorum contrahentium (m.c.l.c.) Ribbing (1909). 
Flechisseur commun profond des phalanges . . Perrin (1892}. 

Fibulo-plantaris ....... Eisler (1895). 

Flexor digitorum profundus ..... Osawa (1902). 

Fibulo-tarsalis ...... McMurrich (1904). 

Together with the following muscle = 

Flexor digitorum profundus .... Humphry (1872). 

Fibulae-metatarsi et digiti I-V . . . Hoffmann (1873-8}. 

This muscle resembles the muscle of the same name in the anterior 
limb and is narrow and ribbon-like. It arises from the mesial side of 



ii8 THE SALAMANDER 

the proximal end of the fibula, and passes directly down the leg to 
the tarsus, where it merges into a flat transverse tendon which is in- 
serted on the distal row of tarsal bones — the basalia. As in the hand, 
the MM. contrahentes digitorum arise in part from this tendon. 

Innervation: From N. fibularis, and mainly from its R. medianus. 

Function : It is the deep flexor of the tarsus. 

MM. contrahentes digitorum (m.c.d.). Ribbing (i 909). 

Flechisseurs profonds des phalanges .... Perrin (1892). 
Flexores breves medii s. contrahentes digitorum . Eisler (1895). 

Flexores digitorum profundi breves .... Osawa (1902). 
Flexor brevis medius ..... McMurrich (1904). 
Together with the former muscle = 

Flexor digitorum profundus .... Humphry (1872). 

Fibulae-metatarsi et digiti I-V . . . Hoffmann (1873-8). 

As already mentioned, this series of small flexor muscles arises partly 
from the tendon of insertion of M. cap. long. muse, contra., and 
partly from the basalia. The slip going to the first digit is slight 
and weak, while its direction is very oblique owing to the fact that 
it arises at the base of the second digit from the fused basale I + 11. 
The slip to the second digit also takes origin from this double basale 
and from basale III, while the third slip arises from basalia III 
and IV. The fourth slip arises only from basale IV, but the fifth 
originates from both IV and V. The insertion of each slip is by 
means of a short tendon on the base of the proximal phalanx of the 
corresponding digit. 

Innervation: By twigs from the plantar nerve loop which pass 
dorsal to the muscles, thus separating them from the MM. flexores 
breves profundi. 

Function : In the case of the first digit the muscle acts mainly as an 
adductor, and only slightly as a flexor, while in the second, third, and 
fourth the action is almost purely flexor, and in the fifth toe both 
adductor and flexor. 

M. pronator prof undus (m.p.p.). Ribbing (i 909). 

Pronateur du pied Perrin (1892 a). 

Rotateur direct du pied Perrin (1892). 

Fibulo-tarsalis ....... Eisler (1895), 

Plantaris profundus I McMurrich (1904). 

Together w^ith M. flex. ace. med. = 

Pronator pedis ...... Humphry (1872). 



THE MUSCLES 119 

and probably 

Femoro-fibulae-metatarsi I, II, and III . . Hoffmann (1873-8). 

Together with M. flex. prim. comm. and MM. flexores accessorii = 

Flexor digitorum sublimis ..... Osawa (1902). 

This Is the deepest of the three pronating muscles of the leg. It 
arises from the mesial side of the fibula, deep to the M. flex. ace. med. 
Its fibres converge towards their insertion on the lateral face of 
the distal end of the tibia, the tibiale, the cart, pre-hallucis, and 
the base of the first metatarsal. The muscle is thus triangular in 
shape. 

Innervation : From N. interosseus. 

Function : It is the chief pronator of the foot. 



Fibulae-tibialis 
Interosseux de la jambe . 
Fibulotibialis s. interosseus 
Interosseus . 



M. interosseus cruris. Ribbing (1909). 

Pronator tibiae . . . Humphry (1872); Osawa (1902) 

. Hoffmann (1873-8) 

Perrin (1892) 

Eisler(i895) 

. McMurrich (1904) 

Like the muscle in the fore-arm this is a thin sheet of muscle joining 
the mesial sides of the two bones in the distal joint of the limb — in 
this case the fibula and tibia. It may be regarded as arising from the 
proximal portion of the fibula and inserted on the distal part of 
the tibia. It lies deep to the M. pronator profundus as seen from the 
flexor side. 

Innervation : From N. interosseus. 

Function : It forms an elastic ligament between the two bones of 
the leg. 

MM. flexores breves profundi (m.f.b.p.). Eisler (1895); Ribbing 
(1909). 

Tarso-metatarsales . . . Humphry (1872); Osawa (1902) 

Tarso-metatarsi I-V ..... Hoffmann (1873-8) 
Flechisseurs primitifs des metatarsiens . . . Perrin (1892) 

Flexor brevis profundus .... McMurrich (1904) 

These form yet another series of short flexor muscles to the digits 
They lie immediately deep to the MM. contrahentes digitorum, 
and are short, bulky muscles arising from the distal faces of the 
basalia. Each muscle, except that of the first digit, has three inser- 
tions, namely, on either side of the corresponding metatarsal and on 
its ventral face. The lateral insertions extend along practically the 
whole length of the bone, but the ventral portion does not extend 



I20 THE SALAMANDER 

more than half-way, i.e. as far as the origin of the MM. flexores 
digitorum minimi. They are necessarily short but nevertheless rela- 
tively powerful muscles, the post-axial slip being rather longer than 
that on the pre-axial side. On the post-axial side of the fifth digit the 
origin of the muscle extends right back along the side of basale V^ 
while a few fibres may even arise from t\v^ fihulare . In the first digit 
there is but a single slip which passes obliquely from the composite 
basale I+II to the ventral side of the first metatarsal. This muscle 
is therefore triangular in shape, and since there is no M. flexor 
digiti minimus in this digit, its insertion is able to extend along the 
whole length of the metatarsal. 

Innervation : From the plantar nerve loop. 

Function: The muscles serve both to flex the tarso-metatarsal joint 
and in some measure also to approximate the digits. 

M. abductor digiti F (m. 2.. d.^.). Eisler (i 895). 

Abductor minimi digiti . Schmidt, Goddard, and van d. Hoeven( 1864); 

Humphry (1872). 
Fibulae-metatarsum V . . . . . Hoffmann (1873-8). 

Deducteur du fibulaire + Deducteur du cinquieme metatarsien 

Perrin (1892). 
Abductor digiti quinti ...... Osawa (1902). 

Abductor ossis metatarsi quinti . . . McMurrich (1904). 

Included in previous muscle by Ribbing (1909). 

This is a superficial muscle lying along the extreme post-axial sur- 
face of the tarsus. It originates from the distal end of the fibula and 
is inserted on the post-axial surface of the Jibu/are and basale V^ and 
on the base oi the. fifth metatarsal. Its insertion is fleshy. 

Perrin distinguishes two muscles according to whether the inser- 
tion is on the tarsal elements or on the metatarsal. There does not 
seem to be any ;/(^/z/r^/ division of the general muscle mass, however, 
and therefore such a distinction seems hardly justified. 

Innervation : From N. fibularis. 

Function : It is the abductor of the fifth toe. 

MM. flexores digitorum minimi. Eisler (1895); Ribbing (1909). 

Metacarpo-phalangeii ..... Humphry (1872). 



Metatarso-phalangei 

Flechisseurs primitifs des phalanges . 

Metatarso-phalangeales . 

Flexor brevis profundus (median slips) 

These are short muscular slips arising from about the middle of the 
^ An obvious /apsus calami. 



Hoffmann (1873-8). 

Perrin (1892). 

Osaw^a (1902). 
McMurrich (1904). 



THE MUSCLES 121 

metatarsals and are inserted on the bases of the proximal phalanges 
immediately deep to the MM. contra, digit. No representative of 
this series is present in the first digit. 

Innervation: From the plantar nerve loop. 

Function : They are the flexors of the first joint of each digit. 

MM. interphalangei (m.i.ph.3,4). Eisler (1895). 

Phalange! ....... Humphry (1872). 

Phalangi I phalanx II digiti III, IV . . Hoffmann (1873-8). 

Flechisseur primitif de la troisieme (quatrieme) phalangine 

Perrin (1892). 
Phalangeales ....... Osawa (1902). 

Interphalangei III und IV .... . Ribbing (1909). 

Two small muscles occur on the third and fourth digits between the 
proximal and middle phalanges. They have tendinous insertions 
on this latter bone. 

Innervation : From the plantar nerve loop. 

Function: They are the flexors of the interphalangeal joint. 

MM. intermetatar sales (m.i.mt.). McMurrich (1903). 

Interossei metacarpales^ ..... Humphry (1872). 

Interossei metatarsals ..... Hoffmann (1873-8). 

Intermetatarsiens ....... Perrin (1892). 

Interossei . . Eisler (1895); Osawa (1902); Ribbing (1909). 

These correspond to the MM. intermetacarpals of the hand. They 
spread web-like across the interdigital space between the metatarsals 
and can only be seen fully by removing the MM. flexores breves 
profundi. They extend rather more distally along the fibular side of 
each metatarsal than along the tibial side. 

Innervation : Also from the plantar nerve loop. 

Function : They serve to approximate, or adduct the digits. 

The limb muscles of Salamandra are to be regarded as being in 
a state of incipient differentiation and specialization. A glance at the 
lists of synonyms will bear testimony to this fact, since in many cases 
scarcely any two investigators share the same opinions as to how 
a given muscle mass should be divided. Thus, while every effort has 
been made to ensure that these lists shall be accurate and complete, 
it is not possible to give the exact equivalent in all cases. 

It will have been noticed that there is a general agreement between 
the arrangement of the muscles in the fore- and hind-limbs, parti- 
cularly in respect of the distal portions of the limbs. One noteworthy 

' Again obviously a lapsus calami. 



122 THE SALAMANDER 

exception must be mentioned, however, namely, the entire absence in 
the leg of any muscles corresponding with the M. flexor antibrachii 
et carpi radialis, the M. flexor carpi ulnaris, and the M. flexor anti- 
brachii ulnaris of the fore-arm. Doubtless correlated with this differ- 
ence is the fibular origin, in the hind-limb, of the M. flex. prim, 
comm. in addition to its femoral origin, since in the anterior limb this 
muscle originates from the humerus alone. 

It is not possible in the present work to compare the muscles of 
Salamandra with those of related groups or with the Mammalia, 
since such a discussion must necessarily be lengthy, and moreover 
is to be found in the works of Ribbing mentioned in the Bibliography. 

BIBLIOGRAPHY OF SECTION IV 

19, 96, 103, 106, 113, 114, 143, 144, 145, 146, 148, 160, 161, 162, 165, 
168, 187, 206, 208, 256, 322, 333, 355, 432, 433, 434, 435, 440, 441, 454, 
457^ 458, 459> 460, 461, 487, 517, 518, 519, 542, 565, 566, 567, 568, 570, 
579, 618, 619, 620, 633, 634, 637, 645, 646, 647, 648, 664, 667, 698, 702, 
705, 708, 720, 764, 794. 



V 
THE NERVOUS SYSTEM 

BRAIN AND SPINAL CORD 

1. Historical. 

Funk, in his general monograph (1827), describes the external fea- 
tures of the brain, but his figures are poor. In 1890 Oyarzun pub- 
lished some account of the histology of the fore-brain, and a year 
later Burckhardt gave a very good account of the brains of Triton 
and Ichthyophis. His work has been used as a basis by most subse- 
quent investigators of the brains of Urodeles. In 1897 Gehuchten 
discusses the habenular ganglia and commissure. Bochenek (1899) 
and Rubaschin (i 903) add contributions to the histology of the fore- 
brain, whilst more recently Rothig (191 1—27) has made a vary care- 
ful and extensive study of the constitution of the several parts of the 
brain of a number of Urodeles, including Salamandra. Driiner 
(1901) gives excellent figures of the brain of Salamandra, both larval 
and adult, in his paper on the muscles and nerves of the hyobranchial 
apparatus, but he does not describe them. 

Some five workers have made a special study of the pituitary body, 
or hypophysis cerebri. The first of these was Haller (1896). His 
work is very good, and he recognizes that the infundibular portion 
undergoes considerable atrophy, but fails to differentiate more than 
one region in the hypophysial part. He was followed by Bochenek 
(1902), who, working with Golgi's neurological methods, concen- 
trated on the distribution of the fibre tracts within the infundibulum. 
Haller returned to the subject with a larger comparative account in 
1909, but it was not until 1921 that a complete description of the 
hypophysis was given by Atwell, while three years later a confirma- 
tory account was published by Sumi (1924). 

The present account deals only with the general structure and 
external features, including the main commissures and cavities visible 
in a median sagittal section. The works above referred to, and others 
of a more general nature included in the bibliography, must be con- 
sulted if further details are required. 

2. Membranes of the Brain and Spinal Cord. 

Our knowledge of the membranes investing the brain and spinal 
cord in Urodeles is still far from being adequate. Some account of 



124 THE SALAMANDER 

them was given by Rex in 1 893, in his paper on the blood-vessels of 
the brain in Triton and Salamandra. Five years later, in 1898, Miss 
Helen O'Neill made a very careful and detailed study of this subject, 
taking Salamandra as her type. She describes two membranes, a 
dura mater — of which there are two layers — and a primary vascular 
coat. Later Sterzi in a series of papers (1899— 1909) reviews the 
structure of the meninges of vertebrates in general, and supposes that 
there is only one primary covering to the brain in Urodeles — the 
' Meninx primitiva' — from which may separate a ' Meninx secundaria' 
(corresponding with O'Neill's primary vascular coat), both of which, 
in higher animals, give rise to the pia mater and arachnoid. More 
recently Flexner (1929) has studied the development of the brain 
coverings in Amphibia in both normal and experimental animals and 
has come to the conclusion that in Amblystoma (taken as a type of 
Urodeles) there are three membranes investing the brain, the dura 
mater, pia mater, and arachnoid, and that the last develops from the 
inner surface of the dura. 

Thus we are faced with three competent observers describing 
respectively two, one, and three primary meninges in the adult 
Urodele. The differences between them are largely those of inter- 
pretation of homologies and mode of origin, since each admits three 
membranes. 

Miss O'Neill's work was done under the supervision of Professors 
Wiedersheim and Gaupp and is the most detailed as far as Sala- 
mandra itself is concerned. It has been thought advisable therefore 
to adhere to her interpretation pending a more satisfactory and deci- 
sive solution of the whole problem. 

Following Miss O'Neill's terminology, the two membranous en- 
velopes surrounding the brain and spinal cord are: 

(i) The dura mater on the outside. A firm tough membrane, con- 
sisting of connective tissue-fibres with flattened cellular elements 
between them. It is slightly pigmented, the pigment cells being of 
the stellate variety. In a few special places, to be enumerated later, 
the dura mater splits so as to form two layers separated by a space, 
the interdural space. The outer layer, next to the bone of the cranium 
or vertebral column, is the periosteal layer \ the inner layer, lying 
nearer to the brain or spinal cord, is the neural layer. 

(ii) The inner envelope or primary vascular coat, which carries 
the blood-vessels. It lies next to the neural tissue itself and 
corresponds with the pia mater of most authors. According to 
Miss O'Neill, two layers are distinguishable: namely, an inner, 
more or less homogeneous, firm layer in which cellular elements 



THE NERVOUS SYSTEM 125 

are scarce, and an outer alveolar layer. The blood-vessels lie 
between the two. 

The space between the dura mater and the primary vascular coat 
is called the, primary subdural space. It is not a homogeneous cavity, 
but is filled with a loose vacuolated tissue which Miss O'Neill likens 
to embryonal 'Schleimgewebe', so that the subdural space in the 
Salamander is represented by a number of small spaces rather than 
by a continuous cavity. 

Miss O'Neill considers this condition primitive, representing a 
phase earlier than the formation of a true 'arachnoid' membrane. It 
should be noted here that, owing to the shrinkage of the neural 
tissue in the processes of fixing and hardening, whether for section 
cutting or dissection, the primary subdural space is always seen more 
or less badly distorted. It is therefore difficult to determine its exact 
relations with accuracy. 

Modifications of the Envelopes surrounding the Brain. The 
splitting of the dura mater and the formation of an interdural space 
in certain regions have already been mentioned. This occurs only in 
the cranium, and does not extend into the vertebral canal. There are 
four such splits, and they are formed by: (i) the saccus endolympha- 
ticus; (ii) the saccus perilymphaticus; (iii) the pituitary body, or 
hypophysis cerebri; and (iv) the paraphysis. 

(i) Spatium interdurale endolymphaticum (O'Neill), as the first is 
called, is the largest. The saccus endolymphatkus is easily seen, on 
removing the roof of the cranium, lying at the sides of the midbrain, 
owing to the fact that it is filled with a white chalky material. It is 
connected with the ear by means of the ductus endolymphatkus^ which 
passes through the cranial wall into the auditory capsule. Both Hasse 
(1873) and Rex (1893) state that it is subdural in position, that is, 
they imagine that the ductus endolymphaticus penetrates the dura 
mater and swells out into the saccus endolyphaticus between this 
envelope and the inner vascular coat. This would put Salamandra — 
and Triton — in a peculiar position, since, in the Frog, both Coggi and 
Gaupp had shown the endolymph sac to be i«/«?rdural. Miss O'Neill 
has demonstrated that this anomaly does not exist, and that in Sala- 
mandra the endolymph sac splits the dura into two layers, and is 
therefore interdural as in the Frog.^ The dorsal and lateral walls of 
the endolymph sac are richly vascular, while the mesial wall — the 
'pars media' of Hasse — lies very close to the brain and to the plexus 
covering the fourth ventricle, so that in the region of this plexus the 
neural layer of the dura and the primary vascular coat are brought 

' For other views on the position of the saccus endolymphaticus see table on p. 128. 



126 THE SALAMANDER 

SO close together that — according to Miss O'Neill — it is not possible 
to distinguish between them. In this region also the saccus endo- 
lymphaticus is continuous right across the dorsum of the brain and 
communicates with the sac of the opposite side. ^ Thus the spatium 
interdurale is also continuous, since it is caused by, and develops pari 
passu with, the development of the saccus. Ventrally the spatium 
interdurale and the saccus terminate lateral to the pituitary body. 
The trigeminus nerve passes through the interdural space to the 
exterior, while the anterior root of the vagus and the acustico-facialis 
root leave the cranial cavity at the border between the spatium 
endolymphaticum and the spatium perilymphaticum. 

(ii) Spatium interdurale perilymphaticum (O'Neill). This inter- 
dural space resembles the former in that it is caused by a saccular out- 
growth from the auditory organ, but it is much more limited in extent. 
The saccus perilymphaticus lies within the spatium interdurale and is 
connected with the ear by the ductus perilymphaticus^ which passes 
through a special opening in the cranial wall — t]\e foramen rotundum 
of Hasse, or better, the apertura ductus perilymphatici (O'Neill). 

Hasse thought that the ductus perilymphaticus was connected 
with the inner envelope of the brain (Gehirnhulle)^ and that the peri- 
lymph flowed into the cavum epicerebrale, that is, into the space 
between the brain and the Gehirnhiille. Miss O'Neill, on the other 
hand, is convinced that this is not the case, but that the perilymph is 
confined to the saccus perilymphaticus within the interdural space. 
In this she is supported by Harrison (1902). She thinks, however, 
that there may be, here and there, some connexion between the peri- 
lymph and the endolymph where the two sacs come into close contact. 

The spatium interdurale perilymphaticum lies ventral to the spa- 
tium interdurale endolymphaticum which is laid over it like a saddle, 
extending beyond it anteriorly and posteriorly. 

(iii) Spatium interdurale hypophyseos (O'Neill). As is well known, 
the hypophysis cerebri or pituitary body develops as an evagination 
from the roof of the mouth, and finally separates entirely therefrom, 
becoming applied to the ventral surface of the brain at the infundi- 
bulum. In doing this it passes through the periosteal layer of the 
dura mater and pushes the neural layer in front of it, thus forming 
and lying within an interdural space — the spatium interdurale hypo- 
physeos. 

' Dempster (1930) states that while the two sacci endolymphatici do communicate in 
some Urodeles they do not do so in Salamandra, where they abut very closely and are 
separated only by an exceedingly thin membrane. He accepts Sterzi's view as to their 
position, but does not mention Miss O'Neill's paper. 



THE NERVOUS SYSTEM 127 

So close does the hypophysis lie to the infundibulum that Miss 
O'Neill states she is unable to detect the neural layer of the dura 
mater between them, and therefore suspects that at this point the 
interdural space and the subdural space may actually communicate 
with one another. 

(iv) Spatium interdurale paraphyseos (O'Neill). The fourth inter- 
dural space is a median dorsal one. The paraphysis is a dorsal out- 
growth from the roof of the thalamencephalon (see under 'Brain'), 
and as it comes into contact with the dura mater during its develop- 
ment, it penetrates the neural layer and spreads out between this 
layer and the periosteal layer, thus producing a space between them. 
It will be noticed that this space is produced in exactly the same 
manner as those already described, but in the reverse direction, 
namely, by a centrifugal outgrowth instead of a centripetal one. 
Since the paraphysis remains attached to the brain, the spatium 
interdurale paraphyseos retains a communication with the subdural 
space. 

The primary vascular coat of the brain entirely envelops this im- 
portant organ. It takes part in the formation of the several plexuses 
— see 'Vessels of the Brain'. 

Modifications of the Envelopes surrounding the Spinal cord. 
There is little or no splitting of the dura mater, with the consequent 
formation of an interdural space, within the vertebral column. The 
only sign of such a condition is towards the anterior end, where the 
V. post-occipitalis passes through the dura on its way out from the 
vertebral column. 

On the other hand, there are three well-defined thickenings of the 
primary vascular coat. 

Two of these, forming a pair of longitudinal bands running one 
on either side of the spinal cord between the dorsal and ventral nerve 
roots, were first described by Berger in 1878. He called each the 
Ugamentum denticulatum. They commence at the level of the second 
vertebra and continue right down the body, protruding a little dis- 
tance into the subdural space and are suspended by a reduplication 
of the vascular coat. 

The third thickening of the primary vascular coat is situated in 
the mid-ventral line beneath the sulcus longitudinalis ventralis^ and 
forms the Ugamentum piae ventrale (O'Neill). The spinal cord is 
somewhat eccentrically placed in the vertebral canal, being rather 
nearer the ventral side. This brings the ligamentum piae ventrale 
into very close connexion with the dura mater, and interrupts the 
subdural space mid-ventrally. 



128 THE SALAMANDER 

Comparison of the three interpretations of the meningeal coverings 



O'Neill {i%c)%) 


Sterzi (i 899-1909) 


Flexner (1929) 


Dura mater: 






{a) periosteal layer 


'Endorachis' 
(periosteum) 


Dura mater 


{b) neural layer 


Meninx primitiva 


Arachnoid 


Interdural space 


Epidural space 


Subdural space 


Primary vascular coat 


Meninx secundaria 


Pia mater 


Saccus endolymphaticus 


Saccus endolymphaticus 


Saccus endolymphaticus 


z»/^r-dural 


epi-dnrdil 


sub-dMXdX 



3. Brain {gross anatomy only) (PI. IX). 

The brain of the Salamander is very long compared with its 
breadth and depth — a feature typical of Urodeles in general. Its 
length, from the anterior end of the hemispheres to the origin of the 
first spinal nerve, is approximately 14 mm.; its breadth, measured 
across the posterior part of the hemispheres, is about 4 mm.; while 
its depth, from the roof of the mesencephalon to the infundibulum, 
is only some 3I mm. 

In dorsal v'lQw (Fig. 51), the following features are conspicuous: 

(i) The olfactory lobes. These are not fused into a median, unpaired 
body as in the Frog, but are lateral structures which merge, without 
any sharp line of demarcation, into 

(ii) The hemispheres of the prosencephalon^ or telencephalon. These 
are long, cylindrical bodies with rounded ends. Together with the 
olfactory lobes they form about 37-5 per cent, of the total length of 
the brain, their diameter being greatest at about two-thirds their 
length from the anterior end. They lie very close to each other 
anteriorly, but their posterior ends turn slightly outwards from the 
middle line. The telencephalon is followed posteriorly by: 

(iii) The diencephalon. The anterior portion of this section — the 
thalamencephalon — is scarcely visible from the dorsal side, being 
covered by the triangular nodus chorioideus and its associated para- 
physis. The paraphysis develops from the anterior wall of the thala- 
mencephalon as a dorsal evagination and becomes associated with a 
rich vascular plexus, forming the nodus chorioideus. In the adult the 
tissue of the paraphysis is largely confined to the posterior region of 
the plexus. It does not penetrate the cranium but is flattened out 
against its roof. The anterior portion of the roof of the diencephalon 
appears at first sight to possess a median cleft. Close inspection, how- 
ever, reveals a weak bridge of nervous material, the hahenular com- 



THE NERVOUS SYSTEM 129 

missure^ followed by a clearer area — the pulvinar. It is from the 
posterior end of this region that the epiphysis (epi.) and its terminal 
pineal body evaginate. They are only weakly developed in the Sala- 
mander and never perforate the cranial roof, but flatten out over the 
surface of the brain between it and the parietals. In the adult the 
epiphysis becomes very closely applied to the surface of the brain, 
but is distinguishable in sections (see Fig. 50). Eycleshymer (i 892) 
states that the paraphysis and epiphysis in Urodela never come into 
close relation as in Reptilia, but remain widely separated. The condi- 
tion in the Salamander supports this statement. 

(iv) The corpus opticus (lob.op.) or mesencephalon follows. It is not 
divided into two distinct lateral lobes as in the Frog, but expands 
gradually from the roof of the diencephalon into a more or less hemi- 
spherical dome. There is a faint groove in the middle line and the 
nervous tissue is thinner there than elsewhere, this feature being more 
obvious in transverse sections. The corpus opticum is far removed 
from the cerebrum. At the posterior border of the mesencephalon 
there is a deep groove between this and the subsequent portion of 
the brain, and from this groove the N. trochlearis (fourth cranial 
nerve) emerges. 

(v) The cerebellum (cerb.) or epencephalon is a very narrow trans- 
verse band of nervous tissue forming the anterior boundary of 
the fourth ventricle, and separated from the mesencephalon by the 
groove above mentioned. This region is surprisingly slender, being 
much smaller comparatively than the corresponding region in the 
Frog. 

(vi) The medulla oblongata (med.ob.) or myelencephalon is a fairly 
long (about 6| mm.) shallow trough of nervous tissue, open dorsally. 
It forms about 42 per cent, of the total brain length. Its anterior end 
is laterally expanded, and tapers gradually into a more or less circular 
cross-section posteriorly, merging imperceptibly into the spinal cord. 
Its cavity — the fourth ventricle — is roofed over dorsally by a rich 
vascular plexus. The trigeminus (Vth) root arises ventro-laterally 
from its anterior angles, and the acustico-facialis (Vllth and Vlllth) 
root appears about | mm. posterior to this. The glossopharyngeus- 
vagus complex (IX and X) arises laterally from the sides of the 
medulla by three roots, of which the middle root is the largest, the 
other two being very fine. 

The following additional features are evident on the ventral sur- 
face of the brain (Fig. 52): 

(i) The emergence of the olfactory nerves (n.i) from the brain. 

4038 K; 



130 THE SALAMANDER 

They are not quite terminal, but arise ventrally a short distance 
from the anterior end of the prosencephalon. 

(ii) The floor of the thalamencephalon (the.) fits in as a wedge 
between the hemispheres, which are to be regarded as lateral out- 
growths from it.^ The optic nerves (n.2) pass from its floor at about 
the level of the posterior ends of the hemispheres. 

(iii) The injundibulum (inf.) is an outgrowth from the floor of the 
diencephalon which extends in a postero-ventral direction so as almost 
to obscure the floor of the mesencephalon. Its posterior end is some- 
what expanded laterally and has a fairly deep median notch. Trans- 
verse sections show that the neural tissue is almost entirely confined 
to the ventral surface, its roof being very thin and membranous. In 
sagittal section it appears as a hollow wedge. From the posterior 
angles of the median cleft the pituitary body is suspended. 

(iv) The pituitary body (pit.), or hypophysis cerebri, is roughly 
a broad oval in surface view, with a median notch anteriorly. The 
lobes at the sides of the notch are distinguished as the pars tuheralis 
(Atwell) (pit.'"), while the main mass is the/>^rj anterior. According 
to Sumi the pars tuheralis arises rapidly at about the time of metamor- 
phosis. In many Urodeles it becomes completely detached from the 
pars anterior, but in Salamandra its separation is only partial. Both 
the pituitary and the infundibulum lie within the basi-cranial fonta- 
nelle of the primary chondrocranium, and are covered ventrally by 
the parasphenoid bone. 

(v) The ventral surface of the medulla calls for no special comment 
save that the roots of the trigeminus and acustico-jacialis are here more 
distinct, and the A^. abducens (VI) is also visible. It arises on either side, 
very close to the middle line, by two roots about a millimetre posterior 
to the acustico-facialis. It is an exceedingly fine nerve and readily 
broken, and even when intact it is not easy to see. 

In lateral view it should be noted that there is only a very slight 
departure from the straight unflexed condition of the embryonic 
brain. 

By direct observation of the sagittal section (Fig. 50) it is not 
possible to make out more than the main features. If the half of the 
brain is immersed in a solution of ferric chloride the iron salt will be 
found to adhere more readily to the grey matter than to the white, 
and thus render the outlines of the cavities more distinct. The posi- 
tions of the commissures have been confirmed, and in some cases 
determined, by the help of transverse sections examined micro- 
scopically. 

' Cf. Graham Kerr on the brain of Lepidosiren, Q.J.M.S., London, vol. 46, p. 448. 



THE NERVOUS SYSTEM 131 

In addition to the commissures and the thickness of the walls of 
the brain in its various parts, the chief features of interest in a 
sagittal section are: 

(i) The. foramen of Monro (fo.M.) connecting the lateral ventricles 
— the cavities of the cerebral hemispheres — with the third ventricle 
— the cavity of the thalamencephalon. This is a long oval slit 
of relatively large dimensions, and through it the blood-vessels 
pass from the nodus chorioideus to the lateral plexus of the first 
ventricle. 

(ii) The remainder of the pituitary body may now be dis- 
tinguished: 

(a) The pars intermedia (Atwell) (pit/). This is very small, 
and lies dorsal to the pars anterior and behind the infundi- 
bulum. 

(Jy) The pars anterior (pit.") is bi-convex in section. It forms by 
far the largest portion of the gland. 

[c] The infundibular portion, or pars posterior^ shows considerable 
retrogression and is almost non-existent. 

(iii) The purely epithelial portions of the brain surface are as 
follows : 

{a) The lamina terminalis (lam.ter.) and antero-dorsal roof of the 
thalamencephalon. 

{h) The dorsal wall of the infundibulum. 

{c) The posterior median portion of the mesencephalon. 

{d) The roof of the fourth ventricle. 

With regard to the commissures on the floor of the foramen of 
Monro, the upper bundle was first homologized with the corpus 
callosum by Osborn (1886). It had previously been thought to be 
homologous with the pars olfactoria of the anterior commissure, but 
Osborn showed that this part was present in the lower bundle of 
fibres as well as the pars temporalis^ and thus the lower bundle is the 
entire anterior commissure. 

Summary. 

The brain of the Salamander is straight, or very slightly flexed, 
and small compared with that of the Frog, but long in comparison 
with its breadth and depth. The pineal organ is much reduced, 
while the paraphysis shows considerable development. The sepa- 
ration of the optic lobes is very incomplete and the cerebellum 
poorly developed. The olfactory lobes are paired and not fused 
to form a median body as in the Frog. The foramen of Monro 
is large. 



132 THE SALAMANDER 

4. Spinal cord. 

The medulla oblongata merges imperceptibly into the spinal cord^ 
which passes right along the body to the tip of the tail, passing 
through the neural canal of the vertebral column. It remains fairly 
uniform in diameter as far as the sacral region, where a slight 
swelling is noticeable in that part of the cord giving off the sacral 
nerves. From this point to the tip of the tail the cord gradually 
diminishes in size. 

Segmental nerves are given off corresponding with each vertebra, 
and arise, with the exception of the first, by both dorsal and ventral 
roots. The details of their course and distribution will be considered 
in a subsequent section. 

In transverse section the spinal cord is eccentrically placed in 
the neural canal towards the ventral side. It is approximately oval 
in shape with a small central canal lined by epithelium, around 
which is grouped the grey matter. The white matter lies towards 
the periphery of the cord. There is a well-marked V-shaped groove 
dorsally — the dorsal fissure or sulcus longitudinalis dorsalis. It is 
shallow, however, and does not extend far in a ventral direction. 
The ventral fissure^ or sulcus longitudinalis ventralis^ is much deeper 
and reaches nearly to the central canal. It is particularly noteworthy 
that the A. spinalis ventralis lies well within the ventral fissure, thus 
tending to open it still farther. 



CRANIAL NERVES (PI. X) 

I. Historical. 

The earlier workers, e.g. Funk and Rusconi, have very little to 
say with regard to the cranial nerves. In 1 840 Vogt gave an account 
of the vagus nerve and hypoglossal which, however, is not very accu- 
rate. Three years later Fischer ( 1 843) published an adequate descrip- 
tion of all the cranial nerves except the IVth, which he failed to find. 
He did not observe the branch from the oculomotorius to the M. 
rectus superior — in fact he states definitely that it does not exist. He 
also omits any description of the branchial branches of the Xth nerve. 
Apart from these omissions he is surprisingly accurate, and is fre- 
quently more correct than some of his successors. In the same year 
Bendz (1843) published an account of nerves IX, X, and hypo- 
glossus, but his work contains a number of errors and is not so 
accurate as that of Fischer. 

In 1 89 1 von Plessen and Rabinowicz gave a description of the 



THE NERVOUS SYSTEM 133 

cranial nerves of the larval Salamander based on the study of serial 
sections, and produced elaborate figures reconstructed from these 
sections. While these authors rectify Fischer's defective account of 
the N. oculomotorius they have nevertheless failed to identify the 
lateralis system which exists in the larval Salamander. The anterior 
ganglion of this system lies just dorsal to the Gasserian ganglion. 
They therefore regard it as an accessory trigeminal ganglion and 
describe the nerves emerging from it as trigeminal nerves. This has 
naturally led to considerable confusion of nomenclature and descrip- 
tion, and a similar confusion exists between the lateralis nerve and 
its course in the vago-glossopharyngeal ganglionic complex. 

These authors also describe and figure an anastomosis between 
the R. palatinus VII and their R. supramaxillaris superior of the Vth 
nerve, which does not exist, while they fail to mention the one which 
does occur between the palatine branch of the Vllth and the ventral 
branch of R. ophthalmicus profundus V. 

The confusion with regard to the lateralis system was cleared up 
by Strong in 1895, but this author is himself in error when he sug- 
gests that the anastomosis between the Vth and Vllth nerves should 
be between R. palatinus VII and R. maxillaris V. He says: Tn 
Wiedersheim's Grundriss this connexion is in one place, by some 
error, spoken of as between R. palatinus and R. ophthalmicus pro- 
fundus.' Actually Wiedersheim is quite right, the difference between 
the topographical relations of this anastomosis constituting an impor- 
tant distinction between the Urodela and Anura. In 1 896 Kingsley 
re-examined the question of this anastomosis by means of sections, 
and shows the true communication to be between the palatine and 
the ophthalmicus profundus, and suggests further that von Plessen 
and Rabinowicz mistook the M. obliquus inferior for an anasto- 
mosis between a branch of the palatine and their R. supramaxillaris 
superior V. 

In 1 90 1 Driiner gave an excellent account, illustrated by a number 
of fine figures, of the visceral nerves (VII, IX, and X), and of the 
muscles they supply. His work is very accurate. 

A year later Hoffmann published a general account of the cranial 
nerves of Salamandra and Triton, both larval and adult. His work 
in respect of the visceral nerves is not so detailed as that of Driiner. 
While on the whole Hoffmann's account is accurate, his text is not 
very easy to follow, probably on account of the absence of any dia- 
grams or figures other than those of microscopic sections. Frequent 
reference to both these papers will be made in the ensuing pages. At 
about the same time several American anatomists began to investigate 



134 THE SALAMANDER 

the cranial nerves of their native Urodeles by means of serial sections, 
paying particular regard to the distribution of their several compo- 
nents. These accounts, notably that of Coghill (1902) for Ambly- 
stoma, have been found to be of considerable use in unravelling 
the intricacies of the nerve relations in the head of the Salamander. 
After making due allowance for the absence of a lateralis component 
in the adult Salamander, the correspondence between the cranial 
nerves of this animal and those of Amblystoma is very close 
indeed. 

The present account is based almost entirely on observations made 
by actual dissection carried out under a binocular dissecting micro- 
scope — checked in a few cases by microtome sections. No attempt 
has been made to elucidate the components of the various nerves, but 
these may be deduced by analogy from the descriptions of American 
Urodeles above referred to, and the composition of each nerve thus 
deduced has been given in parenthesis. It should be noted further 
that where the words 'origin' and 'root' are used they refer only to 
the point of emergence of the nerve from the surface of the brain and 
not to the deeper origin within it. 

All cranial nerves are paired, and those of one side only are 
described. 

2. Description. 

I. N. Olfactorius (n.i). The first cranial nerve leaves the 
olfactory lobe of the prosencephalon from its ventro-lateral border 
and passes directly forwards, through the fenestra olfactoria, into the 
nasal capsule. It is a flat oval in transverse section. Almost imme- 
diately after separating from the brain the nerve gives off a branch 
from its ventral side which turns sharply outwards towards the 
internal narial opening, and apparently supplies Jacobson's gland. 
The main olfactory nerve then divides into a dorsal and ventral 
branch, each of which ramifies over its respective surface of the nasal 
sac, supplying the olfactory epithelium. 

II. N. Opticus (n.2). The optic nerve passes from the floor of the 
thalamencephalon and is inclined slightly anteriorly and dorsally. 
It leaves the cranial cavity by the optic foramen in the side wall of the 
cranium. Since the nerve occupies only a very small part of this 
'foramen' it would perhaps be more correct to call the latter the optic 
fenestra. The nerve is protected within the fenestra, and during 
its passage from the side of the skull to the bulbus oculi, by a 
strong, fibrous, connective tissue sheath, the whole structure 
forming the optic peduncle. 



THE NERVOUS SYSTEM 135 

Eye-muscle Nerves (PI. XIII, fig. 57). 

In addition to the general accounts already referred to, Schwalbe 
in 1879 published a description of the eye-muscle and ciliary nerves. 
He, like Fischer, fails to find the oculomotor branch to the M. rectus 
superior, but he disagrees with him in that he does sometimes find 
a N. trochlearis to be present. He is the first to give an account of 
the ciliary nerves, both superior and inferior, although he is a little 
uncertain of the exact relations of the latter. 

Owing partly to the small size of the animal, and partly to the 
rather large amount of individual variation met with, the investiga- 
tion of these nerves in the Salamander is by no means easy, but with 
care they may be followed fairly satisfactorily with the aid of a bino- 
cular dissecting microscope. 

III. N. Oculomotorius (n.3) (somatic motor). The IlIrd nerve 
leaves the brain from the floor of the mesencephalon and passes out 
of the cranial cavity through a special foramen — For. oculomotorium 
— in the side wall of the cranium, just posterior to the optic fenestra. 
Within the orbit it divides into two rami. 

(a) A ramus superior which turns dorsalwards and enters the M. 
rectus superior along its anterior border. 

(F) A ramus inferior which passes ventralwards, posterior to the 
optic nerve, between it and the M. retractor bulbi, and close to a 
branch of the Vth nerve. It divides into three branches, going respec- 
tively to the inferior and anterior rectus muscles, and to the inferior 
oblique muscle. Throughout its course it keeps close to the trige- 
minal branch above mentioned, and apparently passes across, or 
through it, above the M. rectus inferior. 

(c) Close to the point where the N. oculomotorius divides into 
superior and inferior rami a ramus communicans to the R. ophthalmicus 
profundus V may be found (n.3-5), and usually at about the same 
point a slight swelling of the nerve represents the ciliary ganglion. 
Schwalbe (i 879) reports several patches of ganglionic cells along the 
course of the ramus inferior, of which the largest is in the position 
indicated above. Hoffmann (1902) also confirms the presence of 
ganglia associated with the oculomotor nerve, but says that they sur- 
round the nerve like a sheath and are not within the nerve itself, while 
he considers them to be of a sympathetic nature and to represent the 
most anterior ganglion of this system. Coghill finds in Amblystoma 
that the ciliary ganglion is transitory, and tends to disappear in the 
adult. Hence he concludes that it is at no time functional. 

It is certain, however, that a ganglion does normally occur associated 



136 THE SALAMANDER 

with the oculomotor nerve in the Salamander, and it is probably of 
a sympathetic character, and is not connected with the fibres of the 
nerve itself. The present investigation has, however, not revealed 
its exact relations (cf, p. 183). 

(d) Both Schwalbe and von Plessen and Rabinowicz state that the 
oculomotor nerve sends a branch into the M. retractor bulbi. Now 
while a twig has been found which leaves the nerve together with, 
or close to, the ramus communicans ad V, passing amongst the fibres 
of the above-named muscle towards the bulbus, it is improbable 
that it terminates within the muscle, but seems rather to pass to the 
eye itself and enter the sheath surrounding the optic nerve. Gaupp 
reports a similar twig in the Frog. It may be that this nerve is of a 
sympathetic nature and is associated with the ganglion above re- 
ferred to, but on this point there is no definite evidence. 

The above account agrees fairly closely with Coghill's 'Case A' 
and appears to represent the most usual arrangement, although indi- 
vidual variations are quite common. They chiefly concern the mode 
of branching, and the relations between the ramus inferior and the 
trigeminus branch, and the exact position of the ciliary ganglion. 

IV. N. Trochlearis (n.4) (somatic motor+sensory fibres?). The 
exact relations of this nerve are exceedingly difficult to determine by 
dissection, partly on account of the fineness of the nerve itself and 
the toughness of the tissue in which it is embedded, and partly on 
account of its great variability, even on opposite sides of the same 
animal. The discrepancies between the several published accounts 
testify to this. 

As above mentioned, Schwalbe was the first to recognize its exis- 
tence, but in spite of his account von Plessen and Rabinowicz state 
that the M. obliquus superior is innervated by the trigeminus. Hoff- 
mann (1902) gives a detailed description of the nerve, but describes 
two anastomoses with the Vth nerve, whereas only one has been found. 

So far as the present author has been able to observe, the most 
usual arrangement is as follows. The nerve leaves the brain near the 
middle line immediately in front of the cerebellum, emerging on the 
dorsal side from the deep groove which exists between this part of 
the brain and the corpus opticus. The nerve passes antero-laterally 
through the cranial cavity, and in doing so is often closely applied to 
the under-surface of the parietal bone, so that great care must be 
exercised in removing the roof of the skull to avoid damaging the 
nerve. It emerges from the cavum cranii through a very oblique 
foramen in the parietal, or between the parietal and the orbito- 



THE NERVOUS SYSTEM 137 

sphenoid (cf. Gaupp, 1 9 1 1), at about the level, or just in front of, the 
optic nerve and dorsal to it. The nerve may divide within the skull 
and pass through its wall as a double nerve, otherwise it divides 
immediately on emerging from its foramen, and before penetrating 
the origin of the M. levator bulbi. The division is unequal, one 
branch being very much finer than the other. The finer branch (IVb, 
Hoffmann) passes at once through the origin of the M. levator 
bulbi, and goes to the M. obliquus superior without anastomosing 
with any branch of the trigeminal nerve. It may come into contact 
with such a branch but it is comparatively easily separable from it, 
and no exchange of fibres appears to take place. The larger branch 
usually turns a little anteriorly before penetrating the origin of the 
M. lev. bulbi, and after it has emerged from this muscle it always 
comes into contact, and apparently anastomoses with, one or more 
cutaneous twigs of the R. ophthalmicus profundus V. It may divide 
into several twigs, some of which anastomose with branches of the 
Vth nerve while others pass to the M. obi. sup., or it may anastomose 
as a whole with a branch from the trigeminus, in which case fibres 
arise from the combined nerve and enter the superior oblique muscle. 

The above description agrees fairly well with that given by Hoff- 
mann, except that no anastomosis between the trochlearis and the 
trigeminus ?7iesial to the M. lev. bulbi such as he describes has been 
found, neither in fact has any branch of the Vth nerve been found 
which could fuse with it in this position. It should be noted here that 
Coghill (1902) describes the trochlearis of Amblystoma meeting a 
trigeminal branch mesial to the M. lev. bulbi. He says, 'I find no 
positive evidence of an anastomosis in this case although the nerves 
are in very intimate relation with each other. The fourth nerve is 
penetrated by the trigeminal twig and then passes through the M. 
lev. bulbi and innervates the M. obliquus superior.' This author, 
however, does not describe any fusion of the two nerves lateral to the 
M. lev. bulbi, nor any division of the trochlearis into two distinct 
rami such as occurs in Salamandra. 

From the above details the following facts emerge: 

(i) A N. trochlearis does occur in Salamandra and the M. obliquus 
superior is innervated by it. 

(ii) The whole of the fourth cranial nerve is not distributed to the 
eye muscle, but some part of it goes to the skin or underlying tissues 
via the track of a trigeminal twig.^ 

' If Hoffmann's account is correct these apparently cutaneous branches of the troch- 
learis may actually come from the fifth twig, which, according to him, joins the troch- 
learis mesial to the M. levator bulbi. 



138 THE SALAMANDER 

(iii) The trochlearis normally divides into two distinct rami, one 
of which anastomoses with the Vth nerve while the other does 
not. 

Before the significance of these facts can be adequately appreciated 
a complete reinvestigation of the internal origin and development of 
this nerve would be necessary. 

VI. N. Abducens (n.6) (somatic motor). This is one of the finest 
of all the cranial nerves and one of the most difficult to investigate. 
It arises by two roots from the ventral surface of the medulla very 
far back — in fact posterior to the origin of the IX-X complex — 
close to the mid-ventral line, and passes anteriorly and laterally 
alongside the internal carotid artery, leaving the cranial cavity by a 
special foramen — For. abducentis — just a little antero-mesial to the 
palatine nerve. It enters the orbit mesial to the M. retractor bulbi. 
In the adult Salamander the N. abducens is always quite distinct 
throughout the whole of its course from the trigeminus, and from 
the Gasserian ganglion, although, according to Hoffmann, it 
approaches the latter fairly closely in the larva, while in some Uro- 
deles, e.g. Triton, it actually enters the ganglion and leaves it in 
company with true trigeminus fibres. It supplies the MM. retractor 
bulbi and rectus posterior, and in order to reach the latter muscle 
the nerve frequently bifurcates and passes round either side of the 
M. retractor bulbi, supplying it with numerous fibres on the way, 
or it may remain entire and pass round the lateral aspect of the 
retractor muscle. 

So far as the present investigation goes, the MM. retractor bulbi 
and rectus posterior of the Salamander are only supplied by the 
abducent nerve which is distributed to these muscles alone. 

V. N. Trigeminus (n.5). The Vth nerve of the Salamander is 
probably the most complicated of the cranial nerves. It possesses 
the typical three branches which, however, are not quite typical in 
composition, as will be seen later. The nerve arises from the 
antero-lateral angle of the medulla, and passes immediately into 
the. foramen ■pro-oticum situated just in front of the auditory capsule 
which is formed by the junction of the trabeculae with the roof 
of the ear capsules. Within this foramen lies the large Gasserian 
ganglion from which the branches of the trigeminus nerve arise. 

Von Plessen and Rabinowicz describe two ganglia in connexion 
with the Vth nerve; (i) an accessory (Neben-) ganglion associated 
with a dorsal root, and (ii) a main (Haupt-) ganglion with a ventral 
root. Now their 'Nebenganglion' is really the ganglion belonging 



THE NERVOUS SYSTEM 139 

to the lateralis system and has nothing to do with the trigeminus 
system at all, it being a larval structure which disappears at meta- 
morphosis. The only trace of this system of nerves to be found in 
the adult is a very fine connexion between the acustico-facialis gang- 
lion and the Gasserian ganglion — the remains of the Radix dorsalis 
of von Plessen and Rabinowicz — the whole of the ganglion with its 
R. ophthalmicus superficialis VII (R. frontalis, v. P. and R.), R. 
buccalis (R. supramaxillaris superior, v. P. and R.), and R. mandi- 
bularis externus (R. buccalis, v. P. and R.) having atrophied, with the 
possible exception of a few fibres distributed to the skin above the 
M. levator mandibulae which may represent the remains of the R. 
ophthal. super. VII (cf. Driiner, 1901, p. 537). 

We may now consider the three main rami of the Vth nerve. 

Vj. R. ophthalmicus profundus (m.5a) (general cutaneous). (See 
also PL XIII, fig. 57.) 

R. nasalis . ..... von Plessen and Rabinowicz. 

R. ophthalmicus ........ Hoffmann. 

Comparing this ramus with the typical Selachian arrangement, it 
probably represents both the ophthalmicus profundus and at least 
a portion of the ophthalmicus superficialis V of that group, but since 
almost the whole of the nerve passes ventral to the M. rectus sup. it 
seems justifiable to apply the term 'profundus' to it. 

After leaving the ganglion it passes out from the For. pro-oticum, 
lateral to the orbito-sphenoid cartilage s. trabecula, but mesial to the 
ascending process of the quadrate. (It is thus separated by this pro- 
cess from the other two trigeminal branches.) It then passes directly 
anteriorly, mesial to the levator muscles of the mandible, between 
the R. sup. and the R. inf. of the N. oculomotorius, ventral to the 
M. rectus superior, dorsal to the optic nerve, and between the eyeball 
and the skull. During its passage through the orbit it gives of? 
several — usually three — branches to the skin on the dorsum of the 
head. There is a great deal of individual variation in the precise 
arrangement of these branches, but the following seems to be the one 
most generally met with. It differs slightly from that given by Hoff- 
mann, but the difference is more apparent than real, as this author 
seems to be describing one particular case as if it were a rigid plan 
adhered to by all individuals. 

(i) The first branch (V^^'+V^^\ Hoffmann) probably represents 
the R. temporalis superior (Gaupp) of the Frog. It arises from the 
main stem before it crosses the N. oculomotorius. It passes dorsal 



I40 THE SALAMANDER 

to the M. rectus superior, and immediately above the muscle it gives 

off:— 

(a) The superior ciliary nerve (n.cil.s.) — a fairly large branch which 
either passes over, or through, the muscle and penetrates the sclerotic 
coat of the eyeball. This would appear to be the V^""' of Hoff- 
mann. 

(J?) The rest of the nerve — Hoffmann's V^^' — which is distributed 
to the skin overlying the postero-mesial portion of the orbit. 

It would seem to be this nerve that the earlier authors described 
as supplying the M. rectus superior. 

(ii) Just ventral to the superior rectus muscle a fairly large branch 
arises which almost immediately divides into two — Hoffmann's 
V^^\ Both branches pass dorsal to the M. obliquus superior and are 
distributed to the skin covering the fronto-ethmoidal region, and 
thus would appear to be the equivalent of the RR. frontales et palpe- 
brales superiores anteriores (Gaupp) of the Frog. 

These branches very frequently receive an anastomosis from the 
previous branch, which passes ventral to the M. rectus superior. 

(iii) From the ventral side of the main profundus nerve, just pos- 
terior to the optic nerve, the inferior ciliary nerve (n.cil.i.) is given off. 
It passes between the optic nerve and the M. retractor bulbi and 
receives (or gives T) a fine anastomosis from the oculomotor nerve. 
With this anastomosis are associated both the nerve which runs 
parallel with the optic nerve to enter the eyeball (see also under N. 
oculomotorius) and the ciliary ganglion. 

The inferior ciliary nerve then proceeds lateralwards together with 
the oculomotor, keeping dorsal to the M. rectus inferior. The nerves 
cross one another above this muscle, and the inferior ciliary nerve 
continues in a lateral direction, penetrates the muscle to the ventral 
side, and then enters the sclerotic coat of the eyeball near the inser- 
tion of the muscle. 

Just after penetrating the muscle it gives off a branch which passes 
anteriorly over the floor of the orbit, and is distributed to a small 
section of the skin of the upper jaw between the most anterior branch 
of the maxillary nerve and the most posterior twig of the lateral ter- 
minal branch of the profundus. 

Terminal branches of the profundus (V^*" + V^"^, Hoffmann). 

A little anterior to the point where it crosses the optic nerve the 
ophthalmicus profundus divides into three main terminal branches — 
mesial, lateral, and ventral. These may anastomose variously with 
one another until the anterior limit of the orbit is reached. They all 



THE NERVOUS SYSTEM 141 

pass ventral to the M. obliquus superior, and dorsal to the M. obli- 
quus inferior. 

(iv) Mesial branch (V^'', Hoffmann). The mesial terminal branch 
(n.5a.M.) passes through the Foramen orbito-nasale mediale in the 
antorbital cartilage slightly dorsal to the origin of the M. obliquus 
superior. On entering the nasal capsule the nerve turns mesially and 
then anteriorly, passing over the nasal sac dorsal to the olfactory 
nerve. At a point a little posterior to the level of the external nares 
it breaks up into several fine terminal twigs which penetrate foramina 
in the premaxilla and the nasal bone, and supply the skin of the snout. 

Just before this branch enters the For. orbito-nasale mediale, 
while it is still within the orbit, it gives a small lateral branch which 
fuses with a twig from the lateral terminal branch and then enters 
the nasal capsule separately from the main nerve. It passes dorsal to 
the nasal sac, and turns first mesially, then sharply laterally, and 
finally perforates the nasal capsule laterally to suppy the skin near 
the ductus naso-lacrimalis. 

The branch just described is composed of fibres from both lateral 
and mesial terminal branches in varying proportions, and although 
its course is somewhat peculiar its presence seems to be reasonably 
constant. 

(v) Lateral branch (V^"^', Hoffmann). The lateral terminal branch 
(n.5a.L.) gives off" one or two small twigs within the orbit which turn 
sharply lateralwards, and supply the skin over the upper jaw near the 
anterior angle of the orbit. The main nerve then itself turns lateral- 
wards, penetrates the For. orbito-nasale laterale, and enters the pos- 
tero-lateral corner of the nasal capsule. It crosses the corner of the 
capsule obliquely, more or less lateral to the olfactory epithelium, 
and passes through the fenestra infraconchalis and then through 
foramina in the maxilla to supply the skin of the face laterally of the 
nasal capsule. 

(y\) Ventral branch(y'^^\¥Lo&m2inn). The ventral terminal branch 
(n.5a.V.) enters the nasal capsule by the For. orbito-nasale mediale 
directly between the origins of the superior and inferior oblique 
muscles, a little ventral to the mesial terminal branch, and close to 
the floor of the capsule. After passing through the foramen the 
nerve remains entirely ventral to the olfactory epithelium and, turn- 
ing slightly mesially, anastomoses with the R. palatinus VII just 
mesial to the internal narial opening. In sections ganglionic cells are 
discernible at the point where these two nerves join, and even in dis- 
sections a swelling suggestive of a ganglion is noticeable. The nerves 
resulting from this fusion are of course mixed, and contain both 



142 THE SALAMANDER 

trigeminus and facialis fibres, so that it is not possible to determine 
the precise distribution of each component by dissection. In general 
they spread over the roof of the mouth under the nasal organs (see 
also description of R. palatinus VII). 

V2. R. maxillaris (n.5b.) (mixed, general cutaneous, and visceral 

motor). 

R. supramaxillaris inferior , . . von Plessen and Rabinowicz. 

R. maxillaris superior ....... Hoffmann. 

(Jn the larva this nerve is combined with a lateral line nerve — R. buccalis 
VII — to form the truncus infra-orbitalis ,^ 

After separating from the Gasserian ganglion the maxillary nerve 
enters the antrum petrosum laterale (Driiner) from which it emerges 
between the processus ascendens and the processus oticus of the 
quadrate, and passes, mesial to the M. levator mandibulae externus, 
to the posterior angle of the eye. Here it divides into : 

(i) The superior palpebral nerve (n.s.palp.), (R. dorsalis, Hoffmann), 
('accessory twigs' of the trigeminus, Coghill), which consists of one or 
two fairly strong branches supplying the skin of the upper eyelid and 
also part of that covering the levator mandibulae muscle and ear- 
capsule. The terminal twigs from these nerves appear to anastomose 
with those from the superficial branches of the ophthalmicus pro- 
fundus V. 

(ii) An inferior palpebral nerve (n.i.palp.) (with the remainder of 
the nerve = R. ventralis, Hoffmann) which supplies the skin of the 
lower eyelid. 

(iii) A very fine motor branch to the M. levator bulbi which 
usually arises between the two palpebral nerves. 

(iv) The maxillaris nerve (n.max.) proper, of mixed constitution. 
This nerve passes round the orbit somewhat lateral to the previous 
branch, and gives off sundry twigs as follows : 

(a) From its mesial side, one or two motor twigs to the M. levator 
bulbi. 

{b) About three lateral branches which pass successively to the skin 
of the upper jaw as far as the anterior angle of the orbit. 

Hoffmann describes an anastomosis between the latter branch and 
the R. palatinus VII, but no sign of such a fusion has been found — a 
result which is in harmony with the findings of Kingsley (1896). 
Hoffmann also suggests that the superior palpebral branch (his R. 
dorsalis) is the representative of the Selachian ophthalmicus super- 
ficialis. While it may possibly represent a portion of this nerve, 
it should nevertheless be noted that it emerges from the antrum 



THE NERVOUS SYSTEM 143 

petrosum laterale by a different opening from the ophthalmicus 
profundus, and is separated from it by the processus ascendens of 
the quadrate. 

The homologous comparison of the ophthalmicus profundus and 
maxillaris nerves of Urodeles with those of Selachians and Anura 
is not very easy. On the whole it seems most likely that Coghill's 
explanation is correct, and that the Urodele ophthalmicus profundus 
contains at least a considerable proportion of the fibres forming the 
Anuran maxillaris, thus accounting for the apparent difference in 
the trigemino-palatine anastomosis in these two groups. The Uro- 
dele maxillaris would then be the equivalent of Strong's 'accessory 
ramus' of the Anuran tadpole. This hypothesis also explains the size 
and complexity of the Urodelan profundus nerve, and the smallness 
of the maxillaris. 

V3. R. mandibularis (n.5c.) (mixed, general cutaneous, and visceral 
motor). 

R. mandibularis (partim) . . . von Plessen and Rabinowicz. 

R. maxillaris inferior ....... Hoffmann. 

The remaining trigeminal branch leaves the antrum petrosum 
laterale by the same opening as the R. maxillaris — i.e. between the 
ascending and otic processes of the quadrate, and immediately 
turns ventrally towards the lower jaw. It is flat and ribbon-like in 
shape as it passes through the levator mandibulae muscle. 

(i) Its first branch is a fine twig which turns mesially and dorsally, 
and innervates the M. levator mandibulae anterior — both superficial 
and deep portions. This branch leaves the nerve either just before 
or just after it emerges from the antrum petrosum laterale. The MM. 
levator mandibulae posterior and externus receive several very fine 
twigs from the main nerve as it passes ventralwards through them 
on its way to the lower jaw. 

On reaching the mandible, just anterior to its articulation with the 
suspensorium, the nerve divides into two branches: — 

(ii) A cutaneous branch (cut.) which turns posteriorly and supplies 
the skin overlying the angle of the jaw. From this branch is given off 
an external mandibular nerve — a very fine twig running parallel 
with the lower jaw and supplying the skin overlying the posterior 
two-thirds of the mandible. 

(iii) The other main branch is a mixed nerve, and turns antero- 
ventrally, passing right through the mandible between the dentary 
and Meckel's cartilage. Within the jaw it gives off: 

{a) An internal mandibular^ or inferior dental nerve (n.m.i.) which 



144 THE SALAMANDER 

passes anteriorly in the alveolar canal of the lower jaw, latero-dorsal 
to Meckel's cartilage. It gives fine twigs to the teeth, while its ter- 
minal branches pass out laterally through foramina in the dentary 
to supply the skin covering the anterior part of the lower jaw, and 
may thus be called RR. mentales. It is important to notice that this 
branch throughout the whole of its course remains lateral to the 
primitive jaw. Stadtmuller (1924) describes a rather complicated 
anastomosing system between it and the R. alveolaris VII, which, 
however, is only traceable by means of sections. 

(^)The main branch (n.m.e.) which passes right through the jaw 
and then turns anteriorly along its ventral side. It is a mixed nerve 
and supplies cutaneous branches to the skin between the rami of the 
mandible, and motor branches to the M. intermandibularis, while 
very occasionally its most posterior twigs may encroach on the region 
belonging to the Vllth nerve and enter the most anterior portion 
of the M. interhyoideus (cf. Driiner, 1901, p. 542). From its distri- 
bution it is known as R. intermandibularis. 

VII. N. Facialis (n.7) (mixed, communis, visceral motor). In 
the Salamander the facial nerve leaves the brain in company with 
the auditory nerve and shares a common ganglion — the acustico- 
facialis ganglion (g. 7 + 8) — ^with it. The combined nerve leaves the 
lateral border of the medulla about f mm. posterior to the trigeminus, 
and passes into a recess in the petrosal bone near the anterior end of 
the otic capsule, and ventral to the auditory organ. Within this recess 
the ganglion is situated. The fine R. communicans passing to the 
Gasserian ganglion has already been mentioned (p. 139). In the 
larva, where it is much larger, it arises independently from the me- 
dulla — according to Druner — but in the adult it appears to leave the 
combined acustico-facialis nerve, or the ganglion itself. It is not 
always demonstrable. 

The facial nerve separates from the ganglion and passes directly 
outwards through a canal in the petrosal bone — the facialis canal — 
beneath the auditory organ, and finally enters the antrum petrosum 
laterale (cf. p. 26). 

(i) R. Palatinus (n.pal.) (communis). The palatine nerve is 
given off immediately distal to the ganglion, while the nerve is 
still in the facialis canal, and passes obliquely antero-ventrally 
through a special foramen in the floor of the petrosal bone 
(fo.pal. fig. 2), emerging from the foramen just lateral to the 
origin of the M. retractor bulbi. The nerve then turns antero-mesi- 
ally and crosses the M. retractor bulbi, passing between the muscle 
and the epithelium of the roof of the mouth. At the postero-mesial 



THE NERVOUS SYSTEM 145 

angle of the orbit it turns sharply anteriorly, passing alongside the 
tooth-bearing, backward extension of the pre-vomer. At a point a 
little anterior to the level of the optic nerve the palatine nerve passes 
dorsal to the pre-vomer, between it and the parasphenoid, and runs 
in a groove on the dorsal side of the former bone. 

At this point, or a little posterior to it, a small twig may remain 
outside the groove to pass forwards lateral to the palato-nasal artery. 
This is the lateral palatine nerve (n.pal'. ; Fig. 57). This nerve has not 
always been found, and in any case it is very fine. Nevertheless even its 
occasional presence is of interest in view of the condition in Siren, 
where it normally occurs and fuses with a special branch from the trige- 
minus nerve (cf. H. W. Norris, 1 9 1 3). In the Salamander it enters the 
nasal capsule and fuses with a branch coming from the main palatine- 
profundus anastomosis to form the post choanal nerve (n.p.ch.) (see 
below). Besides this branch the main palatine nerve gives numerous 
fine fibres to the mucous epithelium of the palate, which frequently 
have the appearance of an anastomosing network, and are parti- 
cularly conspicuous around the palato-nasal artery. 

The main palatine nerve continues along the dorsal side of the 
pre-vomer, passing between it and the 'Planum internasale', and 
enters the nasal capsule through the fenestra basalis. Here it is 
immediately joined by the ventral terminal branch of the ophthal- 
micus profundus V at the mesial border of the internal narial open- 
ing. At the junction of these two nerves a fairly conspicuous gang- 
lion occurs. A number of fine twigs are given off by the combined 
nerve, most of which pass anteriorly mesial to the internal narial 
opening, and spread over the roof of the mouth ventral to the olfac- 
tory epithelium, and ventral to the branches of the olfactory 
nerve. One branch, however, is given off laterally and passes 
round the posterior border of the internal nares where it is joined by 
the lateral palatine nerve — when such is present — to form the post 
choanal nerve (n.p.ch,). 

No post-palatine nerve, such as occurs in some Urodeles, has 
been found in the Salamander, and therefore there is no Jacobson's 
anastomosis. This result is in agreement with those of previous 
workers (cf. also Coghill's paper on Amblystoma). 

The remainder of the facial nerve, which may now be called the 
truncus hyomandibularis (n.hy.m.), on entering the antrum petro- 
sum laterale turns posteriorly and leaves the antrum by its posterior 
opening, i.e. between the otic and basal processes of the quadrate. 
Immediately after emerging from the antrum it gives off, antero- 
mesially: 



146 THE SALAMANDER 

R. aheolaris (r.al.) (communis) 

R. mandibularis internus . . .... Hoffmann. 

At the point where this branch is given off, von Plessen and Rabi- 
nowicz, Hoffmann and Driiner report a few scattered clusters of 
ganglion cells, but they are only detectable by microscopic means. 
Von Plessen and Rabinowicz call them the buccal ganglion. Driiner 
is convinced that they do not represent the remains of the ganglion 
laterale VII (connected with the lateral line system), and also finds 
fine sympathetic fibres entering the nerve in this region. Hoffmann 
goes further and suggests that the ganglion may be the forerunner 
of the otic ganglion of mammals. Certain it is that it exists and that it 
has a connexion with the sympathetic system. 

The alveolar nerve turns a little antero-laterally, and, keeping ven- 
tral to the quadrate and pterygoid, passes to the mesial side of the 
lower jaw. At about the level of the processus coronoideus it pene- 
trates a foramen in the pre-articular (fo.i.den; Fig. 7) and comes 
to lie between this bone and Meckel's cartilage — that is, on the 
mesial side of the latter. The nerve then passes anteriorly through 
the alveolar canal and emerges from it, anterior to the pre-articular, 
between the edges of the dentary. Its ultimate branches are distri- 
buted to the epithelium of the mouth-floor, between the M. genio- 
glossus and the mandible. Within the jaw it anastomoses several 
times with the internal mandibular branch of the Vth nerve as above 
noticed, but as a whole these two nerves are separated from one 
another by Meckel's cartilage. 

Homology of Chorda tympani. 

A very great deal has been written on, and around, this subject, 
and the conclusions drawn therefrom have been widely different. 
Only the briefest resume of the discussion may be given here. 

The chorda tympani is typically a mammalian nerve, and receives 
its name from its relations in the human subject, where it appears to 
pass through the middle ear, or tympanum, across the dorsal edge 
of the tympanic membrane. Actually the nerve is outside the tym- 
panum altogether, morphologically speaking, since it is covered by 
the mucous lining of the cavity. After receiving fibres from the otic 
ganglion it joins the lingual nerve — a branch of the Vth nerve — and 
with it enters the inferior dental canal of the lower jaw. 

Now the chorda tympani is a branch of the facial nerve, which 
in turn is a visceral nerve related to the spiracle or its homologue. 
Hence the question arises whether, if a nerve corresponding to the 



THE NERVOUS SYSTEM 147 

chorda tympani occurs in lower vertebrates at all, it is to be looked 
for as a branch of the pre- or post-trematic nerve. In 1 88 i Balfour 
claimed that it was a />r<?-trematic branch of the Vllth nerve. In 
1887 Froriep came to the conclusion that the chorda tympani is to 
be regarded as a ^oj/-trematic nerve, and although others have since 
supported Balfour's alternative suggestion, Froriep's conclusion re- 
presents the view now generally held. 

This discussion with regard to the morphological position of the 
chorda tympani has considerably complicated the second point at 
issue, namely, whether the R. alveolaris VII of the Salamander, and 
other Urodeles, is pre- or post-trematic, and is therefore to be re- 
garded as the true homologue of the chorda tympani or not. 

In 1895 Strong came to the conclusion that the chorda tympani of 
mammals, the R. mandibularis internus VII of Anura, and the R. 
alveolaris VII of Urodeles were all homologous structures. 

In 1 90 1 Driiner, who apparently accepted Froriep's view that the 
chorda tympani is post-trematic, denied its homology with the R. 
alveolaris VII of Salamandra on the ground that he believed the 
latter nerve to be pre-trematic. In both larval and adult Salamander 
the floor of the mouth is somewhat depressed ventrally between the 
mandibular and hyoid arch so as to form a sort of pocket — the plica 
hyomandibularis (plc.hy.m.; Figs. 37 and 38) — and this has been 
supposed to represent a vestige of the gill-slit between these two 
arches. On this supposition therefore Driiner argued that, since the 
R. alveolaris VII is a sensory branch of the hyomandibular nerve 
lying anterior to this supposed hyoid cleft, it must be pre-trematic. 
He accordingly thought the homologue of the chorda tympani was 
wanting in Urodeles. 

In 1902 Coghill, while accepting the view that R. alveolaris VII 
is a pre-trematic nerve, apparently regarded the chorda tympani also 
as a pre-trematic nerve, and, maintaining at the same time a some- 
what cautious attitude, held them to be homologous. He could not, 
however, agree with Strong in regarding these two structures as 
homologous with the R. mandibularis internus of Anura, since he 
believed this nerve to be post-trematic. 

In the same year C. K. Hoffmann (i 902) regarded both the chorda 
tympani and the R, alveolaris VII of Salamandra as pre-trematic 
branches of the Vllth nerve, and held that they were homologous 
structures. 

By 1 903 Driiner's views had undergone a profound change. As the 
result of the investigation of a developmental series of Siredon he 
found that the R. alveolaris VII was morphologically post-trematic 



148 THE SALAMANDER 

and developed posterior to the rudiment of the hyomandibular 
cleft, while the plica hyomandibularis was a secondary structure, 
developing behind the R. alveolaris VII, and in no way related to 
the visceral cleft. The true R. pre-trematicus VII he identified 
with a small branch of the R, palatinus VII distributed to the quad- 
rate. In his later paper, 1 904, he confirmed these results. 

Thus the matter stands, and the accepted views with regard to the 
questions enunciated above may be summarized. 

(i) The chorda tympani is a post-trematic branch of the Vllth nerve. 

(ii) The R. alveolaris VII of Urodeles is the true homologue of the 
chorda tympani of Mammals. 

It should be noted that the above resume is confined to the more 
immediately relevant literature and that all reference to the more 
general, but nevertheless important papers by Cole, Goodrich, 
Herrick, and Ruge, has been omitted for the sake of brevity. 

Very close to the point from which the R. alveolaris separates, the 
R. communicans IX-\-X ad VII (r.9 + 10— 7) joins the main nerve. "^ 
Driiner claims that it brings motor-fibres from the IXth nerve which 
join those of the R. jugularis VII. This is almost certainly incorrect. 
Without direct observation it is unsafe to make a definite statement, 
but it seems very probable that the condition in the Salamander 
resembles that described by Coghill for Amblystoma, viz. that the 
Xth nerve stndiS general cutaneous fibres which enter the R. jugularis 
VII, while the IXth nerve contributes communis fibres to the R. 
alveolaris VII. One feature which strongly supports this assumption 
is that one not infrequently finds the R. communicans bifurcating 
just before reaching the Vllth nerve, and one portion, the smaller, 
passing towards the R. alveolaris, and the other entering the R. 
jugularis about a millimetre further distalwards. It should, however, 
be noted that the same author can find no glossopharyngeal fibres 
in the R. communicans of Triton but only general cutaneous fibres 
from the vagus. 

After the separation of the R. alveolaris from the Truncus hyo- 
mandibularis the remainder of the nerve becomes known as : 

R. jugularis (n.j.) (visceral motor, and general cutaneous derived 
from vagus). 

R. mandibularis externus ...... Hoffmann. 

Its first branch, {a\ arises very close indeed to the point where the 
R. communicans IX +X ad VII enters it, and in fact it usually 

^ The R. communicans may be easily displayed by dissecting the M. depressor 
mandibulae away from its origin on the squamosal and turning it over ventrally. 



THE NERVOUS SYSTEM 149 

appears to spring from the angle formed by these two nerves. It is 
a R. muscularis to the M. depressor mandibulae. 

The main nerve continues ventro-laterally, passing between the 
above-named muscle and the squamosal to the surface of the muscle. 
Here it turns posteriorly, passing right across the muscle, between 
it and the skin. In doing so it gives ofF {h) one or two RR. cutanei 
to the overlying skin. 

At the posterior border of the M. depressor mandibulae the 
nerve makes yet another half-turn, and pursues an antero-mesial 
course, passing dorsal to the M. interhyoideus posterior, and ventral 
to the M. interhyoideus. A few fibres enter the M. interhyoideus 
posterior from its dorsal surface, while others turn back over its 
anterior edge and enter the muscle from the ventral side (i.hy.p.). 
The larger part of the nerve, however, spreads out fan-wise over the 
M. interhyoideus and innervates it (i.hy.). A few of the more lateral 
twigs pass right across the muscle to the space between it and the 
M. intermandibularis, and then turn dorsalwards to supply the M. 
subhyoideus, while a few fine fibres may pass right through the M. 
interhyoideus and enter the M. subhyoideus. 

Driiner (1901) thinks that the glossopharyngeal nerve also contri- 
butes motor-fibres to these muscles by means of the R. communicans, 
but, as stated above, it is exceedingly unlikely that the Salamander 
is so different from all other Urodeles, which have been investigated 
from the point of view of nerve components, as to possess motor- 
fibres in the R. communicans IX + X ad VII. 

Besides the motor branches to the interhyoid muscles described 
above there also arise, from the same fan-like termination of the 
nerve — the Pes anserinus (Driiner) — a number of sensory branches, 
which are (<:) RR. cutanei intermandibulares (Drtiner), supplying the 
skin underlying the muscles. 

Finally a few of the most anterior fibres of the R. jugularis may 
very occasionally extend to the trigeminus region and enter the 
posterior edge of the M. intermandibularis. 

VIII. N. Acusticus (n.8). As above described the auditory nerve is 
indistinguishable from the facial nerve until it separates from the acu- 
stico-facialis ganglion. It leaves this ganglion posteriorly, and enters 
the auditory sac by three foramina at the base of its mesial wall. The 
three foramina correspond to the three divisions of the auditory nerve. 

(i) The R. anterior (n.8. a.) supplies the recessus utriculi and the 
ampullae of the anterior and horizontal canals. 

(ii) The R. medianus (n.8.m.) is a very fine nerve and supplies 
only the sacculus. 



I50 THE SALAMANDER 

(iii) The R. posterior (n.S.Tp.) gives twigs to the ampulla of the 
posterior semicircular canal, to the lagena, and to the macula acustica 
neglecta. 

The R. medianus and its corresponding foramen were first de- 
scribed for Salamandra by Okajima in 1910, and its existence 
was confirmed in other Urodeles and in Anura by Miyawaki 
(1927-9). 

IX and X. NN. Glossopharyngeus and Vagus. 

The roots and ganglia of these two nerves are very closely asso- 
ciated in the adult Salamander, although — according to Hoffmann 
— they are distinct in the larva. It is therefore convenient to regard 
them as forming a complex arising laterally from the sides of the 
medulla oblongata by three roots, which unite and pass out of the 
cranial cavity through t\\e, foramen post-oticum (for. metoticum, Stadt- 
miiller), immediately posterior to the ear-capsule. The composite 
nerve then swells into a large glossopharyngeus-vagus ganglion which 
is thus situated outside the skull, in the angle between the ear- 
capsule and the occipital segment, and at about the level of the 
dorsal surface of the former. To expose it the MM. depressor 
mandibulae, cephalo-dorso-subpharyngeus, cucullaris, and inter- 
transversarius capitis superior must be removed, or turned back. 

IX. N. Glossopharyngeus (n.9) (mixed communis, visceral motor, 
and a few general cutaneous fibres). The glossopharyngeal nerve 
separates from the anterior portion of the ganglion and passes 
lateralwards between the MM. intertransversarius capitis superior 
and inferior, and then turns ventrad, lateral to the M. opercularis 
and mesial to the M. cucullaris. As it proceeds it gives off three 
branches. 

(i) R. communicans ad /^//(seep. 148) separates at about the level 
of the operculum and passes anteriorly round the otic capsule to join 
the facial nerve as described above. This branch probably carries 
only communis fibres from the glossopharyngeus and derives a 
general cutaneous component from the vagus. 

{^\)R.pre-trematicus{n.^rX.<^ (communis). This branch is given 
off very soon after the former, and at first passes anteriorly at right 
angles to the main nerve, but just posterior to the R. jugularis VII 
it turns ventrad and runs parallel with the internal carotid artery, and 
anterior to it. The R. pharyngeus (n.phar.) which, arising at the 
point where it turns ventrad, passes mesially, just anterior to the 
systemic arch, and is distributed to the mucous epithelium of the 
pharynx roof. Driiner describes ganglion cells associated with this 



THE NERVOUS SYSTEM 151 

nerve as well as connexions between it and the sympathetic system. 
It is associated with the pharyngeal branches of the succeeding 
nerves in the formation of a pharyngeal plexus. All these nerves 
are exceedingly fine and very difficult to dissect. 

The main portion of the R. pre-trematicus IX is distributed to 
the epithelium of the mouth-floor directly dorsal to the cerato-hyal 
cartilage. 

(iii) R. -post-trematicu (n.po.t.9) (general cutaneous, visceral motor, 
and communis). After the R. pre-trematicus has been given off, the 
post-trematic branch — constituting the major portion of the nerve — 
passes ventrally, parallel with the internal carotid artery, to the 
dorsal end of the posterior cornu (cerato-branchial I) of the hyo- 
branchial apparatus. Just before leaving the artery it gives off: — 

{a) One or two RR, cutanei jugulares (n.cut.j.) (Driiner) which pass 
to the skin overlying that region. They are exceedingly fine. 

{h) RR. pharyngei dorsales (Driiner), also very fine nerves dis- 
tributed to the dorsal wall of the pharynx and, according to Driiner, 
associated with sympathetic fibres around the internal carotid 
artery. 

In its course from the arterial arch to the cerato-branchial cartilage 
the nerve passes round the dorsal side of the large jugular lymph 
sinus, thus producing a loose loop which allows for the stretching of 
the nerve on the extrusion of the tongue. It then passes along the 
dorso-mesial side of the third visceral arch to the copula, giving off 
numerous twigs. 

{c) RR. musculares^ to the M. subarcualis rectus I. The remainder 
of the nerve now becomes: 

{d) R. UnguaUs^ and passes between the radials of the copula where 
it swells into a small ganglion — Ganglion copulare (Driiner) — from 
which emerge two or more branches passing directly into, and rami- 
fying through the tongue. The nerve is the sensory nerve to the 
tongue. 

Belonging perhaps to both IXth and Xth nerves are the NN. 
Cutanei occipitales^ anterior and posterior (nn.cut.oc.) (Driiner) arising 
from the dorsal side of the IX+X ganglion. These nerves, carrying 
general cutaneous components, pass through the overlying muscles 
— usually somewhat lateral to the M. intertransversarius capitis 
superior — to the skin covering the occipital region. In the larva they 
are associated with a lateralis component which atrophies during 
metamorphosis. 

X. N. Vagus +Accessorius Willisii (mixed communis, visceral 
motor, general cutaneous fibres). The Xth nerve of Salamandra is an 



152 THE SALAMANDER 

interesting example of the transition from a primitive fish-like state 
towards the condition found in more specialized land vertebrates. 
For example, the fourth and fifth visceral arch nerves retain a 
considerable amount of independence, and remain definitely asso- 
ciated with their respective arteries, while the nerve belonging to 
the sixth visceral arch has become more specialized, although still 
easily recognizable, and travels for a considerable distance in the 
track of the intestinal trunk. A careful study of PI. X, fig. 54 will 
make this clearer. 

(i) Fourth ^visceral arch nerve (n.vis.4). 

2nd Gill-arch nerve ....... Druner. 

Nearly always this nerve leaves the ganglion together with the 
fifth visceral arch nerve, but occasionally they remain quite distinct 
from the start. It passes lateral to the M. opercularis, and mesial to 
the M. cucullaris, as do the other visceral nerves emerging from the 
glossopharyngeus-vagus ganglion. Its course round the oesophagus 
lies between the carotid and systemic arches. About 4 mm. distal to 
the ganglion it gives oflF: 

{a) R. pharyngeus (n.phar.) which passes back mesially over the 
roof of the pharynx to enter the pharyngeal complex mentioned 
above. 

(b) RR. cutanei jugulares (n.cut.j.) which are given off at about the 
level of the carotid gland. They correspond with those already 
described for the glossopharyngeal nerve. 

A short distance beyond this point the nerve divides into two ter- 
minal branches (<:) and {d). 

(c) R. muscularis for the M. subarcualis rectus I (s.r. i) passes 
ventral to the common carotid, then, turning lateralwards, enters the 
muscle at the posterior end. It usually unites with a corresponding 
branch from the R. recurrens intestinalis X, and sometimes with one 
from the fifth visceral arch nerve also. According to Druner this 
muscular branch may be missing, but no such case has been met 
with by the present author. The nerve between the arterial arches 
and the muscle is greatly convoluted, thus allowing for the free 
movement of the hyobranchial apparatus. 

(d) R. recurrens (n.rec.4). This is a sensory branch and passes 
dorsal to the common carotid. It ramifies over the middle region 
of the mouth-floor, to the mucous epithelium of which its fibres 
are distributed. Druner reports occasional ganglion cells along 
its course. 

The same author considers that the whole visceral nerve as de- 



THE NERVOUS SYSTEM 153 

scribed above is to be regarded as the R. post-trematicus only, since 
he finds in the larva an exceedingly small R. pre-trematicus, only 
distinguishable as such by its position relative to the gill-slit, so that, 
even supposing that the nerve persists after metamorphosis — which 
is doubtful — it is quite unrecognizable after the gill-slit has dis- 
appeared. 

(2) Fifth visceral arch nerve (n.vis.5). 

3rd Gill-arch nerve . , . . . . . Druner. 

This nerve is very similar to the one just described and, as already 
mentioned, it is usually combined with it for a greater or lesser 
distance from the ganglion outwards. It passes round the oesophagus 
between the systemic and the 'third' arterial arches, and represents 
only the post-trematic branch. It is almost certain that the pre- 
trematic branch is wanting in the adult. Soon after leaving the gang- 
lion it gives the following five branches. 

{ii) R. muscularis for the M. cephalo-dorso-subpharyngeus (c.ph.). 
This branch leaves the nerve posteriorly and enters the muscle from 
its mesial side. 

{h) RR. pharyngei, corresponding to those of the previous nerve, 
come off very soon after, as do also : 

(c) RR. cutanei jugulares to the overlying skin. 

The remainder of the nerve may now divide, as did the previous 
one, into: — 

{d) R. muscularis., passing ventral to the systemic arch to join the 
corresponding branch of the fourth visceral nerve and supply the 
M. subarcualis rectus I, 

(<?) R. recurrens (n.rec.5) passing dorsal to the aortae and distri- 
buted to the floor of the pharynx. More usually, however, the former 
branch is missing. 

(3) Truncus intestino-accessorius (Druner). The main branch of the 
Xth nerve (n.tr.int.acc.) leaves the ganglion more posteriorly 
than the two branches already described and passes in an oblique 
postero-ventral direction, lateral to the M. opercularis, and mesial 
to the MM. cephalo-dorso-subpharyngeus, dilatator laryngis, and 
cucullaris. 

Very soon after leaving the ganglion the nerve gives off three 
branches: 

(d) A R. muscularis anteriorly, which supplies the dorsal part of the 
M. cephalo-dorso-subpharyngeus, and then posteriorly: 



154 THE SALAMANDER 

{b) The R. accessorius (n.acc.) to the M. cucullaris. This nerve 
enters the muscle on its mesial side. It is a fairly large branch. 

ic) The RR. pharyngei are very fine twigs to the pharynx corre- 
sponding with those already described for the other visceral 
nerves. 

Druner (Tafel 29, fig. 37) figures two fine twigs leaving the 
nerve at about this point and entering the M. levator scapulae (oper- 
cularis). In spite of very careful searching these twigs have not been 
found, and one can only suppose that, if they are of normal occur- 
rence, they must be exceedingly fine. He does not mention them in 
the text. 

(d) Almost immediately afterwards one or two RR. musculm-es 
(d.l.)for the M. dilatator laryngis are given off. They are very fine and 
are best seen by lifting the muscle carefully from its dorsal origin, 
when they will be seen stretching across through the connective 
tissue on its mesial side. Unless they are found in this way before the 
muscle is removed it is almost impossible to pick them up afterwards. 

At the ventral edge of the M. opercularis a very fine anastomosis 
with the hypoglossal nerve can sometimes be found, and almost im- 
mediately afterwards is given off": 

{e) The R. recurrens (n.rec.io). This important branch passes 
mesialwards, ventral to the V. facialis, and proceeds to divide as 
follows : — 

(i) A'', laryngeus ventralis (n.lar.v.). 

R. laryngeus ......... Druner. 

This nerve comes off variously, either soon after the R. recurrens 
separates from the intestinal branch, or later as the former nerve 
turns oralwards (i.e. as Druner describes). In the former case it is 
combined with the ramus muscularis for the M. dilatator laryngis. 
It passes directly mesiad, ventral to the muscles, and enters the M. 
constrictor laryngis ventrally. 

(ii) R. muscularis (d.l.) for the M. dilatator laryngis. This twig 
arises soon after the separation of the recurrent branch, either 
separately, or together with the former branch. It turns posteriorly 
and enters the muscle on its ventral side. 

(iii) RR. musculares for the M. cephalo-dorso-subpharyngeus, 
comprising a number of fine twigs entering the muscle from the 
ventral side. 

(iv) Amongst the former twigs a fine branch leaves the main nerve 
and passes along the ventral side of the M. cephalo-dorso-sub- 
pharyngeus to enter the M. transversus ventralis IV. 



THE NERVOUS SYSTEM 155 

(v) A sensory branch to the mucous epithelium of the pharynx and 
mouth leaves the recurrent nerve close to the foregoing branches on 
the ventral side of the M. cephalo-dorso-subpharyngeus. It passes 
dorsalwards to the floor of the pharynx between the pulmonary arch 
and the above-named muscle. It is relatively of fair size, and corre- 
sponds with the recurrent branch of the fourth and fifth visceral 
nerves and, like them, passes dorsal to the arterial arches. 

The remainder of the nerve then turns anteriorly and remains 
ventral to the arterial arches. It passes very close to the truncus 
arteriosus, and, at about the level of the pulmonary arch, it gives off: 

(vi) A fine anterior cardiac nerve (n.c.a.) to the truncus arteriosus. 
The residue of the nerve is now: 

(vii) Thei?. muscularis for the M. subarcualis rectus I. It passes 
anteriorly and fuses with the corresponding branch of the fourth vis- 
ceral nerve, and with that of the fifth visceral nerve also — when such 
is present. 

Driiner reports that a large number of ganglion cells are to be 
found in the region of the R. recurrens, both ventral to the muscles 
and on the branches to the mucous epithelium. 

These various branches of the recurrent nerve are somewhat diffi- 
cult to dissect owing to their small size and rather pale colour. The 
dissection is best carried out from a ventro-lateral aspect. The cucul- 
laris muscle must be removed and the M. rectus cervicis pulled over 
ventrally, and then the nerves may be detached from the connective 
tissue in which they are embedded. 

(/) From the intestinal trunk immediately posterior to the recur- 
rent nerve is given off to the larynx a fine branch which is here 
named dorsal laryngeal nerve (n.lar.d.). It appears to have been over- 
looked by Druner since he does not mention it in his paper. Its 
position relative to the M. dilatator laryngis appears to vary some- 
what, since it has been found both anterior and posterior to the 
muscle. In either case, however, it passes mesially, closely applied 
to the wall of the pharynx, and dorsal to the laryngeal artery. 
Although it appears to terminate in the M. constrictor laryngis and 
sends a branch to the dilatator muscle, its translucent appearance and 
close relation to the pharyngeal wall would lead one to suspect that 
it is probably in the main a sensory nerve to the larynx. 

The intestinal nerve divides into pulmonary and gastric branches 
almost immediately after giving off the branch just described. 

{g) The common pulmonary nerve (n.pul.) is the more ventral of the 
two and soon divides again into two. 

(i) A lateral pulmonary nerve (n.pul.l.), which accompanies the 



156 THE SALAMANDER 

pulmonary artery down the side of the lung and ramifies over its 

surface with the arterial branches. 

(ii) The anterior ■pulmonary nerve (n.pul.a.) passes mesially and 
ramifies over the anterior end of the lung and the muscle-fibres of 
that region. It passes to the point where the pulmonary vein leaves 
the lung on the mesial side, and there gives off two branches. 

{a) One or raoro. posterior cardiac nerves (n.c.p.) which accompany 
the pulmonary vein to the heart. 

{h) "Wit gastric ;/(?r^'^(n. gas.) which proceeds posteriorly for some dis- 
tance, lateral to the lung, and then turns mesially and divides into two. 

(i) A ventral gastric branch (n.g.v.) distributed to the ventro-lateral 
stomach wall, and 

(ii) A dorsal gastric branch (n.g.d.) supplying the dorso-lateral wall. 

SPINO-OCCIPITAL NERVES (PI. X, fig. 54, n.sp.occ.) 

Driiner (Teil i, p. 553 et seq.) describes a fine anastomosis leaving 
the ventral side of the glossopharyngeal vagus ganglion and passing 
round the occipital condyles laterally to join the ventral ramus of 
the first spinal nerve. He considers it is probably the homologue of 
the spino-occipital nerves of Selachians and Holocephali described 
by Fiirbringer (1896). 

His results have been confirmed for Salamandra (n.sp.occ.) and 
a nerve has been found leaving the IX+X ganglion close to 
the branch just described which fuses with the dorsal ramus of 
the N. spin, i, within the M. intertransversarius capitis superior 
(n.sp.occ'). Driiner mentions this connexion as a rare occurrence. 
It is very fine and difficult to dissect so that it is easily missed. 

These nerves may represent the ventral roots belonging to seg- 
ments which have become fused together in the occipital region of 
the skull posterior to the vagus, but Druner was not able to satisfy 
himself entirely on this point as he could not trace their centres of 
origin in the medulla. In this connexion reference should be made 
to the more recent work of Goodrich (191 1) and of de Beer (1926) 
on the segmentation of the vertebrate skull in general. 

SPINAL NERVES 
I. Historical. 

Most authors who have described the cranial nerves, e.g. 
Fischer, von Plessen and Rabinowicz, Druner, and Hoffmann, 
have included the first spinal nerve in their account, since it sup- 
plies certain muscles connected with the hyobranchial apparatus 
and tongue. Details of the discussion relating to the morpholo- 



THE NERVOUS SYSTEM 157 

gical position of this nerve advanced by these and other authors 
are given below. 

The brachial plexus was well described by Fiirbringer in 1873, 
and his account was republished five years later by Hoffmann in 
Bronn's Thier-reich. In this latter work Hoffmann also published 
an account, accompanied by an original figure of the crural plexus. 
His description is not quite complete. 

A number of authors have dealt with various details connected 
with the nerves of the limbs in papers primarily concerned with 
muscles, which are summarized in the muscle-section of the 
present work. Amongst these may be mentioned Sieglbaur, Rib- 
bing, Appleton, and McMurrich. 

Amongst the synonyms given will be found the names of authors 
dealing with Urodeles other than Salamandra, which have been 
included to facilitate comparison. 

The method adopted here is the same as that chosen by Gaupp in 
his Anatomie des Frosches^ v'yl. to describe first a typical spinal nerve 
— for which purpose N. spinalis 10 has been chosen — and then to 
give an account of the ventral rami of those nerves which become 
specially modified, e.g. N. hypoglossus, and the nerves supplying 
the fore- and hind-limbs. 

2. A Typical Spinal Nerve. 

iV. spinalis 10 (PI. XXIV, fig. 79) arises from the spinal cord by 
a dorsal and a ventral root. The dorsal root arises some distance 
posterior to the ventral root, but both pass together through a special 
foramen in the neural arch immediately posterior to the transverse 
process of the ninth vertebra (see also p. 19). The two roots 
fuse within the foramen, and immediately after emerging from it 
swell into a large ganglion — the spinal ganglion 10 (g.sp.io). The 
spinal ganglion thus lies outside the vertebral column, but it is never- 
theless protected by the transverse process anteriorly, and by the 
broad articular processes dorsally, as well as by the tough fibrous sac 
in which it is enclosed. Within this sac, surrounding the ganglion, 
is a spongy fatty tissue, and there is no trace of the chalky masses 
found in this position in the frog. 

From the ganglion emerge the usual dorsal and ventral rami, each 
being composed of motor and sensory fibres, and supplying respec- 
tively the dorsal and ventral body muscles and the skin overlying them. 

R. dorsalis. This ramus arises from the ganglion as several 
twigs. The largest of these emerges dorsally, then turning directly 
caudad, passes alongside the vertebral column almost to the level 



158 THE SALAMANDER 

of the next transverse process. Here it turns sharply dorsally again 
and passes between the MM. interspinals and dorsalis trunci 
to supply the skin of the back. During its course parallel with 
the vertebra it gives off one or two twigs to the above-named 
muscles. This nerve would appear to correspond with the R. cutaneus 
dorsi medialis (n.cut.d.m.) of Gaupp (Frog), carrying with it some 
motor-fibres. 

Another cutaneous branch, rather smaller than the previous one, 
and evidently representing the R. cutaneus dorsi lateralis (n.cut.d.l.) 
of Gaupp (Frog), emerges from the lateral aspect of the ganglion. 
Then, passing towards the transverse process, it penetrates the tough 
connective tissue surrounding the bone and myosepta of the M. 
dorsalis trunci, and supplies the overlying skin. There are also 
two small muscular branches leaving the lateral aspect of the gang- 
lion, one turning dorsally into the main portion of the M. dors, 
trunci, and the other passing into the intertransversarius portion of 
the same muscle. 

R. ventralis. This ramus is considerably larger than the ramus 
dorsalis and forms the main portion of the nerve. It leaves the ven- 
tro-lateral aspect of the ganglion, and, as its name implies, is distri- 
buted to the muscles and skin of the ventral body-wall. It passes for 
a short distance through the sub-vertebral muscles and supplies them 
with fibres, then immediately after, or just before emerging from 
them ventrally, it gives off the R. communicans ad sympatheticus. The 
nerve then passes around the body-wall between the peritoneum 
and the muscles, giving off a cutaneous branch about midway — 
the R. cutaneus abdominis lateralis (n.cut.ab.L). It then supplies 
several twigs to the MM. obliquus externus superficialis and 
profundus, and to the ventral portion of the M. transversarius 
s. obliquus internus. 

At the lateral edge of the M. rectus abdominis profundus it sends 
a branch to the muscle (r.ab."), and then divides into muscular and 
cutaneous branches. The former (r.ab.') passes deep to the muscle 
and supplies the M. rectus abdominis superior, while the latter 
becomes superficial and supplies the skin over the belly — R. cutaneus 
abdominis medialis (n.cut.ab.m.). 

The exceptions to the general plan exemplified by the tenth spinal 
nerve described above will now be considered individually and in 
detail. They may be summarized briefly: 

The first spinal nerve has no dorsal root and only exceptionally 
a ganglion in the adult. There is no dorsal cutaneous branch and 
the ventral ramus supplies the tongue muscles. 



THE NERVOUS SYSTEM 159 

The second spinal nerve is normal with regard to its dorsal root, 
ganglion, and cutaneous branches, but its ventral ramus contributes 
both to the N. hypoglossus — supplying the tongue muscles — and to 
the brachial plexus. 

The third, fourth, and fifth spinal nerves are all exceptional in that 
their ventral rami enter into the brachial plexus and supply the 
muscles of the fore-limb and shoulder. 

The ventral rami of the fifteenth, sixteenth, and seventeenth 
spinal nerves similarly anastomose to form the crural plexus supply- 
ing the muscles of the pelvis and hind-limb. To this plexus the 
fourteenth and eighteenth spinal nerves also contribute, but only in 
a minor degree. 

3. The First Spinal Nerve. 

TV", spinalis 1 (= N. spinalis 2, Hoffmann) (PI. X, fig. 54, n.sp.i). 
There has been some controversy over the morphological position 
of this nerve, together with the related question of the segmentation 
of the occipital region of the skull and of the 'atlas' vertebra (see also 

P- 17)- 

So far as it is possible to separate the discussions concerning the 
nerve itself from those relating to the skeletal structures they may 
be briefly stated as follows. 

In 1878 Hoffmann suggested that the first vertebra of Urodeles 
actually represents two vertebrae fused because it is penetrated by 
the first spinal nerve. In 1902 he confirmed this view by studying 
the larval stages of Triton and Salamandra. He then found the 
ventral root of a transitory nerve emerging between the occipital 
condyles and the vertebral column — N. suboccipitalis — and related 
to the first myotome. He regarded this transitory nerve as the first 
spinal nerve because it emerges between the occiput and the verte- 
bral column, and therefore he argued that the first spinal nerve of 
the adult is morphologically N. spinalis 2. He also found an even 
more transitory nerve (ventral root only) anterior to the N. sub- 
occipitalis belonging to his myotome-o. 

Druner (1904) disagreed with this view. He accepted Peter's 
explanation that the atlas vertebra was single and not two vertebrae 
fused, and said (Teil II, p. 439, footnote): *Nach meinen Befunden 
ist der den i . Wirbel durchsetzende Nerv mit demjenigen identisch, 
welchen C. K. Hoffmann bei jiingern Embryonen zwischen Occipi- 
talknorpel und i. Wirbelbogen austreten sah.' 

Goodrich (191 1) apparently arrived at similar results from the 
study of the larva of Amblystoma. 



i6o THE SALAMANDER 

In 1 9 1 7 Froriep, as the result of studying the larval stages of Sala- 
mandra atra^ returned to Hoffmann's view. He found a sub-occipital 
nerve which he regarded as the first spinal nerve, and came to the 
conclusion that the atlas vertebra is a double one and therefore that 
the first spinal nerve of the adult is morphologically the second. 

More recently, Mookerjee (1930) has shown that in Triton the 
atlas, although it has a double origin, is not strictly a double verte- 
bra (cf. 'atlas' in skeletal section). He finds the sub-occipital nerve, 
however, and calls it N. spinalis i, and calls the hypoglossal nerve, 
N. spinalis 2. 

To summarize: it seems certain, therefore, that the rudiment of 
the nerve which appears between the occipital condyles and the verte- 
bral column should be regarded as N. spinalis i, and that the next 
posterior, which is the first spinal nerve of the adult, is morpho- 
logically N. spinalis 2. Nevertheless since the N. suboccipitalis is 
so very transient and probably never attains functional importance, 
it seems unnecessary to insist on using the strict morphological 
nomenclature, thereby confusing the description of all the remaining 
spinal nerves. In the following account the nerves are accordingly 
numbered as they appear in the adult, leaving the N. suboccipitalis 
out of the series. 

The first spinal nerve arises from the spinal cord by two ventral 
roots only. A dorsal root and a ganglion occur in the larva, but they 
atrophy during metamorphosis. Driiner reports that in three out of 
fifteen adults he finds a trace of the ganglion remaining, but such 
a circumstance has not come under the notice of the present writer. 
Correlated with the absence of the ganglion there is no dorsal cuta- 
neous branch, but the dorsal muscular branches are present and 
innervate the MM. intertransversarius capitis superior and inferior, 
and the M. rectus capitis posterior. 

The R, ventralis passes directly ventral between the subvertebral 
and dorsal musculature, and immediately on reaching the ventral 
surface of the muscles it gives ofF (i) a very fine ramus communicans 
ad sympatheticus (r.com.ad.sym.). It then turns somewhat postero- 
laterally and gives off two more branches. 

(ii) A small twig to the M. intertransversarius capitis inferior 

(i.c.i.). 

(iii) One or two twigs to the M. opercularis (o.) as it passes ventral 
to the edge of this muscle. 

At this point the nerve passes very close to the tenth cranial nerve 
just at the point where the latter divides into gastric and pulmonary 
branches. A very fine anastomosis is usually discernible at the 



THE NERVOUS SYSTEM i6i 

juxtaposition of these two nerves, but it does not appear to be quite 
constant. 

The first spinal nerve now turns anteriorly and in doing so passes 
between the subclavian vein and the small cutaneous vein. 

(iv) Immediately ventral to the subclavian vein a strong anasto- 
mosis from the second spinal nerve is received, so that the vein passes 
through the angle between the anastomosis and the proximal portion 
of the main nerve. The nerve thus formed, containing fibres from 
the ventral branches of both first and second spinal nerves, is the 
N. hypoglossus proper. 

(iv a) From the middle of the anastomosis a fine twig is given off to 
the M. pectori-scapularis (p.s.). 

(v) N. hypoglossus (n.hyp.) (N. hypobranchialis, Driiner). Imme- 
diately after its formation, as the result of the fusion of branches 
from the first two spinal nerves, the hypoglossal nerve turns 
directly forwards along the lateral edge of the M. rectus cervicis. 
It supplies several twigs to both the superficial and deep portions 
of this muscle, then passes ventral to the thyroid gland, and at the 
anterior end of the gland turns somewhat mesially to pass through 
the substance of the M. genio-hyoideus. It supplies several fine 
twigs to this muscle (g.hy.) and emerges from it at the anterior end 
of the cerato-hyal (see Figs. 3 7 and 38) to enter the tongue and inner- 
vate the MM. genio-glossus and hyo-glossus. 

With regard to the homology of the hypoglossus of Urodeles with 
that of the higher vertebrates the statement of Goodrich (P.Z.S. 
191 1, p. 118) may be quoted here. He says '. . . the hypoglossus of 
Amphibia and Amniota may certainly be considered as homologous, 
although not necessarily composed of the same segmental nerves. 
It is owing to the shortness of the skull in Amphibia that the hypo- 
glossal roots come out behind it.' In the same paper he also shows 
that the urodele hypoglossus innervates muscles derived from the 
ventral outgrowths of the second, third, and fourth myotomes. 

4. The Second Spinal Nerve. 

N. spinalis 2 (PL XI, fig. ^z^^ n.sp.2) possesses both dorsal and ven- 
tral roots and also a ganglion, so that its dorsal branches contain both 
cutaneous and motor-fibres, and hence conform to the normal plan. 

The R. ventralis, after emerging from the subvertebral muscula- 
ture, gives off the R. communicans ad sympatheticus and turns postero- 
laterally to the anterior border of the shoulder. Here, according to 
Fiirbringer, it divides into four main branches: (i) N. thoracicus 
superior 2; (ii) N. thoracicus inferior 2 anterior; (iii) N. thoracicus 

4038 M 



r62 THE SALAMANDER 

inferior 2 posterior ; and (iv) a branch to the ventral body muscles and 

the skin of the neck. 

With the exception of the first branch this does not seem to give 
a very accurate picture of the branching of the nerve, which is much 
more variable than Fiirbringer's figure and description suggest. 

(i) A^. thorackus superior 2 comprises two or three fine branches 
which turn dorsalwards and supply, {a) the M. thoraci-scapularis 
(t.s.), and {Jy) the M. cucullaris minor (c') near its insertion. 

(ii) Almost immediately after giving off this branch the nerve 
turns a little anteriorly and sends two fairly stout cutaneous branches 
which spread over the skin at the base of the neck region and over the 
procoracoid and the anterior edge of the scapula. After separating 
from the nerve they pass directly laterally and emerge at the angle be- 
tween the procoracoid and the M. cucullaris. It is very difficult to 
be sure whether or not they supply any motor-fibres to the MM. pro- 
coraco-humeralis and cucullaris, but it is very probable that they do 
not. These nerves supply a region corresponding with that supplied 
by the supra-clavicular nerves of man, and as this name is hardly 
appropriate in an animal not possessing a clavicle, they are therefore 
called here cutaneous cervical^ or NN. cutanei cervkales (n.cut.cer.). 

(iii) After a very fine twig to the M. pectori-scapularis has been 
given off the nerve very soon divides, more or less equally into its 
three terminal branches, as follows : — 

(iv) R. communkans ad N. spinalis J, which has already been 
described — see above. 

(v) A R. muscularis to the MM. rectus cervicis superior and pro- 
fundus. The branch to the former muscle passes mesial to the 
profundus muscle. 

(vi) A R. communicans ad N. spinalis 3 (r. com. 2-3) which arises 
somewhat variously. It may consist of a single bundle, separating 
from the main nerve near the point where it turns anteriorly, 
close to the N. thoracicus superior 2, to join the third spinal nerve 
near the point where the N. supracoracoideus emerges. It bifur- 
cates at this point, one branch entering the N. supracoracoideus 
direct, while the other crosses this nerve and enters the main trunk 
of N. spinalis 3. Quite frequently, however, an auxiliary bundle 
arises separately from the second spinal nerve, and passes obliquely 
to join the N. supracoracoideus just before it passes through the For. 
supracoracoideum in the pectoral girdle. 

The series of anastomoses, Spino-occipital — N. spinalis i ; N. 
spinalis i — N. spinalis 2; and N. spinalis 2 — N. spinalis 3, is fre- 
quently spoken of as the cervical plexus. 



THE NERVOUS SYSTEM 163 

5. Brachial Plexus, and Nerves of the Fore-limb (PI. XI). 

The actual plexus and the nerves supplying the flexor muscles of 
the limb are best dissected from the ventral side, but the nerves to 
the extensor muscles should be dissected dorsally. 

The precise pattern of the plexus varies considerably, but the net 
result obtained is always the same, namely, that most of the fibres 
from N. spinalis 3 pass to the dorsal side of the limb and innervate 
the extensor muscles, while the major portion of N. spinalis 4 remains 
ventral and supplies the flexor muscles. N. spinalis 5 makes only a 
small contribution to the plexus, which mainly enters N. pectoralis. 

Third spinal nerve (ventral branch) (n.sp.3). 

Very soon after emerging from the subvertebral musculature the 
third spinal nerve gives off dorsally one or two fine branches to the 
M. thoraci-scapularis (= NN. thoracicus superior 3, Fiirbringer). 
Shortly afterwards a small branch is given off mesially which divides 
to supply the M. rectus abdominis and the M. obliquus internus 
(origin). 

N. supracoracoideus (12, Fiirbringer) (n.s.cor.) arises at the point 
where the scapula and coracoid join, passes mesially, closely applied 
to the dorsal surface of the latter, and penetrates the For. supra- 
coracoideum to the ventral surface. Here it divides into three 
branches, one branch supplying the M. procoraco-humeralis (p-h.) 
(14, Fiir.), another the MM. supracoracoideus and coraco-radialis 
proprius (s.c.) (13, Fiir.), while the third is a cutaneous branch to 
the skin over the neck and breast (cut.) (15, Fiir.). 

It is this nerve which receives most of the anastomosis from the 
second spinal nerve. 

At about this level, or a little later, N. spinalis 3 divides rather 
unequally into a larger anterior branch and a smaller posterior 
branch. The latter joins the main branch of the fourth spinal nerve 
to form the flexor nerve of the fore-limb (N. brachialislongus inferior, 
Fiirbringer). The former, after receiving one or two small contribu- 
tions from N. spinalis 4, passes to the dorsal side of the limb to 
innervate the extensor muscles (NN. brachiales longi superiores, 
Fiirbringer). 

TV. subscapularis (n.s.sc.) (29, Fiir.) to the muscle of the same 
name leaves the anterior branch of the Ilird nerve just before it 
receives its contribution from the IVth. It is a very fine nerve. 

N. dorsalis scapulae (n.d.sc.) (30, Fiir.) frequently arises as two 
nerves — or as one nerve which divides almost immediately — and 



i64 THE SALAMANDER 

passes round the posterior border of the scapula to its lateral surface, 
mesial to the M. dorsalis scapulae. The smaller posterior nerve 
goes entirely to the muscle. The larger anterior nerve gives a few- 
small twigs to the muscle and then passes to its anterior edge, where 
it divides into three cutaneous and two muscular branches. The 
twolatterenter the adjoining edges of the MM. dorsalis scapulae and 
procoraco-humeralis respectively, while of the cutaneous branches 
{NN. cutanei brachii superiores later ales ^ 32, Fur.), one (a. Fig. c^^) 
supplies the skin covering the lateral surface of the arm, while 
another (j8. Fig. ^^ turns mesially over the procoracoid to the skin 
of the breast, and the third (y. Fig. ^^) turns dorsally to the skin of 
the scapular region. 

N. axillaris (n.ax.) (Osawa) is a fairly large cutaneous nerve 
arising close to the N. dorsalis scapulae. It passes out between the 
M. dors. scap. and the M. dors, humeralis, and supplies the skin 
covering them. 

A^. dorsalis humeralis (n.d.h.) (= N. latissimus dorsi, 34, Fiir.) 
arises from an anastomosis between N. spinales 3 and 4, and appears 
to receive fibres from both spinal nerves. It passes dorsalwards 
alongside NN. dors. scap. and axillaris and enters the M. dorsalis 
humeralis mesially. 

The nerves arising from the brachial plexus distal to this point 
will be dealt with under the headings 'Extensor nerves' and 'Flexor 



Fourth spinal nerve {ventral branch) (n.sp.4). 

The proximal part of N. spinalis 4, after emerging from the sub- 
vertebral muscles, gives one or two fine dorsal branches to the 
M. thoraci-scapularis (= N. thoracicus superior 4, Fur.). Between 
this point and the plexus one or two fine branches are given off to 
the M. rectus abdominis. 

Extensor nerves. 

NN. brachiales longi superiores ..... Furbringer. 
These consist of two^ main nerves which enter the arm posterior 
to the humerus and shoulder-joint, passing between the heads of the 
MM. anconeus scapularis medialis and anconeus coracoideus in 
company with the Vena brachialis medialis. They pass to the dorsal 
side of the humerus. 

' Fiirbringer reports one anomalous case in which the extensor nerve in the arm was 
represented by a single large bundle, instead of the two parallel nerves found normally. 



THE NERVOUS SYSTEM 165 

N. extensorius cranialis (n.ex.cr.). Sieglbaur (1904). 

Musculo-spiral, or radial nerve .... Humphry (187 1). 
N. radialis profundus , . Fiirbringer (1873); Hoffmann (1873-8); 

Ribbing (1907). 
N. radialis superior ...... Osawa (1902). 

This is the larger of the two extensor nerves and is the main con- 
tinuation of N. spinalis 3. Immediately on entering the arm the 
nerve turns slightly anteriorly and gives a branch to the M. anconeus 
humeralis lateralis. It then passes obliquely across the limb between 
the MM. anconeus humeralis lateralis and medialis, remaining 
lateral to the humerus, and emerges at the surface of the muscle on 
the flexor side of the elbow. Here it gives a fairly large branch 
which spreads over the skin on the extensor surface of the fore-arm — 
the A^. cutaneus antibrachii dorsalis (n.cut.abr.d.). A little farther on 
another cutaneous branch arises and passes right along the radial 
side of the fore-arm as far as the wrist, where it becomes known as 
the N. dorsalis manus radialis (n.d.m.r.). On reaching the carpus it 
divides, one small twig continuing along the radial side of the first 
digit, while the rest of the nerve crosses the wrist to the interdigital 
space between the first and second fingers. Here it divides again 
and supplies the skin on the adjoining sides of these two digits. 
Although mainly cutaneous, this nerve also sends a twig — soon 
after its origin at the elbow — to the M. extensor antibrachii et carpi 
radialis, and, as it crosses the carpus, it gives very fine twigs to the 
MM. extensores breves digiti I and II. 

The main extensor cranialis nerve, after giving off these cutaneous 
branches, turns mesially and describes a half-spiral turn round the 
head of the radius, passing between this bone and the M. extensor 
antibr. et carpi rad., so as to lie in the interosseal space between the 
radius and the ulna. As it passes over the radius it gives two twigs 
to the M. extensor antibr. et carpi rad. 

About midway between the wrist and the elbow the nerve divides 
into two branches. 

{a) The N. dorsalis manus intermedins (n.d.m.i.) which remains 
in the interosseal space, and passes along the fore-arm to the 
carpus, where it becomes more superficial, and lies directly under 
the M. extensor digitorum communis. From here it passes to the 
space between the second and third digits and bifurcates to supply 
the neighbouring sides of these two fingers. As it passes over the 
carpus the nerve supplies fine twigs to the MM. extensores breves 
digitorum II and III, and at the proximal part of the wrist, a larger 



i66 THE SALAMANDER 

branch which supplies the MM. ext. brev. dig. I, and abductor 
digiti I. 

{b) The second branch is the N. dorsalis manus ulnaris (n.d.m.u.). 
Almost immediately after its separation it receives one of the 
terminal branches of the N. extensorius caudalis (q.v.). It then runs 
along the fore-arm over the ulna, parallel with the previous branch, 
and gives off one or two twigs which pass outwards, between the 
MM. extensores antibr. et carpi radialis and ulnaris, to supply the 
M. extensor digitorum communis. From these twigs fibres may 
also pass into the M. extensor antibr. et carpi radialis. In the region 
of the wrist this nerve becomes more superficial and lies directly under 
the M. extensor digitorum communis, and, after giving off a branch 
to supply the skin on the ulnar side of the fourth digit, it crosses to 
the space between the third and fourth digits and divides to supply 
the skin on the adjoining sides of these two fingers. It also gives fine 
twigs to the MM. extensores breves digitorum III and IV. 

N. extensorius caudalis (n.ex.cd.). Sieglbaur (1904). 
Posterior ulnar, or better, inferior 

musculo-spiral nerve . ..... Humphry (1871). 

N. radialis superficialis . . Fiirbringer (1873); Hoffmann (1873-8)^ 

Ribbing (1907). 
N. radialis inferior ...... Osawa (1902). 

This is the smaller of the two extensor nerves and is separated 
from the anterior extensor nerve by the Vena brachialis medialis, but 
receives one or two fine anastomosing twigs from it. It arises more 
definitely from the N. spinalis 3-4 anastomosis than does the N. ex- 
tensorius cranialis. 

Soon after entering the arm it gives off a large cutaneous branch, 
N. cutaneus brachii dorsalis (n.cut.br.d.). This branch often carries 
some motor-fibres for the MM. anconaeus scapularis medialis and 
anconaeus coracoideus. Then follows a short branch which divides 
into three and supplies the scapularis medialis, humeralis and cora- 
coideus portions of the anconaeus muscle, while a little farther on a 
small twig is given off to the M. anconaeus humeralis lateralis. Still 
farther distalwards another twig to the M. anconaeus coracoideus 
may frequently be found. 

On reaching the flexor side of the elbow the nerve bends round 
the antero-lateral edge of the M. anc. hum. lat., and passes to the 
extensor side of the fore-arm, crossing obliquely over the origin of 
the M. extensor digit, comm., between the muscle and the skin. At 
the elbow it gives a cutaneous branch to the ulnar side of the fore- 



THE NERVOUS SYSTEM 167 

arm, and then passes round under the M. extensor antibr. et carpi 
ulnaris — i.e. between this muscle and the ulna — into the interosseal 
space of the fore-arm. In doing so it supplies one or two twigs to the 
above-named muscle and finally gives a large anastomosis — amount- 
ing almost to the entire nerve in some cases — to the N. dorsalis manus 
ulnaris. The remainder of the nerve ends in the MM. extensores 
antibr. et carpi radialis and ulnaris. 

The exact relation between the NN. extensorius caudalis and 
cranialis at this point is subject to some variation. In one case no 
certain anastomosis between these nerves in the fore-arm could be 
found. In this example both nerves were approximately equal in 
size all the way along, and the anterior nerve did not divide in the 
fore-arm but formed only the N. dors, manus inter., while the 
posterior nerve ran parallel with it and formed the N. dors, manus 
ulnaris. If any fusion between these nerves occurred at all it was at 
the elbow-joint, in a similar position to that described by Sieglbaur 
(1904) for Necturus, but owing to the ribbon-like, translucent 
appearance of the nerves in this specimen, it was impossible to 
decide whether one was dealing with a nervous connexion or merely 
with the connective tissue capsule of the joint. In any case the 
anastomosis, if it did occur, was very fine and in no way comparable 
with the normal condition described above. 

Flexor nerves. 

N. brachialis (n.br.). 

N. brachialis longus inferior .... Furbringer (1873}. 
After receiving the anastomosis from the third nerve, N. spinalis 
4 gives off posteriorly N.pectoralis (n.pect.) to the pectoralis muscle. 
The N. pectoralis passes round the posterior border of the coracoid 
and enters the muscle on its mesial side. It is accompanied by a 
cutaneous nerve which supplies the skin covering the pectoral 
region. From about this level another cutaneous nerve arises, N. 
cutaneus brachii superior medialis (n.cut.br.s.m.) (42, Fiir.), supplying 
the skin of the axilla, and the postero-mesial aspect of the upper 
arm. 

After giving off these branches, the brachial nerve (which forms the 
main continuation of the fourth spinal nerve) enters the arm ventral 
to the extensor nerves, posterior to the shoulder-joint, and between 
the M. anconaeus coracoideus and the M. anconaeus scapularis 
lateralis, and passes to the ventral side of the humerus. It then 
divides into a R. superficialis and a R. profundus (Furbringer), 



1 68 THE SALAMANDER 

and at the same time describes a half-spiral turn so as to lie on the 
flexor side of the humerus, between the M. coracobrachialis longus 
and the M. humero-antibrachialis inferior. The two rami run 
parallel with one another separated by the A. brachialis. They are 
usually approximately equal in size, but the R. super, may be larger 
than the R. prof. Normally also this latter ramus passes to the elbow 
without branching, but one or two fine muscular twigs may be given 
from it. 

At the point where these two nerves separate, or immediately 
afterwards, one or two branches are given off from the R. super- 
ficialis to the MM. coraco-brachialis longus and brevis; they are 
the NN. coraco-brachiales (n. cor.br.). 

Then follow one or two cutaneous branches which pass out to the 
skin between the M. coraco-brachialis longus and the M. humero- 
antibrachialis. One branch longer than the rest travels along the 
arm to the elbow, and then takes a half-spiral turn to the lateral 
aspect of the fore-arm, crossing the origin of the M. flexor primor- 
dialis communis in doing so. These branches represent the N. cuta- 
neus hrachii inferior lateralis of Fiirbringer, but are here called A^. cut. 
hrachii inf. medialis^ to agree with the blood-vessels. The R. super- 
ficialis also supplies the M. humero-antibrachialis inferior with one 
or two fine branches. 

At the elbow-joint it bifurcates. One branch remains superficial 
and passes along the skin on the radial side of the fore-arm and first 
digit. At the base of the finger it sends a branch mesialwards which 
supplies the short flexor muscles related thereto, and finally joins the 
N. interosseus. This branch of the R. superficialis would appear to 
represent Fiirbringer's A^. cutaneus hrachii inferior medialis^ here 
called N. cut. hrachii inf. lateralis. 

The other branch turns mesially, and passing under the M. flexor 
antibr. et carpi radialis, gives a small branch to this muscle, and 
finally joins the R. profundus. As already indicated this latter nerve 
passes right down the arm to this point without giving off any 
important branch. The nerves arising from this plexus will be 
termed N. ulnaris and N. interosseus.^ 

N. ulnaris (n.ul.). 

R. superficialis ulnaris McMurrich (1903). 

This branch passes distally and obliquely across the fore-arm, 
deep to the MM. flexor primordialis communis and flex, antibr. ulna- 

' Ribbing applies these names to the proximal portions (R. superficialis and R. pro- 
fundus) also, but they seem hardly applicable to this region of the limb. 



THE NERVOUS SYSTEM 169 

ris, but superficial to the M. caput longum musculorum contrahen- 
tium. When it arrives at the mesial edge of the latter muscle, at 
about the level of the middle of the fore-arm, it gives two muscular 
branches. The smaller of these divides into three to supply the M. 
flex. prim. comm. and the MM. flex, antibr. et carpi ulnaris and 
radialis. The other branch R. medianus (Ribbing) = R. superficialis 
medialis, McMurrich (n.ul.r.m.), passes along the mesial edge of 
the M. cap. long. muse, contra., and appears to terminate at the 
point where the M. flexor accessorius medialis joins the tendon of the 
M. flex. prim. comm. It innervates the M. cap. long. muse, contra, 
and the M. flex. ace. med. 

The main N. ulnaris, after giving off the R. medianus crosses the 
ulna, and, following the mesial border of the M. flex carpi ulnaris, 
emerges at the side of the arm between this muscle and the M. flex. 
ace. lat. At the proximal border of the M. flex. ace. lat. a twig enters 
the muscle, and as the nerve passes along its lateral border a fairly- 
constant branch is given off to the skin. At the base of the fourth 
digit another cutaneous branch passes along the ulnar side of the 
finger, while the main nerve turns mesially again and crosses the 
base of the finger, between the MM. eontrahentes digitorum and 
the M. flex. brev. prof. IV to the interdigital space of the third and 
fourth digits. Here it divides again into three main branches, of 
which two are distributed to the skin of the adjoining sides of the 
third and fourth fingers, while the remaining one crosses the base of 
the third digit, between the M. contra, digit, and the M. flex. brev. 
prof. Ill, to join the corresponding palmar branch of N. interosseus. 
Besides these cutaneous branches a number of small muscular 
twigs may be traced into the several short flexor muscles of the 
fourth digit, e.g. M. contra, digit., M. flex. brev. super, and prof., 
M. flex, digit, min., and M. interossei III-IV. 

A^. interosseus (n.io.). Ribbing (1907). 

R. profundus ...... McMurrich (1903). 

N. medianus ....... Sieglbaur (1904). 

This branch almost immediately dips down to the dorsal side of 
the M. interosseus antibrachii, and passes alongside the extensor 
nerves — NN. dors, manus inter, and ulnaris — in the interosseal 
space of the radius and ulna. It lies anterior — radial — to the 
N. dors, manus inter. There is no anastomosis between these nerves 
in the Salamander as there is in Triton (cf. Sieglbaur, 1 904). During 
its course down the fore-arm it gives one or two twigs to the M. 
interosseus antibrachii, and also to the M. pronator profundus. The 



lyo THE SALAMANDER 

nerve becomes more superficial again at the wrist, where it passes 
dorsal to the tendon of insertion of the M. cap. long. muse, contra., 
between the basalia of digits i and 3. At the interspace of these 
digits it bifurcates and each ramus again divides into two, so that 
the posterior ramus supplies the skin on the pre-axial side of the 
third digit — N. digitalis ventralis 3 — and also joins the terminal twig 
of the N. ulnaris, while the anterior ramus similarly innervates the 
post-axial side of the second finger — N. dig. vent. 2 — and then passes 
across to the interspace of the first and second digits and sup- 
plies the skin of their adjoining sides — NN. dig. vent, i and 2 — and 
finally joins the N. cut. inf. lat. It is not certain that this latter fusion 
is quite constant, but it certainly occurs in a sufficient number of 
cases to justify its being considered 'normal'. There is thus a com- 
plete nerve loop across the palm. The N. interosseus also innervates 
the short flexor muscles of digits 2 and 3 by small twigs in the appro- 
priate places. 

The ventral ramus of the Jifik spinal nerve (n.sp.5), on emerging 
from the subvertebral muscles, divides into two branches, and the 
posterior one divides again several times to supply the various 
muscles of the ventral body-wall in that region. The anterior branch 
also divides again, one branch anastomosing with N. spinalis 4 and 
entering the brachial plexus, while the other is distributed to the 
skin of the axilla and the ventro-posterior side of the arm. This 
branch is usually joined by a branch arising direct from the plexus 
3-4 which is somewhat variable in size, and in one case could not 
be found. The resulting nerve may be called N. cutaneus brachii 
ventralis (n.cut.br.v.). 

Fiirbringer seems to have overlooked this nerve unless it is repre- 
sented by one of the 'Aeste fur Bauchmuskeln' which he has not 
followed in detail. This author also figures a condition sometimes 
met with, in which the branch entering N. spinalis 4 bifurcates just 
before doing so, and sends a small twig to join the N. pectoralis. 
From this, and from the fact that the fifth nerve nearly always enters 
the fourth close to the point whence the N. pectoralis emerges, it 
seems safe to conclude that at least some fibres from the fifth spinal 
nerve normally contribute to the pectoral nerve. 

6. Crural Plexus, and Nerves of the Hind-limb (PI. XII). 

The crural plexus of the Salamander is formed by the fifteenth, 
sixteenth, and seventeenth spinal nerves, with small contributions from 
the fourteenth and eighteenth. The main nerves of the plexus are 



THE NERVOUS SYSTEM 171 

sixteen and seventeen.^ It is noteworthy that, as compared with the 
fore-Hmb, the N. femorahs — from the fifteenth and sixteenth nerves 
— enters the Hmb anterior to its articulation with the pelvic girdle, 
while in the fore-limb no nerve passes into the arm in front of the 
glenoid cavity. With this exception there is a marked similarity 
between the two plexuses. For example, the N. obturatorius is 
strongly reminiscent of the N. supracoracoideus, or again, the 
extensor or dorsal nerves of the limb arise from the plexus anterior 
to the flexor or ventral nerves in both limbs. The comparison 
amounts almost to identity in the distal parts of the limb — a feature 
which is of course correlated with the close similarity of the muscula- 
ture of the fore-arm and hand with that of the leg and foot. 

The anastomosis with the fourteenth nerve is slight and involves 
only branches of the N. ileohypogastricus. 

Fifteenth spinal nerve {ventral branch) (n.sp. 1 5). 

This nerve emerges from the penultimate trunk vertebra and 
passes obliquely postero-laterally towards the limb. Its first branch 
is an anterior one to the muscles of the lateral and ventral body-wall, 
the N, ileohypogastricus (n.il.hg.). The actual point at which this 
branch leaves the main nerve varies considerably in different speci- 
mens, since it may be either nearer to, or more distal from the verte- 
bral column than shown in Plate XII, which is about the average 
position. At what may be termed the groin, i.e. in the anterior 
angle between the limb and the body, the remainder of the fifteenth 
spinal nerve divides into three. The first of these branches is the N. 
cutaneus femoris medialis (n.c.fm.m.) (Hoffmann) supplying the skin 
over the antero-dorsal region of the thigh. Hoffmann describes a 
branch to the M. ileo-extensorius from this nerve, but it has not been 
possible to confirm its presence, and it probably represents an indivi- 
dual variation. The remainder of the nerve has been called A^. cruralis 
anterior by Hoffmann ; it gives off' the following two branches. 

N. obturatorius (n.o.) passes ventrally through the M. pubo-ischio- 
femoralis-internus, supplying it with many twigs in doing so (p.i.f.i.), 
and finally penetrates the obturator foramen to innervate the anterior 
portion of the M. pubo-ischio-femoralis externus. The other branch 
fuses with the anterior branch of N. spinalis 1 6, to form : 

N.femoralis (n.fm.). Osawa (1902); Sieglbaur (1904). 

Anterior crural nerve ...... Humphry (1871). 

N. cruralis ....... Hoffmann (1873-8). 

' Sieglbaur gives the sixteenth, seventeenth, and eighteenth for the crural plexus, but 
he probably regards N. hypoglossus as N. spinalis 2. 



172 THE SALAMANDER 

This nerve passes into the leg penetrating the M. pub. isch. fern, 
int. to which it gives a few fine twigs. Just before entering the 
muscle it gives off a branch to the anterior portion of the M. extensor 
iliotibialis (e.il.t.'). The nerve takes a rather winding course in the 
thigh, at first turning sharply posteriorly so as to lie close to the proxi- 
mal end of the femur on its dorsal side. It then turns right round and 
proceeds along the limb, winding itself around the femur so that it 
lies directly anterior to it at the knee. At the antero-dorsal aspect 
of this joint the nerve becomes superficial to the muscles, emerging 
between the M. pub. isch. fern. int. and the M. pubo-tibialis, i.e. 
dorsal to the latter muscle. Several cutaneous branches arise in this 
region which Hoffmann takes to represent the A^. cutaneus femoris 
internus {saphenus minor) (n.c.fm.i.). 

The nerve then continues along the antero-dorsal border of the 
leg immediately under the skin as: 

N. dorsalis -pedis tibialis (n.d.p.t.). Sieglbaur (1904); Humphry 

(1871). 

N. saphenus .... Osawa (1902); Ribbing (1908). 

Just distal to the knee the nerve gives a small twig to the M. exten- 
sor cruris tibialis (e.t.t.), and, at about a third of the distance between 
the knee and the ankle, another branch penetrates the same muscle. 
A considerable proportion of this branch is cutaneous and passes 
right through the muscle to reappear near the middle line on the 
dorsum of the leg, but a number of fibres do actually terminate 
within the muscle itself. 

Ribbing (1908) was the first to call attention to the motor-fibres 
in the N, dors. ped. tib. (his N. saphenus), and they are not common 
to all Urodeles. 

Numerous cutaneous branches are given off to the skin on the 
mesial and dorsal sides of the leg and foot. 

At the base of the first digit the nerve divides, one twig — N. digi- 
talis dorsalis i (n.d.d.i) — passing along the pre-axial side of the first 
toe, while the other crosses the base of the digit dorsally and then 
divides into two so as to supply the adjoining sides of the first two 
digits — NN. digit, dors, i and 2. 

The relations of the N. femoralis vary considerably in different 
specimens, and particularly with regard to the point at which it 
emerges from the thigh muscles. This usually occurs at the knee 
as described, while it may emerge much nearer the body, but in any 
case its course through the thigh forms a half-spiral turn. The 



THE NERVOUS SYSTEM 173 

close correspondence between the course of the N. dors. ped. tib. 
in the leg and that of the N. dors. man. rad. in the fore-arm is obvious. 

Sixteenth spinal nerve {ventral branch) (n.sp. 1 6). 

This nerve emerges from the last trunk vertebra and passes 
somewhat obliquely postero-laterally across the dorsal pelvic region 
to the posterior border of the acetabulum, and there splits into two 
more or less equal parts. The anterior portion, which however may 
be much smaller than the other, turns anteriorly to join a branch from 
the fifteenth spinal nerve and form the N. femoralis as described 
above, while the posterior branch enters the limb close to the 
seventeenth nerve immediately behind the hip-joint. It anastomoses 
closely with N. spinalis 17, but nevertheless its main continuation 
in the thigh is the N. extensorius (n.e.) (cf. N. spin. 3, in brachial 
plexus). The N. ischiadicus ventralis (n.i.v.) also receives a con- 
siderable proportion of its fibres from the sixteenth spinal nerve. 

N. extensorius (n.e.). 

N. fibularis .... Hoffmann (1873-8); Appleton (1923-8). 
N. peroneus ....... Sieglbaur (1904). 

After its formation from NN. spin. 16 and 17 the extensor 
nerve passes dorsal to the M. iliofemoralis, and gives a twig 
to it, and then divides into two rami which pass along the thigh 
parallel with one another immediately ventral to the MM. iliofibu- 
laris and extensor iliotibialis. They supply these muscles (il.f. and 
e.il.t.), and give off one or two cutaneous branches which probably 
represent the N. cutaneus surae lateralis (n.c.s.l.), while they also send 
a fine twig to the knee-capsule. The two rami reunite proximal to the 
knee to form a single nerve, or in some cases the nerve remains single 
throughout. At the knee-capsule it becomes : 

A^. peroneus (n.pn.). 

This nerve leaves the thigh dorsal to the M. iliofib., and, execut- 
ing a half-spiral turn, enters the leg, passing ventral to the M. 
femoro-fibularis between this muscle and the fibula. During its 
passage under the skin, superficial to the origin of the extensor cruris 
et tarsi fibularis, it gives off" one or two muscular branches which 
enter this muscle (e.c.t.f.) on its post-axial side. The nerve passes 
to the interosseal space of the tibia and fibula, and there divides 
into two principal rami which correspond to the NN. dors. man. 
inter, and ulnaris in the fore-arm. They have accordingly been 
named by Sieglbaur N. dorsalis pedis intermedius (n.d.p.i.) and A^. 



174 THE SALAMANDER 

dorsalis pedis fibularis (n.d.p.f.)- The N. dors. ped. fib. — the post-axial 
branch — almost immediately divides again and gives off a branch 
which passes along the leg between it and the N. dors. ped. inter. It 
innervates the M. extensor digitorum communis (e.d.) and MM. 
extensores breves digitorum 3 and 4 (e.b.3 and e.b.4) and sends a 
large branch to join the N. dors. ped. inter. This ramus is not men- 
tioned by Sieglbaur. It is named here R. accessorius n. dorsalis pedis 
fibularis (n.d.p.f.). 

A comparison with the fore-limb suggests that the essential 
difference between it and the hind-limb is that in the former case 
the anterior and posterior extensor nerves remain separate until dis- 
tal to the elbow-joint, so that the extensor communis muscle and the 
ulnar extensor muscles are innervated by sundry twigs from both the 
N. ext. caudalis, and from the N. dors. man. ulnaris — a branch from 
the N. ext. cranialis. In the hind-limb, however, the anterior and 
posterior extensor nerves of the thigh unite proximal to the knee to 
form a single trunk which redivides in the leg, so that the extensor 
communis muscle and the fibular extensors are innervated by the 
N. dors. ped. fib. and its R. accessorius, which separate from the N. 
dors. ped. inter, as a single bundle. A comparison of Plates XI and 
XII will make this clear. 

The N. dorsalis pedis fibularis (n.d.p.f.), after separating from the 
R. accessorius, passes down the leg along the extensor side of the 
fibula, immediately deep to the M. extensor cruris et tarsi fibularis. 
It innervates this muscle by several twigs (e.c.t.f.). At about the 
level of the distal end of the fibula it receives an anastomosis from 
the R. accessorius, while at the distal edge of the M. extensor cruris 
et tarsi fibularis it gives off a cutaneous branch to the skin of the 
post-axial side of the fifth digit — N. digitalis dorsalis 5. The remain- 
der of the nerve crosses the base of this toe, giving a muscular twig 
to the M. ext. brev'. digit. 5, and then divides to supply the adjoining 
sides of digits 4 and ^—NN. digitales dorsales 4 and 5. 

N. dorsalis pedis intermedius (n.d.p.i.). This comparatively large 
nerve passes along the interosseal space of the tibia and fibula 
to the tarsus. It lies immediately posterior to the N. interosseus, and 
during this part of its course it innervates the tibial extensor muscles, 
viz. M. extensor tarsi tibialis (e.t.t.), and M. extensor cruris 
tibialis, the latter twigs coming off" near the origin of the nerve. At 
the ankle it gives twigs to the M. abductor et extensor digiti i 
(a.e.i), and to the MM. ext. breves digitorum of the second digit 
(e.b.2). The nerve then divides into two, one branch — the anterior — 
supplying the adjoining sides of digits 2 and 3 — NN. digitales dors. 



THE NERVOUS SYSTEM 175 

2 and 3 — while the posterior branch is similarly distributed to 
digits 3 and 4, forming A^A^. digitales dorsales 3 and 4. The posterior 
nerve also receives an anastomosis from the R. accessorius n. dors, 
ped. fib. 

N. ischiadicus ventralis (n.i.v.). Appleton (1928). 

Arises from a 16—17 anastomosis as a nerve of moderate size. It 
passes ventral to the M. caudali-femoralis, between it and the M. 
pubo-ischio-femoralis externus. It first gives one or two small 
branches to the M. ischio-femoralis (is.fm.) and then turns antero- 
ventrally between the M. caud. fem. and M. pub. isch. fem. ext., 
giving off a large twig to supply the MM. pubo-ischio-tibialis 
(p.i.t.), pubo-ischio-femoralis externus (p.i.f.e.), and ischio-flexorius 
(pars propria) (is.f.). It then turns more anteriorly and gives off a 
large musculo-cutaneous branch. The cutaneous elements go to the 
skin covering the antero-ventral surface of the leg and form part of 
the A^. cutaneus femoris ventralis (n.c.fm.v.). They emerge between 
the M. pub. isch. tib. and the M. pub. tib., while the muscular 
twigs supply the M. pubo-tibialis (p.t.). The nerve also gives at, 
or near, this point a small branch to the M. pubo-femoralis (p.fm.). 
The remaining portion of the N. isch. vent, now becomes the TV. 
cutaneus femoris ventralis (n.c.fm.v.), and passes along the thigh to 
the knee between the muscles pubo-tibialis and pubo-ischio-tibialis. 
At the knee it becomes superficial and passes right along the tibial 
border of the fore-leg beneath the skin. It is continued to the ex- 
tremity of the first digit as the N. digitalis ventralis i (n.d.v.i). At 
the base of the digit it sends a branch mesialwards over the skin of 
the sole of the foot, from which a twig may arise to anastomose with 
a terminal branch of the N. interosseus, thus completing a plantar 
nerve loop corresponding with the palmar loop of the hand. It is 
very doubtful whether this anastomosis is a constant feature, since, 
when present, it is always exceedingly fine and difficult to dissect 
without breaking. The N. cut. fem. vent, may receive a branch from 
the sciatic nerve in the popliteal region, but this also does not seem 
to be constant. 

Seventeenth spinal nerve (ventral branch) (n.sp. 1 7). 

N. spinalis 1 7 emerges from the sacral vertebra and passes almost 
transversely to the posterior border of the acetabulum, where it 
joins the sixteenth nerve, and, after anastomosing with it, continues 
into the leg as the sciatic nerve, from which arise the ventral or 
flexor nerves of the leg. 



176 THE SALAMANDER 

At the point where it joins the sixteenth nerve a branch arises 
from the posterior edge of N. spinalis 17 which suppHes the muscles 
attaching the tail to the pelvic girdle. This branch is A^. pudendus 
(n.pd.). A little distal to this nerve one or two cutaneous branches 
are given off to the skin covering the ventro-posterior region of the 
thigh — they are the A^. cutaneusfemoris posterior (n.c.fm.p.). 

N. pudendus (n.pd.). Appleton (1928). 

Leaves the seventeenth spinal nerve, as above described, just 
proximal to its junction with the sixteenth, and passes posteriorly 
mesio-dorsally to the M. caudali-femoralis. Almost immediately 
after its origin it gives one or two fine cutaneous branches to the 
postero-dorsal side of the leg. They pass dorsal to the M. caud. fem. 
and form part of the N. cut. fem. post. At about the same level 
a branch from the eighteenth spinal nerve enters the N. pudendus. 
The latter ner\^e then divides into four terminal branches, of which 
one enters the M. caudali-femoralis (cd.fm.), another supplies the 
M. caudali-pubo-ischio-tibialis (c.p.i.t.), and from this a cutaneous 
branch passes between the two muscles above-named to the skin 
at the side of the cloaca. Another branch passes to the cloacal gland 
in the male — or to the equivalent tissue in the female — between the 
MM. caud. pub. isch. tib. and ischio-caudalis, giving a small twig 
to supply the latter muscle in passing (is.cd.). This nerve would 
appear to be the N. perinealis (n.pe.) of Appleton. It is larger and 
better seen in the male. 

A^. sciaticus (n.s.). 

Sciatic nerve ...... Humphry (1871). 

N. ischiadicus ...... Hoffmann (1873-8). 

N. tibialis communis ...... Osawa (1902). 

N. tibialis ........ Appleton (1928). 

The sciatic nerve passes down the thigh in company with the 
sciatic artery, and, a little proximal to the knee, gives off the follow- 
ing branches. First, one or more cutaneous branches which leave 
the thigh between the MM. pub. isch. tib. and isch. flex, and supply 
the skin covering the ventral (flexor) surface of the leg distal to 
the knee; they represent the ISf. cutaneus surae medialis (n.c.s.m.). 
One of these nerves — when there is more than one present — passes 
either close to, or through the 'pars plantaris' of the M. ischio- 
flexorius and supplies it with a small twig (is. P.). The other branch, 
which leaves the sciatic nerve near these cutaneous branches, is a 
muscular one. It divides to supply the M. femoro-fibularis (fm.f.), 



THE NERVOUS SYSTEM 177 

and then enters the M. flexor primordiaHs communis. It may- 
terminate within this muscle or it may pass right through and fuse 
with the N. fibularis (as figured, PL XII). In the latter case a branch 
passes from it to supply the muscle (f.p.). As already mentioned 
an anastomosing branch may pass from the mesial side of the sciatic 
nerve in this region to join the N. cut. fem. vent. 

At some point between the hip and the knee the sciatic nerve 
splits to allow the sciatic artery to pass through it, but the actual 
extent of this split is subject to great variation. It may be quite 
short, when it occurs just proximal to the knee, or it may be of 
moderate length, as in the figure (PI. XII). On the other hand, one 
case has been observed in which two approximately equal nerves, 
representing the sciatic, passed down the thigh parallel with one 
another, and with the sciatic artery between them. One of these 
nerves was derived from N. spin. 17 and the other from N. spin. 16. 
They fused at the knee. This case, therefore, offers an explanation of 
the phenomenon of an artery penetrating an apparently solid nerve, 
and at the same time explains the variation in length of the split in 
different specimens, since this is merely a measure of the degree of 
fusion between the sixteenth and seventeenth nerves. 

The whole sciatic nerve leaves the thigh dorsal to the M. flex, fib., 
and enters the leg between the insertion of the M. pub. isch. tib. 
and the origin of the M. flex. prim. comm. Here it divides — if it has 
not already done so just proximal to the knee — and the divisions 
correspond very closely to the branches of the N. brachialis in the 
fore-arm. They have, therefore, been given analogous names in 
spite of the fact that the term 'N. fibularis' has been applied by some 
authors to an entirely different nerve — ^viz. N. peroneus. Thus the 
two main divisions of the sciatic nerve distal to the knee are here 
called N. fibularis and N. interosseus, while the mesial branch of the 
N. fibularis is termed R. medianus since it corresponds with the 
similarly named branch of N. ulnaris in the fore-arm. 

N. fibularis (n.f.) (mihi). 

External popliteal ...... Humphry (187 1). 

R. lateralis n. ischiadici . .... Hoffmann (1873-8). 

This nerve is not to be confused with Hoffmann's 'fibularis', 
which is the dorsal or extensor nerve and the equivalent of the nerve 
here described under the name 'peroneus'. As stated above, the 
close similarity between the nerve now under consideration and the 
N. ulnaris in the fore-arm seems to the writer sufficient justification 
for giving it an analogous name. 

4038 N 



178 THE SALAMANDER 

It passes down the fibular side of the leg deep to the M. flexor 
primordialis communis, between this muscle and the M. flexor 
accessorius medialis. Directly after separating from the N. interos- 
seus the fibular nerve gives off a R. medianus (n.f.m.), which passes 
along the mesial — pre-axial — side of the M. cap. long. muse, contra. 
It innervates this muscle (c.l.c), as well as M. flex. ace. lat. (f.a.L), 
and terminates finally in the M. flex. ace. med. (f.a.m.). Just after 
it separates from the N. fibularis it gives a small twig to the M. plant, 
super, minor (pi.") of Eisler. 

The main N. fibularis lies post-axial to the M. cap. long. muse, 
contra., close to the fibula. It may send a twig to the M. cap. long, 
muse, contra, (c.l.c), but its main function is to innervate the M. 
flex. prim. comm. (f.p.c), and the short flexor muscles of the fifth 
digit (a.d.5 and e.b.5). It also gives a cutaneous branch to the post- 
axial side of the fifth toe, N. digit, vent. 5. An anastomosis between 
a terminal branch of this nerve and one from the posterior ramus of 
the N. interosseus is usually demonstrable. 

A^. interosseus (n.io.) (mihi). 

Internal popliteal . ...... Humphry (187 1). 

R. medialis n. ischiadici . .... Hoffmann (1873-8). 

N. tibialis medianus ...... Sieglbaur (1904). 

This branch passes down the leg in the interosseal space between 
the tibia and fibula dorsal to the M. interosseus cruris. It supplies 
this muscle (i.e.), as well as the M. pronator profundus (p.p.)- I^ 
appears again from beneath the muscle at the tarsus, between the 
tarsal bones of the second and third digits, but remains dorsal to the 
tendinous insertion of the M. cap. long. muse, contra. In this region 
it divides into three. One branch passes directly to the interdigital 
space of the second and third toes, where it bifurcates into the 
NN. digit, vent. 2 and 3, and the other two branches (which would 
seem to correspond to Humphry's anterior and posterior tibial 
nerves) similarly form the NN. digit, vent, i and 2, 3 and 4, and also 
4 and 5, and in addition give twigs to all the short flexor muscles of 
these digits. The post-axial branch usually joins a terminal branch 
from the fibular nerve, while the pre-axial twig may — but probably 
does not always — anastomose with one from the N. cut. fem. ventr. 

It should be noted that, in the Salamander, there is no exchange of 
fibres between this nerve and the dorsal, or extensor, nerves as there 
is in many Urodeles. 

The ventral ramus of N. spinalis 18 bifurcates, immediately after 
penetrating the ilio-caudalis muscle, and the anterior portion passes 



THE NERVOUS SYSTEM 179 

forwards inside the tough fibrous sheath surrounding the rectum, 
and anastomoses with the N. pudendus from the seventeenth spinal 
nerve. 

The posterior ramus turns caudad, and sends a twig to the M. 
ilio-caudalis (il.cd.), and then penetrates the sheath to supply the 
cloacal gland in the male, or the homologous tissue in the female. 

The ventral ramus of N. spinalis ig besides supplying the sub- 
vertebral muscle also supplies the cloacal gland. 

The remaining spinal nerves, or caudal nerves as they may be 
termed, show gradual reduction in size and complexity towards the 
end of the tail, but are not characterized by any other special modifi- 
cation. 

SYMPATHETIC NERVOUS SYSTEM^ 
I. Historical. 

Only three papers have been published on the sympathetic ner- 
vous system of Salamandra. The first, and best known, is by Anders- 
son (1892). It is somewhat incomplete, but he gives an elaborate 
figure which illustrates the general relations of the abdominal and 
caudal portions very well, though the direct connexion he shows with 
the vagus ganglion must be considered doubtful, while his state- 
ment that a cranial sympathetic is entirely wanting is without doubt 
erroneous. The next account, by Jaquet (1900), is obviously based 
on Andersson's and suffers from the same defects. In 1902 Hoff- 
mann's paper appeared on the development of the sympathetic 
system in Salamandra. This work adds several important details to 
Andersson's account, the most notable of which are the intimate 
association between the sympathetic nerves and the adrenal bodies 
(Nebenniere) and the close connexion of the latter with the veins. 
He denies that the sympathetic chain has any direct connexion with 
the vagus ganglion, and says that it terminates anteriorly with the 
ramus communicans to the first spinal nerve. From this point it 
travels mesialwards along the ventral ramus of this nerve, separates 
from it again at the point where it leaves the dorsal ramus, and 
passes forward to the vagus ganglion through the spino-occipital 
muscles. This certainly misrepresents the facts as far as the adult 
is concerned, since the anterior continuation of the sympathetic 
beyond the ramus communicans to the first spinal nerve is easily 
demonstrable by dissection under the binocular microscope, while 

' It must be noted here that the term 'sympathetic' is used in the anatomicah&ns&,a.nd 
is not intended to include the system as a whole, which is more correctly called the 
autonomic nervous system. 



i8o THE SALAMANDER 

the connexion described by Hoffmann between the vagus ganglion 
and the first spinal nerve is doubtless identical with that interpreted 
here as the spino-occipital nerves (see p. 156). 

2. Description. 

Following Andersson's arrangement, it is most convenient to 
describe the system in three sections, viz. (i) the cephalic -portion^ 
anterior to the ramus communicans to N. spin, i, (ii) the cervical and 
abdominal portions continuing from the posterior extremity of the 
cephalic portion to the commencement of (iii), the caudal portion^ 
lying within the haemal canal. It is further convenient to describe 
sections (ii) and (iii) before dealing with the more difficult and 
obscure cephalic section. 

(ii) The cervical and abdominal portions (PL XVII, fig. 64). In 
general arrangement the sympathetic system consists of ganglia 
arranged in a paired longitudinal series on either side of the dorsal 
aorta, connected serially with each other by a chain of nerve 
fibres, and transversely with the ventral rami of the spinal nerves by 
the rami communicantes. There are also several cross-connexions 
between the two longitudinal series which will be mentioned sub- 
sequently. 

The most anterior ganglion of this section — i.e. the one receiving 
the r. comm. n. sp. i — is very small, as also is the next, but the third 
ganglion, lying immediately anterior to the subclavian artery, is a 
large one. It receives the r. comm. n. sp. 3. This section is called by 
Andersson the cervical portion. Anastomoses between the two sides 
occur in this region, both dorsally and ventrally to the aorta, and they 
usually pass obliquely from the second to the third ganglia. The 
nerve chain of each side in this section is usually double, and often 
treble, while sometimes it becomes so subdivided that it almost dis- 
appears. 

The ganglion associated with the r. comm. n. sp. 3 is the anterior 
subclavian ganglion (Andersson) and from it emerge three, or some- 
times four, nerves. Two of these pass to the gang, subclav. postr. 
(Andersson), one lying dorsal, and the other ventral to the sub- 
clavian artery. The dorsal nerve is very much stouter than the ven- 
tral. The third constant branch is a lateral one passing along the 
anterior side of the subclavian artery to the fore-limb. On the 
animal's right side this nerve bears a small ganglion situated about 
a millimetre from the dorsal aorta. This ganglion has not been 
observed on the left-hand side. It is usually connected with the pos- 
terior subclavian ganglion by a fine nerve which crosses the sub- 



THE NERVOUS SYSTEM i8i 

clavian artery obliquely. The fourth, and more variable, branch to 
leave the gang, subclav. antr. is an anterior visceral nerve, which 
passes alongside the anterior gastric artery to the stomach. This 
also seems to occur more frequently on the right side than on the 
left. Where the anterior visceral nerve does not separate from the 
anterior subclavian ganglion, it does so from the posterior one. 
The nerve described by Andersson as leaving the nerve strand just 
in front of the gang, subclav. antr. and entering N. tr. intest. X has 
not been found. 

It may be noted here that the rami communicantes from the first 
six spinal nerves are not easy to see, since they frequently leave the 
nerve while it is still within the subvertebral muscles, and pass 
mesially through them to emerge close to the vertebral column. 
Andersson fails to find any r. communicans from the fourth spinal 
nerve, but it is certain that such exists in the majority, if not in all cases. 

The ganglion suhclavius posterior (g.scl.p.) marks the anterior end 
of the abdominal portion of the sympathetic chain. It is fairly large 
and frequently gives rise to the anterior visceral nerves (n.vis.a.) — 
see above. The next two ganglia — five and six — are also large, and, 
like all the subsequent ganglia of this section, have adrenal tissue 
intimately associated with them. In this section also the ganglia and 
the adrenal tissue are very closely applied to the post-cardinal veins, 
and frequently envelop them like a sleeve. The longitudinal nerve- 
fibres run parallel with the veins, and there is also a tendency in this 
region, particularly in the anterior two-thirds, for the fibres to pass 
from one ganglion to another in a divided condition, so that the 
strand then appears to be double, treble, or in some higher multiple. 
Sometimes one of the smaller bundles may miss a ganglion and join 
the succeeding one, so that it is not possible to give an exact and 
detailed description of the chain which will serve for any specimen. 
Similarly with the ganglia, they also are very variable both in size and 
arrangement. They normally, but not invariably, occur at the points 
where the rami communicantes join the longitudinal strand. Bearing 
in mind the disjunctive condition of this region of the sympathetic 
chain in Selachians (Young, 1933), it is evident that Salamandra 
forms a truly intermediate type between this and the more regular 
and definite arrangement found in the Frog. There is a distinct 
tendency for the rami communicantes in the posterior region of the 
abdomen, i.e. in the region of the kidneys, to bifurcate and join 
the longitudinal chain at two points. This bifurcation may occur 
right at the point of separation from the spinal nerve, so that the 
ramus communicans then appears double. Whether this subdivision 



i82 THE SALAMANDER 

represents the morphological separation of the pre- and post-gan- 

glionic fibres has not been determined. 

There are two regions of special interest in this section. The first 
is around the ninth and tenth ganglia from which the middle visceral 
nerves arise. These run in company with the arteries and are distri- 
buted to the gonads, spleen, intestine, &c. The details of the visceral 
ganglia have not been worked out, but it would seem permissible to 
refer to the tenth ganglion as the coeliac ganglion. Both sides com- 
municate with one another in this region. The second region of 
special interest is what Andersson calls the plexus iliacus^ but it seems 
preferable to use the term renal plexus^ since this avoids any confusion 
with the spinal nerves. At the point where the post-cardinal veins 
leave the post caval the sympathetic crosses their dorsal aspect and 
comes to lie between the kidney and the post-caval vein, always 
remaining ventral to the aorta. In the male the association of the 
adrenal bodies and the sympathetic with the post caval and its 
tributaries seems to be closer than in the female, a fact which may 
possibly be correlated with the greater development of the vasa 
efferentia in the former sex. Proceeding posteriorly from this point 
the longitudinal strand becomes thicker and the ganglia rather 
larger, particularly in the male. At about the twelfth or thirteenth 
ganglion the strand divides very definitely into two, and, while they 
may be connected by fine strands, they do not re-unite to form a 
single strand until the posterior end of the kidney is reached. The 
more dorsal of these strands (sym.d.) remains associated with the 
dorsal aorta. It receives the rami communicantes and is ganglionated. 
It is always connected with the ventral strand by a pair of strong com- 
missures, one lying anterior and the other posterior to the iliac 
artery. Connexions may also exist in other regions, but they are 
variable. The ventral strand (sym.v.) remains ventral to the aorta 
but dorsal to the post-caval vein, and thus continues the direct line of 
the anterior portion of the strand. It is also ganglionated and remains 
in close connexion with the adrenal bodies. At the posterior end of 
the kidney, around the bases of the superficial renal arteries, the two 
sides of these ventral strands anastomose freely, and form a plexus. 
The pair of extra large ganglia situated in this region virtually ter- 
minate the ventral strands, since a fine nerve leaves each ganglion 
and passes dorsalwards around the posterior margin of each kidney 
and joins the dorsal strand, which then enters the haemal canal and 
continues as the caudal portion. The pair of ganglia just mentioned 
apparently represent the large median cloacal ganglion described by 
Heidenhain (i 890) for Triton, and although they remain paired and 



THE NERVOUS SYSTEM 183 

are much smaller in Salamandra, it is interesting to note that nerve- 
cells do occur along the commissure joining the two sides, so that it 
is easy to see how the unpaired condition has arisen. Heidenhain 
describes the ganglion in the male only, but no appreciable difference 
between the sexes has been noticed in Salamandra. 

The sympathetic nerve for the hind-limb leaves the ventral strand 
at the plexus around the iliac artery. Fine fibres leave both the an- 
terior iliac ganglia and entwine the artery, finally uniting to form a 
single strand at the edge of the kidney. From about the point at 
which the longitudinal nerve strand divides into dorsal and ventral 
portions as described above, i.e. at about the twelfth or thirteenth 
ganglion, a fine nerve is given off which passes laterally, and appar- 
ently supplies Miiller's duct and the Wolffian duct. It is larger in the 
female. There appears to be a nerve plexus around these ducts but 
the details have not been elucidated. 

(iii) The caudal -portion is relatively simple and consists of a double 
nerve chain lying along the caudal artery. The ganglia are approxi- 
mately segmental in arrangement. Anastomoses between the two 
sides are fairly frequent — otherwise there are no special features to 
note. 

(i) The cephalic portion. After receiving the ramus communicans 
from the first spinal nerve, at about the point where the systemic 
arches join to form the dorsal aorta, the sympathetic chain follows 
each systemic arch around the pharynx, remaining rather towards 
the posterior side of the vessel. On reaching the point where the 
A. pharyngea ascendens is given off the nerve is deflected along it, 
and a portion enters the hyomandibular trunk of the facial nerve close 
to the point where the ramus communicans from NN. IX and X is 
received, and where the R. alveolaris is given off. At this point also 
a few nerve-cells occur (cf. p. 146). Another branch of the sympa- 
thetic crosses over the systemic arch and follows the A. petrosa 
lateralis into the antrum. It has not been determined precisely what 
happens to it, but presumably it enters the trigeminus ganglion, 
and, passing by way of the ophthalmicus profundus and the ramus 
communicans V ad III, reaches the ciliary ganglion on the oculo- 
motor nerve. This point, however, has not been definitely estab- 
lished. The portion of the sympathetic passing round the systemic 
arch also appears to be connected with the pharyngeal rami of the 
branchial branches of the vagus nerve — in fact this has been de- 
scribed by Druner. The presence of nerve-cells along certain rami 
of the cranial nerves, and in the ganglion copulare and the palatine 
ganglion, has already been noted (cf. pp. 141, 145, 150-2). 



1 84 THE SALAMANDER 

The appearance of the nerve during its course round the systemic 
arch is very variable. Sometimes the fibres are so diffuse as to render 
their dissection a virtual impossibility, but in cases where they are 
amalgamated into a single, or at least into a double strand, a dissec- 
tion of as much of the system as has been described above is by no 
means impossible, though not easy. An idea of the difficulty may 
be gauged from the measurements given by Hoffmann, who esti- 
mated that the diameter of the strand is at most 0-05 mm. and is 
frequently barely 0-035 mm. 

BIBLIOGRAPHY OF SECTION V 
17, 19, 21, 22, 37,45,46,52,64,65, 66, 78,94, loi, 102, 113, 114, 115, 
134, 135, 143, 144, 145, 146, 166, 187, 192, 199, 200, 201, 202, 208, 226, 
235, 236, 250, 251, 262, 291, 292, 303, 304, 305, 306, 307, 308, 323, 333, 
351, 374, 378, 404, 440, 44I5 489, 510, 516, 524, 525, 537, 543, 544, 545, 
551, 57^y 579, 615, 619, 628, 629, 630, 631, 632, 634, 656, 658, 686, 698, 
705, 725, 726, 727, 728, 731, 732, 738, 739, 759, 765, 794, 806, 807, 808, 
809,810, 828. 



VI 
THE VASCULAR SYSTEM 

BLOOD-VASCULAR SYSTEM 

I. THE HEART 

I. Historical. 

One of the earliest investigators to attempt to describe the structure 
of the heart of Salamandra is Fritsch (1869), but his work and figures 
are unsatisfactory. He omits the sinus venosus entirely, and figures 
the divisions of the truncus arteriosus wrongly. In 1873 Langer- 
hans published a brief account of the inter-auricular septum which 
he wrongly described as perforate. Nine years later (1882), Boas 
published his classic work Uber den Conus Arteriosus und die Arterien- 
bogen der AmphibieUy which included a general account of the heart as 
a whole, in both larva and adult. His work is well known and forms 
the basis of most text-book descriptions. Rose (i 890) gives a general 
comparative account of the heart in vertebrates, including Salaman- 
dra, but has few original observations to make. He agrees with 
Langerhans that the auricular septum is perforate. Langer (1894) 
made a thorough study of the structure and development of the 
bulbus cordis and truncus arteriosus, and draws a comparison be- 
tween these structures and those of fishes, particularly the Dipnoi. 
Greil (1903) has also studied the development of the truncus arteri- 
osus, of which he gives a good account. The rather considerable 
gaps which thus existed in the available literature on the anatomy 
of the heart of Salamandra have been filled recently, to a large extent, 
by Zullich (1930). His account is detailed and illustrated by a 
number of very good figures. Unfortunately he does not seem to 
have investigated the venous trunks for himself, but has copied 
Wiedersheim's erroneous figures. Nevertheless he gives a good 
account of the systolic rhythm and discusses the separation of the 
blood. 

As regards the physiological and functional aspects, the classic 
work of Briicke (1852) may be mentioned. He saw the ductus 
Botalli between the pulmonary and systemic arches, and realized that 
it was perforate. He further undertook injection experiments to try 
to find whether the venous blood was separated from the arterial, 
and came to the conclusion that very little, if any, such separation 



1 86 THE SALAMANDER 

occurred. He thought that the acute angle at which the ductus 
BotalU meets the systemic arch would cause the former to be 
closed when the pressure in the latter rose beyond a certain mini- 
mum, and thus an admixture of blood in the two arches would be 
prevented. Suchard (1902) continued and extended Brucke's in- 
vestigations. He thought that separation did actually occur but 
that it was only partial. He experimented by injecting a coloured 
fluid into the ventricle of an anaesthetized animal and found that it 
first entered the pulmonary arch, and therefore concluded that the 
first blood to leave the ventricle at the commencement of the systole 
would likewise enter the pulmonary arch. More recently, Nakano 
(19 1 3) and Haberlandt (19 16) have conducted physiological experi- 
ments on the heart of the Salamander, but their results lie rather out- 
side the scope of the present work. The experiments were mainly 
of an electrical nature and were designed to elucidate the nature and 
sources of the nerve stimuli controlling the heart movements. Ziil- 
lich's contribution to the physiological aspect has already been 
mentioned and will be referred to again on p. 191. 

2. Description (PI. XIII, figs. 59 and 60). 

The present investigation was commenced before Ziillich's paper 
appeared, and it was decided to reconstruct the anterior end of the 
ventricle, bulbus cordis, and truncus arteriosus by the wax-plate 
method in order to elucidate the details of their structure. The fol- 
lowing description is based on a study of this model and of the 
sections from which it was built, as well as on the dissection of other 
specimens. 

External Features. 

The heart of the Salamander is relatively much smaller than that 
of the Frog, while the ventricle (ven.) is shorter and has a more 
rounded apex, which is displaced a little to the right of the middle 
line. As seen in the ordinary course of dissection in a fresh or pre- 
served specimen, the anterior face of the ventricle, instead of lying 
in a transverse plane as in Rana, is inclined obliquely to the left, 
and the auriculo-ventricular opening, instead of being at about the 
same transverse level as the opening of the bulbus cordis, is posterior 
to it, while at the same time both auricles are displaced to the left. 
The bulbus cordis'^ (b.c.) arises from the right antero-ventral angle of 
the ventricle and is inclined slightly to the left. It expands distally, 
where it merges into a short, thick truncus arteriosus (t.a.), from which 
' Often incorrectly described as the truncus arteriosus, see p. 1 89. 



THE VASCULAR SYSTEM 187 

arise normally four arterial arches, although occasionally only three 
may be present. These arteries do not^ as in the Frog, leave the 
truncus as a single vessel, divided internally, but are separate from 
the beginning, and although a certain amount of connective tissue 
envelops their bases, it is clearly distinguishable from the arterial 
walls. In this connexion Ziillich is a little misleading, since he refers 
to a 'Truncus arteriosus dexter' and 'sinister' (cf. his Abb. i with 
Fig. 60 of this work). 

The auricles (or atrid)^ as already stated, are displaced to the left. . 
They are separated from the ventricle by a deep groove — the sulcus 
coronarius — which is especially apparent from the ventral side. Dor- 
sally it is covered by the sinus venosus. It is not usually possible to 
distinguish accurately, from the outside, the partition between the 
auricles — the septum atriorum — but the left auricle is considerably 
smaller than the right. The sinus venosus (s.v.) is a large triangular 
sac which may normally be seen from the ventral side, lying dorsally 
and to the left of the heart (right as viewed), although obviously the 
exact size and shape will depend mainly on the amount of blood it 
happens to contain. It is much larger than in the Frog. The apex 
of the sinus venosus is directed posteriorly and receives the post-caval 
vein, while the two ducts of Cuvier enter the basal angles. On 
removing the heart, or by turning the apex of the ventricle forwards 
so that the dorsal side may be seen, there are exposed to view the 
right ductus Cuvieri lying across the anterior rim of the ventricle, 
and the pulmonary vein passing forwards along the right margin of 
the sinus venosus to enter the left auricle. 

Internal Details. 

In the detailed description of the internal structure and relations 
of the several parts of the heart, the order followed will correspond 
with the course of the blood through that organ, namely, from the 
sinus venosus to the truncus arteriosus. 

Sinus venosus (s.v.). The cavity is undivided and the walls are 
very thin and devoid of striped (cardiac) muscle elements. It joins 
the right auricle at the anterior, right-hand, basal angle, quite close 
to the point of entry of the right duct of Cuvier, and the cavities of 
the sinus venosus and the right auricle are there confluent. The 
opening — the ostium venosum sinus (Ziillich) — is situated near the 
septum atriorum, and is guarded by a large single flap valve which 
forms an antero-dorsal outgrowth from the septum atriorum, and 
prevents regurgitation of the blood from the auricle to the sinus 
venosus. It lies to the left of the ostium. 



1 88 THE SALAMANDER 

TYvt pulmonary vein (v.pul.) lies close against the dorsal wall of the 
sinus, and anteriorly it actually lies within the wall. It enters the left 
auricle at its extreme right dorsal angle. The opening is not valved, 
so that regurgitation of the blood into the vein can only be prevented 
by the contraction of the surrounding muscles of the auricular wall. 

The auricles (aur.) are separated from one another by a septum 
composed of a muscular network covered on either side by the car- 
diac endothelium. Thus the septum within the meshes of the net- 
work is exceedingly thin, and frequently appears to consist merely 
of the double layer of endothelial cells. It is probably due to this 
construction that the older authors thought it was perforate. The 
walls of the auricles are also formed of a muscular network, but it is 
much denser, so that, although the thickness varies greatly, they are 
never so extremely thin as the septum atriorum. The inner surfaces 
of the walls of the auricles are not smooth, neither are the cavities 
entirely simple, since the muscle bands form numerous ridges, and 
even isolated trabeculae, which are detached from the wall for a 
greater or shorter distance, thus forming a number of small pits and 
cavities within the wall. This is particularly the case in the left 
auricle and the upper part of the right. Owing to this peculiar 
formation of the auricular walls certain bulges appear on the external 
surfaces of the contracted auricles which have been named 'recesses' 
by Rose. Thus the recessus sinister lies at the ventro-posterior portion 
of the left auricle, the recessus dexter is the right ventro-posterior 
part of the right auricle, the recessus intermedius is the main cavity, 
and the recessus dorsalis the right antero-dorsal portion of the same 
auricle. 

The auriculo-ventricular opening is a single oval aperture with the 
long axis in the horizontal plane, and the free edge of the septum 
atriorum stretching along the short axis in a dorso-ventral direction. 
Thus the blood from both auricles has to pass through a single 
opening into the ventricle. Regurgitation is prevented by a pair of 
flap valves, one dorsal and one ventral. Nevertheless, in the speci- 
men modelled, there is a hint of a double nature of the opening, 
since at its left end there is a shorter pair of valves, partially 
separated from the main large pair by a fold. It is perhaps not safe 
to generalize from a single specimen, and it is not practicable to 
detect this feature except from sections. In any case the valves 
must be regarded as a single pair from a functional point of view. 
Nevertheless it is interesting to note that Osawa describes two pairs 
of valves in Cryptobranchus^ where the heart is much larger and the 
valves easier to see, and they consist of one large and one small pair. 



THE VASCULAR SYSTEM 189 

The ventricle (ven.), like that of the Frog, is a muscular sponge 
with a central cavity, but the sponge is much less dense than in that 
animal, and the central cavity is comparatively smaller. From the 
work of Benninghof (1920-1) there would appear to be some order 
in the seeming chaos of muscle bands composing the sponge of the 
ventricle, but the questions raised by this author are too special to 
be entered into here. The blood in many places comes very close to 
the external surface of the ventricle, and in a freshly killed specimen 
it is clearly visible within the meshes of the sponge, as the ventricular 
wall is semi-transparent. The 'central' cavity does not extend any 
distance caudad of the auriculo-ventricular opening, and is a more or 
less L-shaped canal leading from that opening to the bulbus cordis. 
For particulars of the vena cordis see p. 1 9 1 . 

Of the rather complicated structure interposed between the ven- 
tricle and the aortae, often loosely referred to as the 'truncus arteri- 
osus', only the distal portion is actually the truncus arteriosus, homo- 
logous with the ventral aorta of the fish. The proximal portion is the 
bulbus cordis. 

The bulbus cordis (b.c.) is a long, narrow, slightly twisted tube, 
cylindrical at the ventricular end and expanding in a horizontal plane 
to a broad oval distally. It is delimited from the ventricle by the pre- 
sence of three pocket valves which prevent the reflux of the blood 
into the ventricle. When closed each forms a cup with the con- 
vexity towards the ventricle. They are non-muscular and thin-walled, 
with but few trabeculae stretching across their concavity from the 
bulbus wall. The dorsal valve is distinctly larger than the other 
two. The limit between the bulbus cordis and the truncus arteriosus 
at the distal end of the former is also marked by pocket valves, 
of which there are four — two dorsal and two ventral. They are 
also non-muscular but are V-shaped rather than cup-shaped, and 
are prolonged some distance in a caudal direction along the wall of 
the bulbus, forming ridges thereon. This is especially pronounced 
in the case of the right dorsal valve which is contiguous with the so- 
called spiral-valve — or better septum bulbi — a longitudinal septum 
partially dividing the cavity of the bulbus into right and left cham- 
bers. The septum bulbi is much more rudimentary than in the Frog 
and does not extend any distance anteriorly into the truncus, and 
it is very doubtful whether it has any influence on the separation of the 
venous from the arterial blood (cf. p. 1 91 et seq^. It passes across the 
dorsal wall of the bulbus from left posterior to right anterior. The 
distal valves frequently have small knobs of hyaline cartilage of vary- 
ing size embedded in them. They are also much thicker than the 



I90 THE SALAMANDER 

proximal row, while numerous trabeculae pass across their concave 
aspects from the wall of the truncus. The bulbus cordis is cardiac in 
origin, its muscular walls being of the same nature as, and continuous 
with, the ventricle. 

The truncus arteriosus (t.a.) represents a greatly shortened ventral 
aorta enclosed in a sheath of cardiac muscle which grows forwards 
round it from the bulbus, the tissue pertinent to the truncus itself 
being arterial tissue containing unstriped muscle-fibres. The sheath 
of cardiac muscle envelops the whole of the truncus posterior to the 
aortic arches, and it is prolonged in the middle line so as to form a 
strap passing between the carotid arches from the dorsal to the ven- 
tral side. On the dorsal side, within the cavity thus formed between 
the strap and the truncus, is the central lymph heart (cf. p. 260). 
Ziillich did not realize the true significance of the lymph heart and 
strap, but describes it as an 'elastische Polster', and says (p. 205), 
'Die funktionelle Bedeutung scheint lediglich die zu sein, das dieses 
elastische Kissen den abgehenden Arterien Schutz gegen Knickung 
gewahrt, bzw. eine nachgiebige, federnde Abstutzung gegen den 
dorsal daruberliegenden Osophagus bietet.' He does not appear to 
have been acquainted with Greil's work, although he states that the 
'Polster' contains lymph. 

As regards the cavity of the truncus arteriosus, two regions are 
discernible; a short undivided section next to the bulbus cordis — 
the truncus impar — and the main section which gives rise to the eight 
aortae by the ingrowth of septa from its walls. These septa have 
received different names from various authors. Greil's nomenclature 
is adopted here, and it is not thought necessary to give other syno- 
nyms as they are quite obvious. The septum between the carotid 
arches is the septum caroticum\ that separating the carotid from the 
systemic arch on either side the septum carotico-aorticum ; the septum 
aortico-pulmonale divides the systemic arch from the 'third' arch, and 
the latter is in turn separated from the pulmonary arch by the 
septum accessorium. The septum pulmonale separates the right pul- 
monary arch from the left. It should be noted that there is no great 
prolongation of the horizontal septum so as to divide the chamber 
into a cavum pulmo-aorticum and a cavum aortico-caroticum as there 
is in the Frog. In fact there is no well-marked difference in the 
length of any of the septa, and they may not be quite symmetrical on 
the two sides. The septum pulmonale is the longest and is closely 
followed by the septa aortico-pulmonale and caroticum, while the 
septa carotico-aortica and the septa accessoria are shorter and approxi- 
mately equal to one another in length. 



THE VASCULAR SYSTEM 191 

A F. cordis from the ventricle enters the right ductus Cuvieri on 
its ventral aspect at the right margin of the ventricle. 

3. Functional. 

The problem of whether there is any separation of the oxygenated 
and de-oxygenated blood in the Salamander is a vexed one. As 
already mentioned, Briicke — who was one of the earliest workers to 
attack the question — came to the conclusion, as the result of injection 
experiments, that in all probability no such separation did occur, 
while Suchard, who followed him, thought that there was some slight 
separation. Noble (1925), as the result ofinjecting Indian ink into the 
pulmonary veins of a number of Amphibia, both Urodeles and Anura 
— but not Salamandra — came to the conclusion that in those animals 
which possessed lungs, two auricles and a septum bulbi, the oxygen- 
ated and de-oxygenated blood remained distinct, and that the former 
never entered the pulmonary arch, whereas in those Amphibia which 
are devoid of lungs, and in which the septum bulbi and inter-auri- 
cular septum have also atrophied to a greater or lesser extent, no 
such separation could be demonstrated. He therefore ascribed to 
the septum bulhi the important function of bringing about this separa- 
tion. All injection experiments are open, to a varying extent, to the 
objection that it is impossible to execute them without causing 
traumatic shock to the heart, which is very sensitive to such stimulus. 
It is, therefore, difficult to be sure that one is obtaining a normal 
systolic rhythm under these conditions. Noble's results are con- 
sistent, and would seem to be as free as possible from this objection. 
Nevertheless they are susceptible of another explanation than that 
which he puts forward. 

Ziillich (1930) gives a detailed and tolerably complete account 
of the course of the blood through the heart in Salamandra, and con- 
cludes that while the blood from the right and left auricles does to 
a large extent remain distinct as it passes through the heart, the sep- 
tum bulbi is in no way responsible for this — a conclusion with which the 
present writer is in agreement. A careful examination of the structure 
of the bulbus cordis and truncus arteriosus makes it impossible to 
conceive that the septum bulbi can have any effect whatever on the 
separation of the two kinds of blood in the Salamander, although it 
probably has in Rana and other Anura where it extends farther into 
the truncus impar.^ 

' Since this was written a paper has appeared by Vandervael (1933) setting forth con- 
clusions different from those given here. His experiments consisted of observing the 
circulation of the blood through the heart and arterial trunks of the Frog by means of 



1^2 THE SALAMANDER 

The course of the blood through the heart Is briefly as follows. 
The left auricle fills before the right. The systolic rhythm com- 
mences with the sinus venosus, which empties itself completely into 
the right auricle, and, as a result, the inter-auricular septum is pushed 
over into the left auricle, which accordingly commences to discharge 
its contents into the ventricle. The two auricles then contract almost 
simultaneously, the left leading ever so slightly, and thus the blood 
from the left auricle passes into the proximal or left-hand side of the 
ventricle, and that from the right into the central cavity and distal 
or right-hand side. The first blood to leave the ventricle is that 
from the right auricle, then a mixture from both right and left 
auricles, and finally that from the left auricle only. Now the resis- 
tance offered by the several vessels leaving the truncus is not identical 
in each case. The greatest resistance is offered by the carotid arch, by 
virtue of its labyrinth, and the least by the pulmonary arch. As 
therefore, the systole spreads on the ventricle, the first blood to 
leave would enter the latter vessel, and the last would go to the head 
via the internal carotid artery. The difference in pressure between 
the pulmonary and systemic arches must be very slight since they are 
connected by a perforate ductus Botalli, but the actual capacity of the 
pulmonary system is relatively small, and hence it speedily fills and 
allows the greater part of the blood to flow into the other two arches. 

Noble's experimental results, which are simply that none of the 
ink injected into the pulmonary vein ever entered the pulmonary- 
artery, are thus easily explained without involving the septum bulbi 
at all. There is another important feature to be noted which seems 
to have been overlooked by previous writers, namely, that as shown 
on p. 274 the bucco-pharynx is as intimately concerned with respira- 
tion in the Salamander as the lung, and this region, although sup- 
plied mainly by the pulmonary arch, returns most of its blood into 
the jugular veins and hence into the r/^/z/ auricle, so that although the 
blood of the left side of the heart is purely oxygenated blood, that of 
the right side must also contain a considerable amount of oxygen. 
Thus, by virtue of this and the systolic rhythm, the spongy ventricle 
and the resistance of the carotid labyrinth, the Salamander possesses 

transillumination from the bulb of an ophthalmoscope lamp placed under the heart. He 
observed no perceptible difference in the colour of the blood in the three arches, and also 
found that Indian ink injected into the pulmonary vein vi^as distributed simultaneously 
and equally to each of the three arterial arches. He therefore concluded that the septum 
bulbi was not responsible for any selective distribution of the two kinds of blood, and 
in fact went further, and claimed that the blood from the right and left auricles does not 
remain distinct in its subsequent passage through the remaining portions of the heart. 



THE VASCULAR SYSTEM 193 

a mechanism whereby the brain is assured of a supply of pure 
oxygenated blood from the left auricle, and the body and respiratory 
systems are supplied with a mixed blood, of which that going to the 
former contains rather more oxygen than that to the latter system. 

This hypothesis, by discounting the importance of the septum 
bulbi of Salamandra in effecting such separation of the blood, does 
not explain why the septum should be absent from those Amphibia 
which do not possess lungs, nor indeed why it should be present in 
those which do. On the other hand it must be remembered that 
the septum bulbi is but an exaggeration of the tendency to become 
prolonged into ridges exhibited by all the distal valves. It seems 
therefore not improbable that the inception of the septum bulbi may 
be referred to similar mechanical stresses to those which, in the other 
valves of the distal series, occasion the development of small rods of 
hyaline cartilage (cf. p. 189). Whether this is so or not, a careful 
observation of the contracting heart and bulbus cordis makes it cer- 
tain that the presence of the septum enables the bulbus to empty 
itsQ^ completely of blood, and thus ensures that the last drops of pure 
oxygenated blood shall be forced into the vessels. In the case of an 
apulmonate species, where the blood in the ventricle is homogene- 
ously constituted with regard to oxygen, this complete emptying of 
the bulbus would be of much less importance. IntheAnura,e.g.Rana, 
the septum bulbi has developed more extensively, and the septum 
principale of the truncus arteriosus approaches close to its anterior 
end, so that the 'spiral valve' is able to execute the additional function 
of ensuring more effectively than pressure alone can do that the 
blood carrying least oxygen shall be delivered to the pulmonary artery. 

4. The Pericardium. 

The heart is surrounded by a coelomic space, th.Q pericardial cavity, 
which is bounded externally by a tough, fibrous, transparent mem- 
brane — th& pericardium. The cavity is completely separate from the 
pleuro-peritoneal coelom, there being no trace of any pericardio- 
peritoneal canal. 

The pericardium is attached to the heart at the anterior end of the 
truncus arteriosus, at the ductus Cuvieri close to their formation 
by the fusion of the venous trunks, and also to the dorsal wall of 
the sinus venosus and to a part of the dorsal wall of the ventricle. 
This attachment of the ventricle to the pericardium is an annular one, 
involving a relatively large area (see Fig. 59). It is also present in the 
Dipnoi and in many Reptiles^, but only as a ligamentous strand. In 
' Cf. also Hoffmann in Bronn's Thierreich, Band VI, Reptilia. 
4038 o 



1^4 THE SALAMANDER 

these latter animals Fritsch (1869) calls the ligament the 'Guberna- 
culum cordis'. It is, of course, the remains of the dorsal mesocardium 
and is frequently referred to as the mesocardial ligament. The exten- 
sive area involved by this pericardio-ventricular connexion in Sala- 
mandra must be regarded as a very primitive feature. The auricles 
are free, as is also the whole ventral surface of the heart, except at the 
anterior end of the truncus arteriosus, at the ductus Cuvieri, and at 
the post-caval and pulmonary veins. 

The pericardium has also important connexions with the MM. 
obliquus internus and rectus abdominis, which are discussed on 
p. 270 in relation to the question of the diaphragm. 

II. BLOOD-VESSELS. 
I. Historical. 

Funk (i 827) is neither accurate nor detailed regarding the blood- 
vessels — a situation that Rusconi (1854) made a special point of 
amending. Rusconi's figures of the aortic arches in both larva and 
adult are very good and are reproduced by Hoffmann in Bronn's 
Thierreich. He does not describe the remainder of the arteries in 
any detail, but he gives some account of the main veins, and is the 
first to describe the vessel derived from the left vitelline vein which 
now bears his name, but was called by him the longitudinal gut vein. 
Jourdain (1859) gives a good description, with figures, of the renal- 
portal system, but he figures the connexion between the V. iliaca 
communis and Jacobson's vein wrongly. 

The first investigator to study the vascular system of the Salaman- 
der apart from the general anatomy was Hochstetter (1888). He 
gives a very good account of the veins of the abdomen and of the 
renal-portal system, but does not deal with the details of the cephalic 
vessels nor with those of the limbs. He recognized the true nature of 
the post-cardinal veins (Rusconi had called them the veins of the 
oviduct), and his work on the development of these vessels and of 
the abdominal vein is too well known to need comment. In 1 894-5 
Zuckerkandl, in his work on the anatomy and development of the 
arteries of the fore-arm, deals with Salamandra as a type of Urodeles, 
but his description is brief. Bethge (1898) supplements Hoch- 
stetter's account to some extent in a paper entitled 'Das Blutgefass- 
system von Salamandra maculata, Triton taeniatus, und Spelerpes 
fuscus'. He is the first to mention the vertebral artery — A. verte- 
bralis collateralis — and the lateral vein — V. cutanea magna — but 
on the whole his description is very general. He is more concerned 



THE VASCULAR SYSTEM 195 

with the respiratory function of the vessels he is describing than with 
their exact anatomical relations. He improves on existing accounts 
by giving some description of the vessels of the head region, but in 
this he is not quite accurate. He figures the lingual vein as passing 
ventral to the arterial arches and says that it breaks up into a 'Rete 
mirabile' in the region of these vessels. 

The only vessel that can be identified with the vein he describes 
is the V. thyroidea^ while the 'Rete mirabile' must be the thyroid 
gland itself. It is astonishing that he should have fallen into this 
error, as the development of the thyroid gland had been well de- 
scribed only four years previously by Maurer, but his figure allows no 
other explanation, since the thyroid vein is the only vein in that 
region passing ventral to the aortae. Bethge accurately figures and 
describes the proximal portion of the true lingual vein — which 
passes dorsal to the arterial arches — as the V. pharyngea. 

Only two more authors need be noted here, namely, Choron- 
shitsky (1900) who gives a good account of the development of the 
mesenteric veins and of the hepatic portal system, and Konigstein 
(1903) who describes the pulmonary and related arteries. 

The vessels of the brain were first described by Scholb in 1882. 
His figures are very beautiful but not quite accurate, particularly 
in respect of the veins, and his work was amended later by Rex 
(1893). 

2. General Features. 

The condition of the blood-vascular system in the Salamander is 
extremely interesting, and it is surprising that no really adequate 
account of it exists, for we have in one and the same animal what 
amounts to two systems of blood-vessels — that of the fish and of the 
land vertebrate, and particularly is this so in the case of the venous 
system. The Salamander thus forms a nearly perfect transitional 
type, being in this respect much less specialized than the Frog. 

The chief features of interest may be tabulated thus: 

Arterial System. 

(i) Persistence normally of four arterial arches in the adult, 
(ii) Occurrence of a perforate ductus Botalli between the pul- 
monary and systemic arches, 
(iii) Limitation of the area served by the carotid arch, 
(iv) Distribution of a branch of the systemic arch to the palate, 
(v) Distribution of branches of the pulmonary arch to the 
oesophagus and pharynx. 



196 THE SALAMANDER 

Venous System. 

(i) Persistence of the post-cardinal veins, which may even in 

some cases retain their primitive connexions with Jacobson's 

veins, 
(ii) The persistence of one of the vitelline veins of the embryo — 

the left — in a perfectly functional condition (see Rusconi's 

vein, p. 238). 
(iii) The obvious derivation of the venous system of the head 

from Selachian-like sinuses, 
(iv) The occasional absence of the middle portion of the post- 
caval vein, and corresponding enlargement of one of the 

post-cardinals. 

As might be expected in an animal retaining so many primitive 
features there is a considerable amount of individual variation to be 
met with, and it is difficult to give a general account which will cover 
all cases, but it is hoped that the following description, based mainly 
on the dissection of specimens which have been injected with 
Prussian blue (gelatine and liquid) injection masses, will enable all 
the important vessels to be identified and followed. 

All vessels may be taken as paired unless definitely stated to be 
otherwise. 

The vessels of the cranial and vertebral cavities, both arteries and 
veins, are described at the end of the present main section under 
'Vessels of the Brain'. 

3. Arterial Arches (PL XIV). 

The Salamander, after metamorphosis, normally retains all four 
of the branchial arteries present in the larva, and rarely three only 
are to be found. In the latter condition it is the 'third' arch which is 
missing, its place being occupied by a strand of pigmented connec- 
tive tissue bearing the same relation to the M. cephalo-dorso-sub- 
pharyngeus and to the systemic arch as the artery does when present. 
The other arches present are — naming from anterior to posterior — 
(i) the carotid^ (2) the systemic^ and (4) the. pulmonary. 

The carotid — third visceral — arch (A. Ill) is not normally con- 
nected with the systemic by a ductus Botalli, but in one specimen 
such a connexion has been found on one side. It was exceedingly fine, 
however, and only just visible under the dissecting microscope. 

The systemic — fourth visceral — arch (A. IV) is considerably larger 
in diameter than the other arterial arches while it also receives the 
blood leaving the heart by the 'third' arch. Its branches deliver blood 



THE VASCULAR SYSTEM 197 

to nearly the whole of the body except the brain, the lungs, and the 
pharynx. 

The third — fifth visceral — arch (A. V) varies considerably in size. 
Occasionally it is absent, or it may be quite well formed and compar- 
able in diameter with the common carotid and pulmonary arches, but 
normally, however, it is much smaller. It leaves the truncus arteri- 
osus somewhat postero-ventrally relative to the systemic arch, and, 
running parallel with the latter, ultimately fuses with it at the level 
of the dorsal side of the pharynx, just proximal to the entry of the 
ductus Botalli. 

The pulmonary — sixth visceral — arch (A. VI) in the Salamander 
supplies the walls of the pharynx as well as the lungs. It does not run 
quite parallel with the other arches, but after leaving the truncus 
arteriosus it turns at first slightly posteriorly, then bends anteriorly 
to the anterior border of the M. dilatator laryngis, at which point the 
ductus Botalli is given off. The latter passes mesially to the muscle 
and enters the systemic arch just distal to the point where the 'third* 
arch enters. The ductus Botalli is perforated by a functional lumen, 
and in fact one or two small cutaneous branches are given off from 
it laterally, as well as the lateral pharyngeal artery from its mesial 
side. The pulmonary arch then turns sharply posteriorly again 
towards the lung. 

4. Branches of the Carotid Arch (Common Carotid) (PI. VI, figs. 
37, 38, and PI. XIV). 

(Note. — The terms ''Internal'' and ^External' are applied to the two divisions 
of the Common Carotid for sake of convenience^ although their distribution is much 
more restricted than is the case in the Frog.) 

The common carotid emerges from the truncus arteriosus some- 
what antero-dorsal to the systemic arch and passes laterally for about 
4 or 5 mm. to the carotid gland, where it divides into its external and 
internal branches. 

A. The external carotid (a.car.e.) artery leaves the gland on its 
mesial side anterior to the entry of the common carotid, and runs 
back parallel with the latter for a millimetre or so before turning 
anteriorly. From this section of the vessel arises: 

(i) Ramus muscularis for the MM. rectus cervicis and interhyoideus 
posterior. This branch divides almost immediately, so that the 
branch for the former muscle passes mesially, dorsal to the thyroid 
vein, while the other enters directly into its muscle, which covers this 
neighbourhood ventrally. 



198 THE SALAMANDER 

After turning anteriorly the carotid artery passes along the lateral 
border of the thyroid gland to which it sends : 

(ii) The thyroid artery — a short but fairly broad vessel which breaks 
up immediately within the gland into a rich network. 

On reaching the level of the second hypobranchial cartilage the 
main artery divides into three (sometimes four) branches as follows : 

(iii) The sublingual artery (a.subl.), the more superficial of the two 
anterior branches, passes along the lateral border of the geniohyoid 
muscle to within about 3 mm. from the tip of the lower jaw. Here it 
divides into four rami. 

(a) R. Ungualis which passes mesially and enters the tongue 
together with the hypoglossal nerve. 

{¥) R, muscularis which passes anteriorly and enters the M. genio- 
glossus. 

(c) R. communicans which passes laterally and sends a branch to 
the M. subhyoideus, then turning anteriorly and passing to the 
middle line round the lower jaw, where it may fuse with its fellow on 
the other side, although it may also anastomose with the mandibular 
artery. Whether these anastomoses always occur is uncertain, since 
failure to find them in every case may rather be due to faulty in- 
jections than to the absence of the vessels. They are very fine vessels. 

{d) R. intermandihularis arises soon after the origin of the sub- 
lingual artery at the level of the first hypobranchial cartilage and 
passes ventrally into the underlying muscle of the same name. 

(iv) The lingual artery (a. ling.), the other anterior branch, runs more 
or less parallel with the previous artery but deep to the geniohyoid 
muscle. It enters the tongue at two or three points between the 
radials of the copula, i.e. together with the glossopharyngeal nerve. 

(v and vi). Two muscular branches which pass laterally to the 
lateral edge of the M. subarcualis rectus I, which they supply. These 
two branches may be represented at their origin by a single vessel for 
a greater or shorter distance. 

All the branches of the external carotid artery are somewhat con- 
voluted and very extensible — a necessary feature in view of the large 
amount of movement to which the parts they supply are subjected. 
They are only loosely held by connective tissue and are easily dis- 
sected. The external carotid artery as a whole has been called the 
Arteria lingualis by some authors — e.g. Hoffmann — but this would 
seem to be a misnomer. 

B. The internal carotid artery (PL XIII, fig. 58 ; PI. XIV, fig. 6 1) 
(a.car.i.) continues the direction of the common carotid as far as the 
floor of the otic capsule. Here it divides, giving off the A. petrosa 



THE VASCULAR SYSTEM 199 

lateralis (a.pet.l.) (Driiner), which turns dorsalwards around the 
posterior border of the processus basilaris and enters the antrum 
petrosum laterale by the 'faciaHs' opening (see below). The remain- 
ing branch of the internal carotid then bends somewhat mesially 
across the ventral face of the Proc. basilaris and enters the carotid 
canal (Fig. 2, can. car.) — a groove in the floor of the cranium covered 
ventrally by the parasphenoid bone. There is no definite 'carotid 
foramen' in the Salamander's skull. The artery enters the cranial 
cavity through the hasicranial fenestra and immediately gives ofl^ the 
A. carotis cerehralis (a.car.ce.) (Scholb.) — see 'Vessels of the Brain'. 

The remainder of the vessel turns anteriorly along the edge of the 
basicranial fenestra as far as the foramen for the N. oculomotorius. 
It then penetrates this foramen and passes into the orbit as the A. 
ophthalmica (see next page). 

(i) A. petrosa lateralis (Driiner) (a.pet.l.). This small but important 
vessel is probably to be regarded as the homologue of the stapedial 
artery of Mammals. As already described it passes into the antrum 
peirosum laterale around the posterior border of the processus basi- 
laris, mesial to the corresponding vein, and ventral to the columella. 
Within the antrum the artery loops over the dorsal edge of the vein 
so as to lie lateral to it. It leaves the antrum between the Proc. 
oticus and the Proc. ascendens together with the mandibular and 
maxillary branches of the Vth nerve, and divides into the following 
two main branches: 

{a) A. temporalis (a.t.), which accompanies the second division of 
the trigeminal nerve to the posterior angle of the eyelids, and here 
divides into two equal branches, one of which, the superior orbital 
artery^ passes around the upper eyelid and anastomoses with the R. 
ophthalmicus superior posterior of the A. ophthalmica. The other 
turns ventrally and supplies the lower eyelid and the tissues of the 
posterior end of the upper jaw, and represents a portion of the inferior 
orbital artery (a.o.i.), the other part being represented by a branch of 
the A. mandibularis. 

{F) A. mandibularis (a.m.). This branch of the A. petrosa lateralis 
turns ventralwards and accompanies the mandibularis branch of the 
trigeminus. Very soon after its origin it gives off a branch anteriorly, 
which passes through the levator mandibulae muscle and is dis- 
tributed to the roof of the mouth at the postero-ventral corner of the 
orbit and also to the inflected portion of the skin which occurs in the 
coronoid region. This branch has not always been found, but failure 
to find it is more probably due to faulty injections than to its absence. 
It is very easily damaged in removing the muscle through which it 



200 THE SALAMANDER 

passes, and consequently may sometimes be overlooked. It can only 

be regarded as forming a portion of the inferior orbital artery. 

In this connexion it is interesting to compare the relations found 
in the human embryo and in Sphenodon (see O'Donoghue, 1921, 
p. 192). In the human embryo the inferior orbital artery is given off 
from the mandibular branch of the stapedial artery, while in Spheno- 
don it is one of the two end branches of the A. temporalis. In Sala- 
mandra both these conditions are present in the one individual. 

The mandibular artery proper continues ventro-laterally through 
the levator mandibulae muscle close to the nerve, and, like the nerve 
itself, divides on reaching the lower jaw into a R. lahialis (Osawa) 
(a.lab.), to the skin on the lateral aspect of the posterior portion of 
the lower jaw, and ai^. aheolaris (Osawa) (a.alv.), which passes through 
the inferior dental canal within the jaw itself to supply the teeth, &c. 

In addition to the two main branches just described the A. petrosa 
lateralis gives one or two RR. musculares to the several portions of 
the M. levator mandibulae. 

Abnormalities. Two rather interesting anomalies have been met 
with in this region, viz : 

(i) A case in which the A. petrosa lateralis occurred as a branch 
of the A. pharyngea ascendens instead of the internal carotid, the 
latter vessel giving off no branch at all before entering the cranial 
cavity. The specimen was well injected, and the relations were quite 
distinct, while the branches of the A. petrosa lateralis were normal 
and both sides of the specimen were alike. 

(2) A case in which a branch from the A. petrosa lateralis was 
given off almost immediately after its origin from the internal carotid 
to the lateral pharyngeal wall. This is the only trace that has been 
found of a pharyngeal artery stated by Bethge to occur as a normal 
branch of the internal carotid. Unfortunately the specimen was not 
injected, and the vessel was nearly empty of blood, so that its exact 
extent and relations could not be determined (see Fig. 58). 

As already described the internal carotid artery terminates within 
the orbit as the A. ophthalmica (a.oph.). After leaving the cranial 
cavity by the oculomotor foramen the ophthalmic artery passes along 
the optic peduncle, and a short distance from the bulbus oculi divides 
into four or five branches. It sends two or three branches to the eye 
itself, and a reflex muscular branch into the MM. retractor bulbi and 
rectus posterior. The main portion of the artery turns dorsalwards 
and divides into three vessels : 

(i) R. muscularis^ which curls around the anterior aspect of the 



THE VASCULAR SYSTEM 201 

optic peduncle and is distributed to the MM. rectus superior, 
anterior, and inferior. 

(ii) R. ophthalmicus superior posterior (a.oph.s.p.). This branch 
passes directly dorsalwards and is distributed to the upper eyelid and 
the skin above the skull in the posterior orbital region, as well as to 
the tissue surrounding the eye itself. It fuses with the superior 
orbital branch of the A. petrosa lateralis. 

(iii) R. ophthalmicus superior anterior (a.oph.s.a.). The third ter- 
minal branch of the ophthalmic artery passes dorsalwards together 
with the previous one and is distributed to the anterior portion of the 
upper eyelid and the skin of the head, as well as to the general tissues 
around the eye. It passes round the orbit along the dorsal border of 
the orbito-sphenoid and ultimately fuses with a branch of the palato- 
nasal artery and enters the nasal capsule. In addition it supplies both 
the oblique muscles of the eye. 

5. Branches of the Systemic Arch (PL XIV). 

The systemic arch runs in company with the carotid arch in an 
antero-dorsal direction to the occipital region of the skull, and then 
bends round mesially to unite with its fellow from the other side in 
the middle line to form the dorsal aorta (q.v.). The vessels given 
off from this arch are four in number. 

A. A cutaneous branch passes dorso-laterally between the two por- 
tions of the M. cephalo-dorso-subpharyngeus to which it gives a 
small branch. It is distributed to the thymus and paratoid glands. 
This vessel arises very close indeed to the point where the 'third' 
arch enters the systemic, and in one specimen it appeared to be 
actually continuous with this vessel. In view of the fact that it 
divides the pharyngeal constrictor into two sections dorsally, just as 
the 'third' arch does ventrally, it would appear to be actually the 
peripheral continuation of this arch, but only an investigation of the 
conditions prevailing at the period of metamorphosis would settle 
this point. 

B. A. pharyngea ascendens (Osawa) (a.ph.asc). 

A. maxillaris externa Bethge (1898). 

A. mandibulo-jugularls Druner (1904). 

This artery arises laterally close to the previous one at about the level 
of the posterior end of the ceratohyal and passes antero-laterally, 
mesial to the M. depressor mandibulae, to the posterior edge of the 
squamosal. It has six branches: 

(i) RR. cutanei dorsales (Osawa) of which one passes lateralwards 



202 THE SALAMANDER 

between the M. depressor mandibulae and the squamosal to the over- 
lying skin and the anterior region of the paratoid, while the other 
passes posterior to the muscle and is distributed to the paratoid 
gland. 

(ii) R. hyomandibularis (? R. branchialis, Osawa) (a.hym.) which 
passes lateralwards between the depressor muscle and the squamosal 
to the surface of the muscle, when it turns, coursing posteriorly 
across the muscle (in company with the N. jugularis — Vllth nerve), 
to its posterior edge. It then turns round anteriorly and mesially to 
be distributed to the dorsal surface of the M. interhyoideus pos- 
terior, while a further branch of this artery forks around the M. 
interhyoideus and is distributed to the posterior region of the lower 
jaw and the tissue directly mesial to it. 

(iii) R. mandibularis externus (a.m.e.) which passes to the surface 
together with the previous branch, but then turns anteriorly and 
passes mesial to the jugal tendon, along which it sends a branch, 
while the remainder of the artery is distributed to the lateral surface 
of the lower jaw. It is a very small vessel, very much smaller than 
the artery of the same name in the Frog. 

(iv) R. pharyngeus which passes postero-ventrally to the posterior 
end of the ceratohyal, and then loops forwards over the mouth-floor. 
This artery seems very variable in size and has not always been found. 

(v) R. mandibularis internus which passes antero-ventrally to the 
mesial surface of the lower jaw. It is distributed to the skin of the 
ventral surface of the head, to the MM. intermandibularis and inter- 
hyoideus, passing right round the lower jaw to anastomose with its 
opposite fellow in the middle line and also with the ramus communi- 
cans of the sublingual artery (q.v.). It usually arises combined with 
the mandibularis externus for a greater or shorter distance. 

(vi) A small muscular branch passing into the mesial surface of the 
depressor mandibulae. 

It will be noticed that there is a striking resemblance between the 
distribution of the A. pharyngea ascendens, as described above, and 
that of the R. auricularis of the A. cutanea magna of the Frog, as 
described by Gaupp in his Anatomic des Frosches. The A. cutanea 
magna, however, is a branch of the pulmonary arch, while the A. 
pharyngea ascendens arises from the systemic arch. Nevertheless its 
origin is very close to the point where the ductus Botalli from the 
pulmonary enters the systemic arch. In the Frog there is no con- 
nexion between the pulmonary and systemic arches in the adult, but 
there is in the larva. In view of these facts, therefore, and bearing in 
mind that small cutaneous branches do actually arise from the ductus 



THE VASCULAR SYSTEM 203 

Botalli In the Salamander, it seems not improbable that the A. cutanea 
magna of the Frog may be the morphological equivalent of the 
ductus Botalli+the A. pharyngea ascendens of Salamandra. The 
differences in detail between the branches of the A. pharyngea ascen- 
dens and those of the R. auricularis may easily be accounted for 
by the difference in the structure of the auditory organ in the two 
animals. 

C. The -palato-n a sal artery (PI. XIX, fig. 70) (a.pal.nas.) is, in the 
Salamander, a branch of the systemic arch. It arises dorsally close to 
the middle line, in the region of the occiput, and either close to, or 
jointly with the vertebral artery. Almost immediately it gives off a 
small twig to the dorsal wall of the pharynx, and then bends round 
laterally, following the curve of the systemic arch as far as the palatine 
foramen, when it turns sharply round mesially to the inner border 
of the orbit. At this bend four branches arise. 

(i) RR. pharyngei^ forming a rich network over the dorso-lateral 
pharyngeal wall. Usually two in number. 

(ii) R. -pterygoideus (a.pt.), a small branch passing along the ptery- 
goid process of the palato-pterygoid and shortly after, at a point ven- 
tral to the origin of the M. retractor bulbi a maxillo-palatine artery 
is given off. 

(iii) A. maxillo-palatina (a.max.pal.), arising laterally and running 
in company with the maxillary vein. It is distributed to the roof 
of the mouth underlying the postero-lateral corner of the orbit and 
then turns anteriorly and is continued a short distance along the 
posterior region of the maxilla. 

(iv) The posterior palatine artery, arising mesially and ramifying 
over the whole of the posterior region of the palate. 

At the inner posterior angle of the orbit the palato-nasal artery 
turns anteriorly and gives off the anterior palatine artery and then 
continues forward as the orbito-nasal artery. 

(v) The anterior palatine artery (a. pal. a.) also turns anteriorly 
and runs alongside the palatine vein and nerve supplying the mucous 
membrane of the middle and anterior regions of the roof of the 
mouth. The rich vascular network around the vomero-palatine teeth 
is also supplied by this artery. 

(vi) The orbito-nasal 2iYte.Yy must not be confused with the orbito- 
nasal of the Frog, which runs dorsal to the eye-muscles. It passes 
obliquely in an antero-dorsal direction across the surface of the orbito- 
sphenoid to the dorsal side of the inner anterior angle of the orbit, 
where it gives off a small twig, viz. : 



204 THE SALAMANDER 

(a) R. descendens which passes vertically downwards and spreads 
over the palate in that region. 

The orbito-nasal artery then passes through a foramen in the pro- 
cessus antorbitalis, and divides into two terminal branches. 

{}}) A mesial nasal artery which immediately divides again into 
dorsal and ventral rami which bifurcate and spread around the 
mesial border of the nasal capsule. 

{c) A posterior nasal artery (a.na.p.) which passes round the orbital 
border of the nasal capsule. Soon after its origin it gives off four 
twigs as under. 

(i) The anterior orbital artery which passes backwards out of the 
same foramen as that by which the main artery (i.e. the orbito-nasal) 
entered. It is distributed to the pre-orbital glands of that region and 
also sends a branch to the overlying skin, and another which anasto- 
moses with the dorsal branch of the ophthalmic artery, that is, with 
a branch of the internal carotid (see p. 201). 

The posterior nasal artery then proceeds to the lateral border of 
the nasal capsule where it divides into the remaining three twigs. 

(2) The lateral nasal artery (a.na.l.) which supplies the lateral 
border of the nasal capsule. 

(3) The mesial fnaxillary artery (a.max.m.) which turns posteriorly 
and supplies the middle region of the upper jaw. 

(4) The anterior maxillary artery (a. max. a.), turning anteriorly to 
supply the upper jaw in the region of the snout. 

It will doubtless have been noticed that the vessel which has 
just been described as the palato-nasal artery bears a strong re- 
semblance to one described by Gaupp for the Frog, and called by 
him the arteria occipitalis. Particularly is this so as regards its origin 
and its nasal branches. This agreement was considered by Bethge to 
be sufficiently close to justify his adopting the same name for the 
vessel in the Salamander, viz. arteria occipitalis. The A. occipitalis 
of the Frog, however, is a dorsal vessel, its orbito-nasal branch pass- 
ing 'iiber den M. rectus oculi superior (Gaupp), while in this animal 
also the palate is supplied by the palatine artery from the internal 
carotid. In view of these facts a non-committal name based on its 
distribution has been adopted here. 

D. Vertebral artery (a.vert.) {arteria vertebralis collateralis^ Bethge). 
It arises from the systemic arch close to, or together with, the palato- 
nasal artery just lateral to the occipital condyles, and, after running a 
short distance caudalwards, turns sharply in towards the vertebral 
column and then runs close alongside it posteriorly right down the 
body. It commences at about the level of the first vertebra, and 



THE VASCULAR SYSTEM 205 

passes, ventral to the spinal nerves, through special foramina in the 
transverse processes of the ribs (see p. 19). Since the artery receives a 
number of reinforcements from the dorsal aorta (q.v.), which in the 
lumbar region are quite as strong or stronger than the original artery, 
it is probably more correct to regard the vessel as a series of segmental 
arteries which have fused with one another, rather than as a single 
vessel originating at the base of the skull. The latter view is, how- 
ever, more convenient for descriptive purposes. A number of lateral 
vessels are given off to the muscles of the back. 

(i) Costal arteries arise alongside or within the transverse processes 
of the vertebrae. In the latter case they soon emerge, after giving off 
a few small twigs to the Haversian canals of the bone, to run parallel 
with the ribs. 

(ii) Intercostal arteries arise about midway between the previous 
ones, that is approximately opposite articulations between the verte- 
brae. They do not run directly lateralwards but in an oblique pos- 
tero-lateral direction, the obliquity becoming more marked in the 
lumbar region. They are also distributed to the muscles of the back. 

(iii) Cutaneous branches arise chiefly from the costal arteries and 
pass directly dorsalwards to supply the skin of the back, also giving 
a branch to the large cutaneous poison glands which occur along the 
mid-dorsal line. The capillary network forms a sort of vascular cup 
around the bases of these glands. 

(iv) Occasional anastomoses with the intraspinal arterial system. 

It must be observed that the arrangement of costal and intercostal 
vessels above described is not absolutely rigid. Many intermediate 
conditions are found, but on the whole these would seem to conform 
to a basic plan in which have been interpolated secondary additions. 

In contradistinction to the Frog it should be noted that the verte- 
bral artery is not responsible for the blood-supply to the ^'^;//r(^/ body- 
wall, since the epigastric artery (q.v.) discharges this function. The 
arteriae ahdominales of the Frog are therefore not present in the Sala- 
mander as factors of the vertebral artery. 

The vertebral artery also takes some share in the blood-supply of 
the anterior region of the oviduct, but, since these vessels usually 
arise from anastomoses with the dorsal aorta, they will be dealt with 
under the branches of the latter vessel together with the other ovi- 
ducal arteries. 

The Dorsal Aorta. 

The dorsal aorta (d.a.) arises, as previously described, by the fusion 
of the paired systemic arches in the mid-dorsal line immediately 



2o6 THE SALAMANDER 

ventral to the base of the skull in the region of the occiput, that is 
some seven millimetres or so anterior to the origin of the subclavian 
arteries, and a very considerable distance before the mesenteric 
vessels are given off. The fusion of the aortic arches in the Sala- 
mander is thus much more complete than in the Frog and there 
is no question of any asymmetrical origin of the coeliaco-mesenteric 
vessels. 

Branches of the Dorsal Aorta 

(PI. XIV, fig. 6i ; and PL XVII, figs. 64 and 64a) 

These may be arranged in three groups according to their distri- 
bution, as follows: 

Group I. Vessels of the Alimentary Canal (all of which leave 
the aorta in the mid-ventral line). 

(i) AA. gastricae anteriores (a.g.a.) (Hyrtl, Osawa, Bethge). One or 
two small vessels arising at about the level of the subclavian arteries, 
and passing obliquely backwards through the mesentery to their 
distribution over the dorsal wall of the stomach. 

(ii) A coeliaco-mesenteric complex^ comprising one large vessel, 
which may be called the coeliaco-mesenteric artery, and a series of 
smaller ones arranged in 'anterior' and 'posterior' bundles. 

The coeliaco-mesenteric artery (arteria intestinalis, Hochstetter) is 
a very large vessel almost equal to the diameter of that from which 
it springs. It arises at about the level of the tenth vertebra and oppo- 
site the posterior end of the stomach, immediately posterior to the 
junction of the post-caval and post-cardinal veins, and then passes 
ventrally through the mesentery to supply the stomach, spleen, pan- 
creas, liver, duodenum, and the anterior section of the intestine, by 
means of the following four branches. 

{a) A. gastrico-lienalis (a. g. lien.) or A. gastrica posterior (Hyrtl, 
Osawa) which, as its name implies, is distributed to the spleen and 
stomach. Its course runs anteriorly through the mesentery separat- 
ing these two organs, more or less parallel with the vein of the same 
name, but nearer the stomach than the latter vessel. It supplies the 
dorsal stomach wall only, by means of a series of branches arising at 
intervals of about a millimetre along its length, and approximately 
at right angles to it. The spleen is supplied by a similar series of 
branches arising from the opposite side of the vessel. 

(^) The A. duodeno-hepatica (a.d-h.) or A. mesenterica prima (Hyrtl, 
Osawa) which passes ventrally through the distal edge of the pan- 
creas alongside the portal vein. Very soon after its origin one or two 



THE VASCULAR SYSTEM 207 

branches arise which are distributed to the distal end of the duo- 
denum, while a few small vessels pass into the pancreas from the 
main artery during its passage through the gland. The next impor- 
tant division of the vessel occurs about 5 or 6 mm. farther on when 
it bifurcates into dorsal and ventral hepatic arteries. The dorsal 
hepatic artery passes straight to the posterior end of the liver and 
then turns anteriorly along the dorsal surface of this organ on the 
right of the vein formed by the union of the abdominal and portal 
veins. Besides supplying the liver this vessel also sends branches to 
the ventral stomach wall, which pass along the mesentery together 
with the factors of the above-mentioned vein returning the blood 
from this section of the stomach-wall. There are usually two arterial 
branches to each venous factor. A small cystic artery to the gall- 
bladder also arises from this branch. 

The ventral hepatic artery sends numerous branches to the duo- 
denum in the region of the bile-duct, and then loops round the duct 
to enter the liver immediately ventral to it. 

(/) The A. duodeno-pancreatica (a.d-pan.) or A. mesenterica prima 
(Hyrtl, Osawa) which passes through the pancreas proximal to the 
portal vein, and besides giving off considerable branches to the gland 
itself is distributed to the proximal region of the duodenum. 

id) The mesenteric branches (a.mes.a.) of the coeliaco-mesenteric 
artery which vary from two to four in number. They leave the artery 
posteriorly, the last one coming off very soon after its origin from 
the dorsal aorta, and are distributed to the proximal portion of 
the intestine. 

After the large coeliaco-mesenteric vessel just described there 
arise from four to eight anterior mesenteric arteries (a.mes.a.) (AA. 
mesenteriae accessor'iae — Osawa). These are given off serially from 
the section of the dorsal aorta corresponding approximately with the 
eleventh to the fourteenth vertebrae inclusive, and supply the distal 
portion of the intestine as far as the rectum, frequently spreading 
over the anterior end of this also. The interval between these vessels 
is only about a millimetre or a little more, and they tend to run 
obliquely forwards through the mesentery so as to be gathered to- 
gether into a bundle just posterior to the coeliaco-mesenteric artery. 
They then bend back posteriorly to their distribution. 

Distinguishable from the foregoing only by the fact that they arise 
after an interval of about 5 mm., and that they are not grouped quite 
so closely into a bundle, are about three posterior mesenteric arteries 
(a.mes.p.). They enter the dorsal side of the rectum. The interval 
between the origins of these vessels is rather greater than between 



2o8 THE SALAMANDER 

the anterior arteries above referred to. They are obviously equivalent 
to the AA. mesentericae posteriores of Osawa. 

The distinction between anterior and posterior mesenteric arteries, 
as already indicated, must be largely arbitrary, the whole obviously 
forming a serial complex. It is for this reason that they have been 
regarded as regions of a 'coeliaco-mesenteric complex'. 

Group 2. Vessels of the Urino-genital System. 

(i) Ovarian or spermatic arteries. The arteries to the gonads are 
essentially the same in both sexes. They consist of eight or ten fine 
vessels, i.e. four or five to either side, which arise mesially from the 
ventral side of the dorsal aorta and pass laterally towards their re- 
spective gonads through the mesovarium or mesorchium. Those of 
the right side are more anterior than the corresponding ones of the 
left owing to the asymmetry of the gonads, but in general they arise 
from the same section of the aorta as the anterior mesenteric vessels. 
They pass mesially to the kidneys, and tend to bifurcate very soon 
after their origin into two parallel but fairly widely separated vessels, 
which may occasionally bifurcate again. They frequently run in com- 
pany with a factor of the post-caval vein. The fat-bodies are supplied 
by branches from these arteries. 

(ii) The renal arteries (a.r.) (AA. renales — Osawa) consist of a 
number of extremely small vessels which arise from the aorta itself, 
from the oviducal, ovarian or spermatic arteries, or even from the 
dorso-lumbar arteries. At the posterior end of the kidney are a 
pair of much larger superficial renal arteries (a.r.s.) (A. suprarenalis 
— Osawa) which spread over the ventral face of the kidneys, their 
ultimate branches being connected with a number of small Mal- 
pighian bodies around the postero-lateral margin. The small deep 
renal arteries enter the kidney from the dorsal side. 

(iii) The uterine or oviducal arteries. While these are essentially 
vessels of the female sex they are also present in the male, and are 
then distributed to the vestigial Miillerian ducts and ureters, so that 
they are strictly homologous in the two sexes. They are, however, 
very much smaller in the male than in the female. The following 
description is based on the conditions in the female. They arise in 
three regions, but are not entirely symmetrical on either side. 

(a) The posterior oviducal arteries (a.od.3) arise from a common 
stem which leaves the dorsal aorta in the midventral line between 
the posterior lobes of the kidneys, just posterior to the left iliac artery. 
They are distributed to the expanded posterior end of the oviduct or 
'uterus', passing along its ventral side and giving off numerous fine 



THE VASCULAR SYSTEM 209 

branches which anastomose frequently, forming a fine vascular net- 
work, amounting almost to a 'rete mirabile'. 

Besides giving off the posterior oviducal arteries their common 
stem also gives rise to one or both superficial renal arteries, and to a 
pair of mesial do acal 3.rtenes (Figs. 64 and 64a) (a.cl.m.). These are 
essentially vessels of the male sex although also present in the female. 
In the male, these vessels, together with the superficial renal arteries, 
are the principal branches of the common stem, while the only 
vessels which may be considered the homologues of the oviducal 
branches of the female are two very small twigs arising from the 
superficial renals to the ureters and Miiller's duct. In the female 
it is the cloacal branch which is vestigial. 

In the male (Fig. 64a) after giving off the renal vessels, the cloacal 
artery proceeds dorso-posteriorly to the edge of the M. caudali- 
femoralis, to which it sends a branch, and it also gives a branch to the 
muscles of the back. The artery then turns sharply ventralwards 
along the anterior aspect of the cloacal gland, giving off two large 
branches which enter the gland mesially. A small continuation of 
the vessel finally anastomoses with a cloacal branch of the vesicular 
artery (q.v.). 

The drawings (Figs. 64 and 64a) have been carefully made from 
a single specimen of each sex. Small individual variations from them 
may therefore be expected. 

(b) The middle oviducal arteries (a.od.2) usually arise from the 
same section of the aorta as the posterior mesenteric vessels, but 
their exact point of origin varies considerably. They are usually 
asymmetrical, but either side may be the more anterior. From what- 
ever level they may arise, however, they always run parallel with the 
aorta for a greater or shorter distance before turning lateralwards, 
dorsal to the kidneys, to pass via the mesentery to the oviduct 
at approximately the same level. Their branching commences within 
the mesentery before reaching the oviduct. Although they are vessels 
of considerable size they appear to supply only a comparatively short 
section of the latter organ, namely, just anterior to the vascular net- 
work of the posterior arteries. Renal arteries are given off by these 
vessels from the section which runs parallel with the dorsal aorta. 
It is probable that the posterior and middle oviducal arteries are 
together equivalent to Osawa's AA. uterinae. 

{c) The anterior oviducal 2Lrtene.s {?A. oviductus — Osawa) (a.od.i) 
frequently arise in conjunction with a dorso-lumbar vessel, but they 
may come off independently from the dorsal aorta, or from the sub- 
clavian arteries, or even from the vertebral artery without any direct 



2IO THE SALAMANDER 

connexion with the dorsal aorta. The anterior oviducal arteries are 
thus subject to a considerable amount of variation, and it is not 
possible to give a general description of them. The illustration (Fig. 
64) shows one arrangement. It will be noticed that in this case the 
artery on the right side has no direct connexion with the dorsal 
aorta at all, but arises anteriorly from the vertebral artery, and pos- 
teriorly from one of the dorso-lumbar vessels which connect this 
artery with the dorsal aorta, while the artery on the left arises directly 
from the aorta posteriorly, and from a dorso-lumbar vessel anteriorly. 
In this case also each artery is in two sections, while in some speci- 
mens they are continuous. In one specimen, a female, they arose 
anteriorly from the subclavian arteries, and posteriorly from the 
dorsal aorta direct. They were continuous, and a connexion with the 
vertebral artery occurred at about the same level as the anterior origin 
of the vessels illustrated. 

It will be noticed that they entwine the post-cardinal veins, and 
their association with these vessels is very close indeed, so that they 
are not easily distinguishable from them. It was doubtless the 
branches of these arteries that Rusconi mistook for factors of the 
post-cardinal veins, which led him to misname them the veins of 
the oviduct, and although Hochstetter was well aware of the true 
homology of the veins he also states that they receive numerous 
factors from the oviduct. Now while two or three factors do enter 
the post-cardinal veins from these organs they can hardly be de- 
scribed as numerous, and it is possible that Hochstetter too may 
have been confusing the arterial branches. 

In the male these arteries are very small, and their branches form 
a fairly large-meshed regular network over the vestigial Miiller's 
duct, giving a very pretty injection. 

Group 3. Vessels distributed to the Limbs and Body-wall, &c. 

(PI. XIV). 

A. A. subclavia (a.scL). The subclavian artery leaves the dorsal 
aorta at the level of vertebrae 3-4, and passes slightly obliquely to- 
wards the axilla. Its first branch is given off about 2 mm. from its 
origin. 

{a) A. thoracica (Osawa) (a.th.). This vessel leaves the posterior 
aspect of the subclavian artery, passes dorsal to the fourth spinal 
nerve, and supplies the muscles of the body-wall immediately pos- 
terior to that nerve. 

At the axilla there arises from the ventral aspect of the subclavian 
artery a short branch which almost immediately divides into the 



THE VASCULAR SYSTEM 211 

A. e-pigastrica anterior and the A. hrachialis. Dorsally there arise 
two smaller branches, the A. subscapularis^ and the A. circumflexa 
scapulae. 

{b) The A. epigastrica anterior (a.e.a.) (A. cutanea magna, Bethge) 
passes ventralwards round the body, deep to the M. pectoralis, but 
lateral to the longitudinal muscles of the ventral body-wall. At the 
lateral edge of the M. rectus abdominalis superficialis it turns abruptly 
caudad along the ventral parietes, between the MM. rect. abd. 
super, and rect. abd. profundus, and deep to the M. obliquus externus. 
It gives off branches segmentally at each muscular inscription, and 
unites finally with the A. epigastrica posterior — a branch of the 
iliac artery — thus forming a complete arterial loop along the ventro- 
lateral aspect of the trunk. Within the arm-pit the A. epigas. antr. 
gives the following two branches: 

(i) A. supracoracoidea (a. sup. cor.) passing, with the nerve of the 
same name, over the dorsal surface of the coracoid, through the 
coracoid foramen, to supply the muscles arising from that bone. 
This branch is always small and is sometimes absent. In the latter 
case the muscles are supplied by a branch from the A. pectoralis 
which passes forwards over the posterior margin of the coracoid on 
its ventral aspect. The supracoracoid artery may apparently some- 
times arise from the A. subclavia direct. 

(ii) A. pectoralis (a.pect.). Leaves the anterior epigastric artery just 
before it turns caudad, and supplies the M. pectoralis and the skin 
covering it. One of its branches — which sometimes arises separately 
from the epigastric — turns anteriorly and enters the muscles cover- 
ing the ventral surface of the coracoid. 

(<:) A. circumflexa scapulae (a.circ.sc.) arises dorsally from the sub- 
clavian artery and passes round the lateral face of the scapula in 
company with the V. circumflexa scapulae. It gives off the following 
six branches: 

(i) A. profunda brachii (Osawa) (a. pf.br.). This artery enters the 
arm dorsally just posterior to the insertion of the M. dorsalis hume- 
ralis, and passes along the dorsal side of the humerus in company 
with the extensor nerves. It supplies the several heads of the M. 
anconaeus and the skin overlying it, and terminates finally in the 
skin within the bend of the elbow — R. cutaneus brachii posterior 
(Osawa). 

(ii) A. subscapularis (Osawa) (a.s.sc.) passes round the mesial side 
of the scapula to the muscle of the same name. 

(iii) and (iv) RR. musculares to the MM. dorsalis humeralis and 
dorsalis scapulae. These vessels enter the muscles mesially. 



212 THE SALAMANDER 

(v) A. trapezia (Osawa) (a.tr.) to the skin covering the anterior 
part of the shoulder and to the M. cucullaris. 

(vi) A. procoraco-humeralis (Osawa) to the muscle of the same 
name and to the skin covering it. 

(d) A. hrachialis (a.br.) is the main artery of the fore-limb and forms 
the direct continuation of the subclavian artery. It enters the arm 
in company with the flexor nerves (N. brachialis). During its 
course through the arm it gives a branch to the M. humeri anti- 
brachialis inferior, and some of the ultimate twigs from this branch 
penetrate the muscle and supply the overlying skin. It varies some- 
what as to its exact point of emergence from the main vessel — it may 
leave it at about the middle of the arm, or it may not emerge until 
near the elbow. In the latter case it runs backwards parallel with the 
A. brachialis before entering the muscle. 

In the bend of the elbow the brachial artery gives off the A. 
radialis (Zuckerkandl), which passes along the radial side of the fore- 
arm and first digit between the NN. cutaneus inferior lateralis and 
dorsalis manus radialis in company with the radial vein. From this 
artery, very close to its origin, arise several twigs to the skin, and a 
recurrent branch. 

(i) AA. cutanei antibrachii^ medialis and /^/,?r^/zV(Gaupp, Frog) with 
several branches supplying the skin on the posterior and flexor sides 
of the elbow. One branch, longer than the rest, extends down the 
flexor side of the fore-arm with the N. cutaneus brachii inferior 
lateralis. 

(ii) A. recurrens radialis^ a small vessel supplying the muscles on 
the flexor side of the elbow. Its name indicates the fact that it turns 
backwards in a proximal direction. 

The direct continuation of the brachial artery is the A. interossea 
(Zuckerkandl) or A. mediana (Osawa), from which arises the ulnar 
artery. 

(i) A. ulnaris (Zuckerkandl, Osawa). This artery and the one 
from which it arises correspond in course and distribution with the 
nerves of the same names. The ulnar artery remains on the flexor 
side of the M. interosseus and divides to supply the MM. flexor 
primordialis communis, flexor antibrachii et carpi ulnaris, and the 
caput longum musculorum contrahentium. Immediately after its 
origin it gives a i^.r^c«rr^;?j uharis which penetrates, and supplies, the 
muscles towards the extensor side of the elbow. A branch from it 
passes forwards superficially along the ulnar side of the limb to join 



THE VASCULAR SYSTEM 213 

a second branch of the ulnar artery — R. recurrens ulnaris anterior — 
which leaves the A. ulnaris at about the middle of the fore-arm. 

After giving off the ulnar artery the A. interossea passes to the 
extensor side of the M. interosseus. Immediately after doing so it 
gives off a branch which passes out at right angles, through the origin 
of the M. extensor antibrachii ulnaris, to the skin of the ulnar side of 
the elbow and fore-arm. It is the A. collateralis ulnaris^ and its twigs 
fuse with those of the R. recurrens ulnaris and also with a terminal 
twig of the A. profunda brachii which probably represents the R, 
collateralis radialis. 

At about the same level the A. interossea gives off a large branch 
which, after giving a small twig to the M. interosseus, turns dorsally 
and comes to lie immediately underneath the M. extensor digitorum 
communis. It is the second main branch of the interosseal artery. 

(ii) The A. interossea externa (Zuckerkandl), or A. perforans 
superior (Osawa), spreads over the dorsum of the fore-arm and carpus, 
and supplies the extensor muscles. Three of its branches are worthy 
of note. First, the R, recurrens posterior passes backwards along 
the fore-arm and fuses with a branch — R. collateralis posterior — 
from the A. profunda brachii. The second passes over the carpus 
and supplies the M. abductor et extensor digiti I, and may finally 
unite with the A. perforans carpi. The third passes out to the skin 
on the ulnar side of the wrist and sends a branch to the ulnar 
side of the fourth digit, where it joins the arcus dorsalis from the A. 
perforans carpi. 

The A. interossea proper passes along the fore-arm, just dorsal to 
the M. interosseus, without further branching. At the level of the 
ulnare + intermedium it divides, and the larger branch penetrates 
a foramen in this bone as the A. -perforans carpi (q.v.), while the 
smaller continues the direction of the main vessel for a short distance 
and then spreads over the palm of the hand supplying the short flexor 
muscles of the digits — A. volaris. 

The A. perforans carpi^ after reaching the dorsal side of the hand, 
runs forwards to the level of the bases of the metacarpals, and then 
divides into two branches. Of these the pre-axial divides again so as 
to supply the adjoining sides of digits i and 2, and 2 and 3 — the 
pre-axial side of the first digit being supplied by the radial artery. 
The post-axial branch of the A. perforans carpi divides to supply 
the adjoining sides of digits 3 and 4, and gives off a twig which 
crosses the base of the fourth digit to anastomose with a branch of 
the A. interossea externa and supply the post-axial side of that 
finger. 



214 THE SALAMANDER 

B. AA. dor so-lumh ales (a.d-lum.). The dorso-lumbar arteries arise 
from the dorsal aorta at frequent intervals and pass directly dorsal- 
wards to fuse with the vertebral arteries. In the anterior region of 
the trunk they are small, but increase in size and importance 
towards the sacral region. The anterior vessels take some share in 
supplying the anterior oviducal arteries (q.v.), while the posterior 
ones may give off small renal arteries to the kidneys, but with these 
exceptions their main function is to supply the muscles and skin of 
the back. Not infrequently a small dorso-lumbar artery may be found 
leaving the proximal portion of the common iliac artery (see also A. 
vertebralis, p. 204). 

C. A. iliaca communis (a. il.com.). The common iliac artery leaves 
the dorsal aorta at the level of the sacral vertebra and passes across 
the dorsum of the pelvis to the leg. Its direction is slightly oblique 
in a posterior direction parallel with the transverse process of the 
vertebra and with the ilium. During its course it gives a small twig 
to the muscles of the back (see above). Just posterior to the hip-joint 
the common iliac artery breaks up into four branches. It is not easy 
to distinguish amongst them the familiar 'external' and 'internal' 
iliac arteries, but it would seem justifiable to regard the branch which 
passes antero-ventrally across the mesial face of the ilium as the 
external iliac^ and the other three as representing together the internal 
iliac. These three branches are, the A. vesicalis^ the A. pudenda^ and 
the A. ischiadica. 

(a) A. iliaca externa gives off its first branch almost immediately 
after separating from the iliac artery. 

(i) The A.glutea (a.gl.) or {J) A. femoralis (Osawa) is a medium- 
sized vessel passing along the thigh antero-dorsal to the A. ischia- 
dica and the sciatic nerve, and postero-dorsal to the femur. It sup- 
plies the MM. extensor ilio-tibialis and ilio-femoralis, and then 
continues down the thigh close to the femur to the knee. Here it 
gives off a R. cutaneus cruralis posterior^ and then bends round the 
knee with the N. peroneus to join the R. recurrens of the A. interos- 
sea externa. It should be noted that the gluteal artery may sometimes 
leave the A. iliaca communis direct, from the angle between this 
vessel and the external iliac. 

In the Frog the A. glutea is a branch of the A. ischiadica and is 
relatively much more limited in distribution, but it should be remem- 
bered that, in that animal, t\\Q femoral ^LYtCYj — i.e. the ^r^-^;czW vessel 
— is the main artery of the leg and the A. ischiadica is confined to 
the thigh, whereas in the Salamander the reverse is the case. The 
position of the A. glutea posterior to the hip-joint and postero-dorsal 



THE VASCULAR SYSTEM 215 

to the femur excludes the possibility of any homology with the 
femoral artery such as Osawa's description seems to suggest. 

(ii) The A. obturatoria (a.ob.) or A. pelvica (Osawa) is a small 
vessel arising from the external iliac at about the same level as the 
gluteal, or occasionally from a common stem with this vessel. It 
passes ventralwards over the dorsal surface of the pubo-ischium and 
supplies the origin of the M. pubo-ischio-femoralis internus, and, 
after passing through the obturator foramen, the origins of the 
muscles arising from the ventral face of the pubo-ischium, e.g. the 
MM. pubo-ischio-femoralis externus, pubo-ischio-tibialis, &c. 

The external iliac then turns rather more ventrally and passes 
round the pelvis in company with the pelvic vein, crossing lateral to 
this vessel in the inguinal region. At the anterior border of the limb 
it gives off(iii) a small branch which enters the thigh between the 
MM. pubo-ischio-femoralis internus and extensor ilio-tibialis, and 
there divides into a branch supplying these muscles — particularly the 
former — and a cutaneous branch passing out to the skin — the A. 
cutanea femoralis lateralis. Although it is only small and compara- 
tively insignificant the position and distribution of this vessel would 
suggest that it is to be regarded as the A. femoralis. The greater 
portion of the region characteristic of this artery is supplied by a 
circumflex and recurrent vessel from the A. ischiadica (q.v.). 

(iv) The remaining portion of the external iliac is the posterior 
epigastric artery (a.e.p.), which, like its anterior counterpart, runs 
along the lateral body-wall at the level of the ventral border of the 
oblique muscles, just outside the peritoneum. It gives off segment- 
ally RR. abdominales to the muscles of the body-wall, and finally 
anastomoses with the posterior extremity of the anterior epigastric 
artery so as to form a complete arterial loop along the side of the 
trunk (cf. p. 211). 

{})) The A. vesicalis (a.ves.) passes ventro-mesialwards to the 
neck of the bladder, and care must be taken on stretching open the 
pelvis — after having split the pubic symphysis — to avoid rupturing 
the artery. On reaching the bladder it breaks up into a tuft of five 
or six small arteries, of which the majority supply the bladder, but 
one or two twigs spread over the ventral side of the rectum, while 
one branch passes back towards the cloaca and supplies the cloacal 
gland. This last branch sometimes fuses with the mesial cloacal artery. 

(c) The A. pudenda (a. pud.) turns postero-ventrally and divides 
into two main branches, although other accessory twigs may occur. 
Of these one is a muscular branch and is distributed to the M. 



2i6 THE SALAMANDER 

ischio-femoralis and the ventral muscles of the tail — MM. caudall- 
pubo-ischio-tibialis and caudali-femoralis. The other branch is the 
lateral cloacal artery and is distributed to the gland, while a few twigs 
go to the skin at the base of the tail. 

(d) The fourth and largest branch of the A. iliaca communis forms 
the direct continuation of that vessel, and is the A. ischiadica^ or 
sciatic artery (a. is.). It enters the limb posterior to the acetabulum, 
and at first it is ventral to the corresponding vein, but within the 
thigh it twists round to the dorsal side. 

(i) Its first branch is given off at about the middle of the thigh, and 
corresponds in position and distribution with the y^.^ro///;z^^/<?;;2(9m 
of the Frog, but it is a much smaller and less important vessel. 
Only a small portion of it is distributed to the M. ischio-fiexorius, 
the major portion passing to the skin on the postero-mesial aspect of 
the thigh as the A. cutanea femoris posterior (Osawa) (A. cut. fem. 
medialis, Gaupp). 

(ii) The most important branch of the A. ischiadica in the thigh is 
the A. articularis genu tibialis (Osawa). This vessel passes to the 
anterior aspect of the thigh round the ventral side of the distal end 
of the femur, and comes to lie between the MM. pubo-ischio- 
tibialis and pubo-ischio-femoralis. Here it breaks up into some four 
or five branches which supply the muscles named and the MM. 
pubo-tibialis and pubo-femoralis. Also there are one or two 
cutaneous branches which doubtless represent the A. cutanea genu 
medialis superior (Gaupp) of the Frog. Close to the origin of the 
A. art. gen. tib. a small twig arises for the M. femoro-fibularis. Two 
other branches are worthy of mention. The first of these is the R. 
ascendens (Osawa), a recurrent branch which passes back up the 
thigh and is distributed to the MM. pubo-ischio-femoralis internus 
and externus. Whether the twigs of this ramus anastomose with 
those of the A. femoralis is uncertain, but it is possible that they may. 
The second branch is the R. descendens (Osawa). This passes over 
the insertion of the M. pub. isch. tib. to the tibial side of the knee, and 
continues down the leg in company with the N. saphenus to the pre- 
axial side of the first digit. It supplies several cutaneous twigs to the 
extensor side of the leg. 

(iii) Another branch arising from the A. ischiadica just proximal 
to the foregoing is the A. articularis genu fibularis (Osawa). This 
vessel leaves the sciatic artery postero-ventrally and supplies the pars 
plantaris of the M. ischio-fiexorius and the insertion of the M. pub. 
isch. tib. as well as the skin on the posterior aspect of the thigh 
and knee. 



THE VASCULAR SYSTEM 217 

The sciatic artery (A. ischiadica) now passes into the leg behind 
the knee and there divides into two terminal branches, viz. the A. 
plantaris (iv), a very short vessel which almost immediately divides 
into lateral and medial rami, and the A. interossea (v) which, like 
the nerve of the same name, passes to the extensor side of the leg. 

The R. plantaris lateralis (Manno) passes to the tibial side of the 
leg, and sends a branch which becomes superficial at the lateral 
border of the insertion of the pars plantaris of the M. ischio-flexorius, 
and then gives off a R. descendens and a R. ascendens. The R. 
descendens passes down the leg to the sole of the foot, while the R. 
ascendens turns backwards to the lateral aspect of the knee and is 
distributed to xhe pars plantaris of the M. ischio-flexorius and to the 
insertion of the M. pub. isch. tib. The remainder of the R. plantaris 
lateralis continues down the leg on the mesial side of the tibia to the 
sole of the foot, and supplies the short flexor muscles of the tibial side. 
During its passage through the leg it gives a twig to the M. pronator 
profundus and to the M. interosseus. 

The R. plantaris medialis (Manno) passes out to the fibular side of 
the leg, deep to the M. flexor primordialis communis, and immedi- 
ately superficial to the M. caput longum musculorum contrahentium, 
in company with the N. fibularis. As it turns distalwards down the 
leg it gives a fairly large cutaneous branch to the flexor surface — A. 
cutanea cruralis medialis (Gaupp). It also supplies the muscles named 
above and then comes to lie — together with the accompanying nerve 
— close against the surface of the fibula. At the tarsus a cutaneous 
branch is given off to the skin on the fibular side of the ankle and a 
deep muscular branch to the MM. flexores accessorii, medialis and 
lateralis. It then continues along the fibular border of the tarsus to 
the base of the fifth metatarsal, where it turns mesially to supply the 
short flexor muscles of the fibular side of the foot. 

(v) A. interossea (Zuckerkandl). 

A. perforans cruris ....... (Manno). 

A. tibialis medialis ....... (Osawa). 

Immediately after reaching the extensor side of the M. interosseus 
the interosseal artery gives off a superficial branch. 

(yd) A. interossea externa (O'Donoghue, in Sphenodon). 

R. dorsalis cruris ...... (Zuckerkandl). 

A. perforans superior ....... (Osawa). 

This artery supplies the skin on the dorsum of the leg — A. cutanea 
cruralis lateralis — and the extensor muscles. It lies on the tibial side of 
the A. interossea and is separated from it by the corresponding vein 



2i8 THE SALAMANDER 

and nerve. One noteworthy branch Is the R. recurrens which runs 
back around the proximal end of the fibula close to the N. peroneus 
and anastomoses with the distal extremity of the gluteal artery. ^ 

The A. interossea proper passes right down the leg without 
branching, and, at the distal border of the M. interosseus, passes to 
the ventral side of the tarsus, and then, as the A. perforans tarsi 
(Manno) (A. perforans inferior, Osawa), penetrates to the dorsal 
side of the foot through the foramen between the tibiale and inter- 
medium. Here it is known as the A. dorsalis pedis and divides into 
four branches, each of which divides again to form the AA. digitales 
supplying the adjoining sides of the toes. 

6. Branches of the Pulmonary Arch (PI. XXIII, fig. 77). 

The ventral oesophageal (or pharyngeal) artery (a.oes.v.) arises 
mesially from the pulmonary arch at the posterior border of, and 
ventral to, the dilatator laryngis muscle. It passes anteriorly, dorsal 
to the M. cephalo-dorso-subpharyngeus and ramifies over the whole 
ventral surface of the pharynx and the floor of the mouth as far as 
the root of the tongue. Its most posterior branch is the laryngeal 
artery supplying the muscles of the larynx. 

The dorsal oesophageal {or pharyngeal) artery (a.oes.d.) arises later- 
ally from the pulmonary immediately opposite the ventral vessel, and 
bends dorsally round the oesophagus to meet a similar vessel from 
the opposite side, with which it fuses, and then continues as a median 
artery for some distance down the oesophagus posteriorly. These 
arteries thus form a Y-shaped vessel from which branches ramify- 
over the whole of the dorsal surface of the oesophagus and pharynx. 

The pulmonary artery (a.pul.), as the continuation of the arch may 
now be called, passes posteriorly, rather closely embedded in connec- 
tive tissue, and enters the lung on its dorso-lateral border about a 
quarter of the way down. It commences to divide just before its entry 
into the lung. 

Unless the small cutaneous branches of the ductus Botalli may be 
taken to correspond with the cutaneous artery of the Frog, this vessel 
is not represented in the Salamander (cf. pp. 202—3). 

7. The Venous System. 

The Venous Trunks (PI. XV). 

To be logical the course of the veins should be traced from their 
distal origins towards the heart, but, since it is impossible to see the 

'^ The A. interossea externa may separate from the A. ischiadica before the AA. 
interossea and plantaris, the actual order of branching being somewhat indefinite. 



THE VASCULAR SYSTEM 219 

smaller factors without the aid of a good dissecting microscope such 
as is not usually available to the ordinary student, the detailed de- 
scription of the venous system is prefaced by the following short 
account of the sinus venosus and the main venous trunks entering it. 

The sinus venosus forms a triangular swelling on the anterior end 
of the post-caval vein, almost directly after the latter emerges from 
the substance of the liver, on the left side of the heart. The base of 
the triangle is anterior. It may be seen from the ventral side, after 
opening the pericardium, by gently pressing back the left auricle. 
The ducts of Cuvier enter the basal angles, while the post-caval vein 
enters the apex. 

The left ductus Cuvieri is either very short or entirely wanting, since 
the union of the post-cardinals with the anterior veins — the internal 
and external jugulars — occurs very close indeed to the sinus. 

The right ductus Cuvieri^ on the other hand, is quite long, and ex- 
tends across the dorsal side of the heart — at about the level of the 
anterior end of the ventricle — since the union of the venous trunks 
on this side is symmetrical with that on the left, and occurs outside 
the pericardium. Thus a long ductus Cuvieri is necessary to reach 
the sinus venosus, which is situated well to the left of the middle line. 
It may easily be seen by turning the tip of the ventricle forwards. 

The venous trunks uniting to form the ducts of Cuvier on either 
side are four in number, viz. the external jugular, the internal jugular, 
the subclavian, and the post-cardinal. The exact order in which these 
veins join up varies a little, but they may be easily identified as 
follows. The external jugular (v.j.e.) is the most anterior, and com- 
prises (i) the common facial vein (v.f.c.) arriving laterally from the 
paratoid region, and (ii) the lingual and thyroid veins arriving from 
a mesial direction, the thyroid vein lying ventral ^.nd the lingual vein 
dorsal to the arterial arches. They unite to form a common vessel 
immediately posterior to the arches, and the vessel thus formed 
usually unites with the common facial vein to form a short trunk — 
the external jugular proper — which, in turn, joins the subclavian 
vein. It may, however, occasionally enter the subclavian separately 
from the facial vein. 

The subclavian vein (v.scl.) is the largest and most prominent of 
the vessels entering the ductus, and also, with the exception of the 
thyroid vein, the most ventral, while its direction is almost exactly 
transverse, so that it is easily found and forms a useful landmark for 
identifying the other vessels. 

The internal jugular (v.j.i.) arrives from an antero-dorsal direction 
and joins the subclavian vein close to the external jugular. It is most 



2 20 THE SALAMANDER 

easily found from the posterior aspect of the subclavian vein, between 

this vessel and the post-cardinal vein. 

T}\Q post-cardinal vein (v.card.p.) lies very close to the oesophagus, 
round which it passes, and is the most posterior of the veins entering 
the ductus Cuvieri. It is only likely to be confused with the internal 
jugular, but it is more posterior than this vessel and more mesial, 
while its direction is slightly po5tero-dors?i\ as compared with the 
obliquely antero-dorsal direction of the internal jugular. 

A rather tough connective tissue enclosing lymph spaces sur- 
rounds the whole of the venous complex just described, and, as the 
walls of the vessels are very thin, considerable care must be exercised 
in cleaning them to avoid rupture. 

The ducts of Cuvier are often referred to as the pre-caval veins, 
but in view of the persistence of the post-cardinal veins in a functional 
condition, and the retention at the same time of their primitive con- 
nexions, it would seem preferable to call the vessel formed by the 
union of the post-cardinal vein with the derivative of the anterior 
cardinal vein (the internal jugular) the ductus Cuvieri. 

8. Description of the Veins. 

For descriptive purposes the venous system may be divided into 
groups according to distribution and function as follows:^ 

Veins entering the Heart direct. 

Veins of the Head. 

A. Internal Jugular (see footnote). 

B. External Jugular. 

Veins of the Trunk and Viscera. 

A. Post-caval. 

B. Post-cardinals. 

C. Subclavian. 
Veins of the Lungs. 

Veins of the Portal Systems. 
Hepatic Portal System. 

A. Abdominal Vein. 

B. Portal Vein. 
Renal Portal System. 

A. Caudal Vein. 

B. Common Iliac Veins. 

C. Pelvic Veins. 

'^ The veins of the cranial cavity, factors of the internal jugular, are described at the 
end of the present section under 'Vessels of the Brain'. 



THE VASCULAR SYSTEM 221 

Veins entering the Heart direct. 

Veins of the Head (PI. XV and PI. XIX, fig. 70). 

A. F. jugularis interna (internal jugular vein) (v.j.i.). This 
important vein may be said to be formed by the fusion of two 
main factors. One of these lies latero-dorsal to the aortic arches and 
returns the blood from the occipital region and the muscles of the 
*neck', and may be called the V. jugularis lateralis (v.j.L). The other 
lies ventro-mesial to the internal carotid and the dorsal part of the 
systemic arch and drains the palate. This factor is the F. pharyngo- 
palatina (v.ph-p.) (Druner). There are anastomoses between these 
two vessels that make it difficult to decide exactly which factors 
belong to each, and particularly is this so in the case of the V. petrosa 
lateralis, as it discharges into both vessels. It has been included 
among the factors of the V. pharyngo-palatina, as the contribution 
to this vessel is larger, and it also anastomoses with some of its 
palatine factors. 

Owing to the large amount of individual variation met with, par- 
ticularly in the palatine region, it is very difficult to give a detailed 
description of the factors of the veins of the head. Only the more 
constant features have been noted therefore, together with some 
indication of the types of variation to be expected. 

Factors of the F. jugularis lateralis. 

(a) RR. musculares from the MM. cucullaris and opercularis. 
These enter the vein variously as it passes along the ventral edge of 
the muscles. 

{jy) F. occipitalis (occipital vein). This vein arises just behind the 
head, between the MM. inter-transversarius capitis superior and 
inferior, by factors from these muscles and from the M. opercularis. 
It also receives : 

(i) A small F.post-occipitalis from the cranial cavity, which passes 
through a foramen in the first vertebra, as in the Frog. 

(ii) A F. cranialis post-otica (sinus jugularis. Rex) from the region 
of the medulla. This factor emerges from the cranial cavity together 
with the ninth and tenth cranial nerves. 

(iii) A R. muscularis from the M. cucullaris, passing dorsal to the 
X+X ganglion. 

(iv) An anastomosing factor from the vertebral vein. 

The occipital vein joins the main vessel ventral to the IX+X 
ganglion. 

(c) F. vertehralis (v.v.). The vertebral vein is one of the most 



222 THE SALAMANDER 

important factors of the V. jugularis lateralis. It collects the blood 
from muscles of the neck region on the dorsal side of the vertebral 
column. It passes anteriorly dorsal to the M. opercularis and the Xth 
nerve, and mesial to the I Xth nerve to join the main vessel posterior 
to the IX+X ganglion. 

Very frequently, but perhaps not quite constantly, the vertebral 
veins may be traced right through the dorsal muscles to the ventral 
side, appearing just anterior to the transverse process of the second 
vertebra. They then pass backwards alongside the vertebral column 
to join the post-cardinals at the point where they turn lateralwards 
to loop around the oesophagus. An anastomosis from the palatal 
region is also frequently received at the point where they emerge on 
the ventral surface of the muscles. The connexion with the post- 
cardinal veins is primitive, and in the Frog, where the post-cardinals 
are absent, the vertebral veins discharge entirely into the internal 
jugular veins. 

(d) Another considerable tributary of the V. jugularis lateralis is 
an anastomosis with the V. pharyngo-palatina. It arises somewhat 
variously, either from the V. pharyngo-palatina itself, or from its 
mandibular factor, or even from the V. petrosa lateralis. It enters 
the V. jugularis lateralis ventrally just posterior to the columella. 

(e) The only other tributary of note is the dorsal branch of the V. 
■petrosa lateralis. This vessel leaves the V. petrosa lateralis within the 
antrum, and emerges between the columella and the processus oticus, 
looping round the ^orWside of the columella to enter the V. jugularis 
lateralis dorsally. 

Factors of the V. pharyngo-palatina . 

(a) V. petrosa lateralis (Driiner) (v.pet.l.). 

V. capitis lateralis ....... Goodrich. 

V. lateralis ........ Osawa. 

This is the vessel which passes through the space between the palato- 
quadrate and the otic capsule — the antrum petrosum laterale. It 
drains the cranial cavity and the orbit by several factors as follows : 
The peri-orbital network (p.o.n.) (see also PI. XIII, fig. 58) is a 
wide venous girdle completely surrounding the bulbus oculi. It may 
be said to originate at the anterior border of the orbit where it re- 
ceives the V. nasalis (v.na.) from within the nasal capsule, and a 
medium-sized cutaneous vein from the skin overlying it — F. fron- 
talis (v.fr.). The network lies mainly, but not entirely, deep to the 
eye-muscles, and drains them as well as the tissues of the orbit 
generally. It is hardly possible to differentiate any particular vessel, 



THE VASCULAR SYSTEM 223 

but one rather large vein, which may be termed the F. orbito-nasalis 
(v.o.na.) leaves the anterior end of the plexus and passes postero- 
mesially deep into the orbit to a point close to the optic fenestra, 
where it receives the V. ophthalmica (v.oph.) from the bulbus oculi. 
The ophthalmic vein passes inwards from the eye along the optic 
peduncle. The V. orbito-nasalis then turns dorsalwards and dis- 
charges into the posterior end of the peri-orbital network and more 
particularly into the V. orhitalis superior^ (a factor of the V. petrosa 
lateralis), and into the superior facial vein (a factor of the external 
jugular). 

The V. orhitalis superior (v.o.s.) lies deep to, and drains, the M. 
levator mandibulae anterior and the skin overlying it. It also drains 
the deeper parts of the dorsal portion of the peri-orbital network as 
already described. 

The V. orhitalis inferior (v.o.i.) is the ventral counterpart of the 
above and drains the deeper regions of the peri-orbital network ven- 
trally as well as a considerable portion of the levator bulbi muscle. 

Another important tributary of the V. petrosa lateralis is the V. 
anastomotica (v.anas.) joining it to the palatal vessels. Sometimes it 
joins the V. palatina itself, more frequently the maxillo-palatine vein, 
and sometimes both vessels. 

The V. cranialis pro-otica (Gaupp, in Frog) (v.cr.pr-ot.) or V. cere- 
bralis anterior (van Gelderen) is the chief vein draining the cranial 
cavity. It leaves the skull together with N. ophthalmicus profundus 
V and joins the V. petrosa lateralis mesially just as it enters the 
antrum. 

Within the antrum a factor is received which enters the cavity 
between the Proc. oticus and the Proc. ascendens; the V. petrosa 
lateralis itself enters the antrum ventral to the Proc. ascendens, 
between it and the Proc. basilaris. It drains the M. levator mandi- 
bulae. Another small vein leaves the cranial cavity in company with 
the Vllth nerve and joins the V. petrosa lateralis near the posterior 
extremity of the antrum. It may be called the F. cranialis mesotica 
(v.cr.m-ot.) and obviously corresponds with the V. cerebralis medialis 
of van Gelderen. At about the same level a small factor draining the 
loose connective tissue and skin immediately posterior to the ear can 
usually be found. 

Van Gelderen has made a comparative study of the origin of the 
veins draining the cranium in vertebrates, and it is apparent, from 
his account, that the condition in Salamandra must be looked on as 
primitive. 

(J?) Tht factors from the palate entering the F. pharyngo-palatina are 



224 THE SALAMANDER 

extraordinarily variable. The whole region is highly vascular, and 
anastomoses are frequent, so that the several main vessels may vary 
considerably in size and extent from specimen to specimen or even 
on the two sides of the same individual. 

The most mesial of them is the V. palatina (v.pal.). It arises 
by many fine tributaries from the region of the internal nares and 
from the mucous membrane covering the pre-vomer, and, follow- 
ing the curve of the posterior tooth-bearing process of this bone, 
drains the rich vascular network surrounding the bases of the 
teeth. Just in front of the dorsal portion of the systemic arch the 
palatine vein drains a venous plexus forming a coarse network on the 
roof of the pharynx in the middle region between the occiput and 
the posterior ends of the pre-vomerine teeth (pi. ph.). It then turns 
sharply laterally, and, following the curve of the systemic arch, joins 
the pharyngo-palatine vein at the side of the pharynx a short distance 
before this vein itself unites with the V. jugularis lateralis to form 
the internal jugular. 

The vein just described lies mesial to the nerve and to the palato- 
nasal artery, but there is also a vein lateral to these structures which 
may accordingly be called the lateral palatine vein (v.pal. 1.). It re- 
ceives its blood from the mesial part of the floor of the orbit — from 
the M. levator bulbi, and the mucous membrane of the roof of the 
mouth — and may be distinguished from the V. orbitalis inferior in 
that it lies ventral to the M. levator bulbi. It nearly always joins the 
V. anastomotica of the V. petrosa lateralis and not infrequently the 
V. palatina also, but either of these connexions may be absent. 

The V. maxillo-palatina (v. max. pal.) drains the anterior part of the 
upper jaw, and, at about the middle of the orbit, turns mesialwards 
and crosses obliquely towards the postero-mesial corner of the orbit. 
Here it may join the V. anastomotica and so empty itself into the V. 
petrosa lateralis, or it may pass somewhat more lateralwards and dis- 
charge into the V. pharyngo-palatina direct, while both connexions 
may be present. When it enters the V. pharyngo-palatina direct it 
does so close to the point where the V. petrosa lateralis and the V. 
mandibularis profunda are received. 

The V. mandibularis prof unda (v.m.pf.) arises from an anastomosis 
with the internal mandibular vein (a factor of the external jugular, 
q.v.) just posterior to the N. intermandibularis. It passes backwards, 
dorsal to the MM. interhyoideus and interhyoideus posterior, receiv- 
ing blood from the former muscle, as well as numerous tributaries 
from the floor of the mouth between the lower jaw and the base of 
the tongue. Sometimes two vessels may enter the V. pharyngo-pala- 



THE VASCULAR SYSTEM 225 

tina from the floor of the mouth, one from the mandibular region, 
and the other from the tissues on the dorsal side of the cerato-hyal, 
but this would appear to be merely a variety of the type where both 
regions drain into one vessel. 

Close to the point where the above vessel is received a small F. 
pterygoidea (v.pt.) also enters. It arises in the posterior region of 
the upper jaw and passes backwards along the ventral side of the 
pterygoid. 

B. The external jugular may be regarded as comprising three main 
venous factors: 

(I) The F. facialis communis (v.f.c), Vena jugularis externa 
(Driiner), or Vena jugularis interna.'' (Bethge). This vessel collects 
from three main regions, the orbit, the superficial intermandibular 
region, and the thymus. It arises in the orbit. 

The superior 2.nd inferior palpebral veins (v.palp.s. and v.palp.i.) 
collect blood from the upper and lower eyelids respectively. These 
vessels unite at the posterior angle of the orbit where they are joined 
by several small factors from the glandular tissue of that region to 
form the superior facial vein. 

(i) The F. facialis superior (v.f.s.) passes backwards superficially to 
the level of the thymus gland. It is closely applied to the skin, from 
which it receives numerous branches, particularly from the large 
paratoid gland of that region. In the region of the thymus it joins 
its ventral counterpart — the inferior facial vein. 

(ii) The F. facialis inferior (v.f.i.) appears to arise at the tip of the 
lower jaw from the genio-glossus muscle, and from an anastomosis 
with the corresponding vessel of the opposite side, as the internal 
mandibular vein. The F. mandihularis interna (v.m.i.) passes through 
the loose connective tissue on the ventral side of the mandible. 
Numerous factors are received from the intermandibularis muscle 
and from the skin of the intermandibular region. At the posterior 
border of the muscle an anastomosis is received from the V. mandihu- 
laris profunda — a factor of the Internal Jugular — and from now on 
the vessel is spoken of as the inferior facial vQ\n (v.f.i.). Continuing 
along the ventral surface of the mandible it passes dorsal to the M. 
interhyoideus and ventral to the M. interhyoideus posterior, receiv- 
ing factors from both muscles. At the posterior angle of the lower 
jaw it turns abruptly dorsalwards to join the superior facial vein at 
the anterior border of the thymus as above described. 

Numerous factors from the thymus gland enter the combined 
superior and inferior facial veins to form the common facial vein, 
which then proceeds ventro-mesially to enter the ductus Cuvieri as 



226 THE SALAMANDER 

already described. A small laryngeal v€\n can usually be found enter- 
ing the vessel just before it merges into the ductus. 

(II) The V. lingualis (v.ling.) or Vena pharyngea? (Bethge). This 
is essentially the vein of the tongue, emerging from that organ 
between the anterior and posterior radials of the copula, and running 
posteriorly along the mesial border of the first hypobranchial carti- 
lage. At the point where the second hypobranchial cartilage unites 
with the first branchial arch the vein loops back on itself a little so 
as to allow it to stretch when the hyobranchial apparatus is pro- 
tracted, while at this point also it receives a factor from the M. 
subarcualis rectus I. The vein then leaves the cartilage and turns 
somewhat mesially, passing dorsal to the arterial arches very close to 
their origin from the truncus arteriosus. As the vessel passes over the 
arteries it divides into two or more smaller veins which reunite to 
form a single vessel on the posterior side of the pulmonary arch, thus 
forming a closed venous loop. The reason for this is fairly obvious. 
The space between the arterial arches and the pharynx is very re- 
stricted, so that the vein must be subjected to considerable pressure 
both from the blood in the arteries and from food passing down the 
pharynx, and it is manifest that the blood would flow with more free- 
dom under these circumstances if it were contained in several small 
vessels instead of in one large one. At this point also several large 
vessels from the ventral pharyngeal wall enter the vein, but the details 
are particularly difficult to determine owing to the fact that the blood 
or injection mass gets squeezed out of the veins by pressure from 
the arteries. Emerging on the posterior side of the arterial arches 
the vein turns sharply laterally and is soon joined by the thyroid 
vein, and sometimes also by the common facial before entering the 
ductus Cuvieri. 

Just at the point where the lingual vein turns laterally to meet the 
thyroid vein it receives the efferent trunk from the central lymph 
heart, constituting one of the most important connexions between 
the lymphatic and venous systems. It is obviously this trunk which 
Driiner refers to as 'Vena reuniens' (see also p. 260). 

(Ill) The V. thyroidea (v.th.) or V. lingualis (Bethge). The thy- 
roid vein arises in the loose connective tissue between the hyobran- 
chial muscles just anterior to the thyroid gland as the V. thyroidea 
advehens (Druner) and passes posteriorly, ventral to the gland, re- 
ceiving on its way numerous factors from the rich vascular network 
of the gland itself. At the posterior margin of the thyroid gland a 
cutaneous factor is received. This small vein arises in the skin in the 
neighbourhood of the gular fold and passes to the region of the os 



THE VASCULAR SYSTEM 227 

triangulare in approximately the middle line. It then turns lateral- 
wards, dorsal to the M. genio-hyoideus and ventral to the M. rectus 
cervicis to join the thyroid vein. It is very variable in size and 
extent. 

After receiving the above-mentioned vessel the thyroid vein con- 
tinues posteriorly as the V. revehens (Druner), ventral to the aortic 
arches, to join the lingual vein. It occasionally unites with the facial 
vein just before this vessel joins the lingual. 

It is not difficult to see how the system of the external jugular has 
been derived from the sinuses of the fish.^ Thus the superior and 
inferior palpebral veins draining the peri-orbital network, and the 
superior and inferior facial veins appear to be derived, in part at least, 
from the orbital, post-orbital, and hyoidean sinuses of the Selachian, 
while the lingual and thyroid veins represent the inferior jugular 
sinus. 

Veins of the Trunk and Viscera (Pis. XV and XXII). 

A. The post-caval (v.c.p.) vein is a single, unpaired, more or less 
median vessel which first originates among the Dipnoi, and takes over 
the function of the post-cardinals by returning the blood from the 
posterior region of the body to the heart. It is general among the 
Amphibia and all higher groups. In the Salamander the transference 
of function is not complete, since the post-cardinal veins persist in 
the adult and are quite functional. In a few cases the post-caval may 
be wanting, with a consequent enlargement of one or other of the 
post-cardinal veins. These cases throw considerable light on the 
probable mode of origin of the post-caval, but the question will be 
reconsidered after giving a description of the vein as it is normally 
found in the adult Salamander. 

The post-caval vein arises near the posterior end of the kidneys 
by the fusion in the middle line of a fairly large renal vein from either 
kidney. It continues anteriorly, ventral to the dorsal aorta, to the 
level of the coeliaco-mesenteric artery, receiving during its course 
numerous renal factors from the kidneys and two or three genital 
(spermatic or ovarian) factors from the left gonad and fat-body. The 
vein now divides: one half passes dorsally and almost immediately 
divides again as the post-cardinal veins, while the other, turning some- 
what to the right, passes antero-ventrally into the liver as the true post- 
caval. The genital factors from the right gonad enter at about this 
point, either into the post-caval, or into the stem which is combined 
as the post-cardinals. The post-caval enters what appears to be 

I Cf. 0'Donoghue(i9l4). ^.... 



228 THE SALAMANDER 

a dorsal out-growth of the right lobe of the liver, the 'post-cavaF 
lobe (Hohlvene Fortsatz, Hochstetter), which grows right down 
to the point where the post-cardinals are constituted and invests the 
vein very closely. 

The post-caval becomes visible again on the ventral surface of the 
liver, just to the right of the gall-bladder, and it may be traced as a 
dark line passing obliquely across the liver to emerge free once more 
at the left edge of the pericardium, and, after a free course of about 
6 mm., it enters the sinus venosus as already described. At the 
junction between the post-caval vein and the sinus venosus a pair of 
small veins enter, either together or separately. These have been 
called by Hyrtl the 'Venae abdominales anteriores', and Joseph for 
the sake of convenience adopts the same name. They are, however, 
the mesial branches of the anterior epigastric veins. They arise on 
either side by the bifurcation of the anterior epigastric veins into 
mesial and lateral factors (see under factors of subclavian vein, p. 2 30) 
at about the level of the posterior border of the coracoid. They pass 
mesially, dorsal to the rectus abdominis muscle, to unite and enter 
the sinus as described, or they may enter separately. The right 
anterior epigastric vein is necessarily longer than the left, and re- 
ceives a factor in the middle line dorsal to the sternal plate. This 
small vein arises in the skin ventral to the sternum and passes either 
posterior to, or through the cartilage to reach the dorsal side. It would 
therefore seem best to call it simply the sternal vein. 

If now the liver tissue be carefully teased away from that portion 
of the post-caval which passes through it, which is best done with 
blunt needles or seekers, numerous hepatic veins will be found enter- 
ing it from the liver. They enter from both sides, but, on the left, 
there is usually one larger than the rest which is referred to as the 
hepatic vein (v.h.s.). Very occasionally this larger vein may enter on 
the right side. 

We may now consider the cases where the post-caval vein is 
apparently wanting. There have been two such recorded, one by 
Hochstetter (1888), and the other by Joseph ( 1 902), while Romeiser 
(1905) summarizes these two cases and describes a similar occur- 
rence in Necturus. Both the cases mentioned are essentially similar 
except that in one it is the right, and in the other the left, post- 
cardinal that hypertrophies to carry the additional blood. An ex- 
amination of the liver in these cases revealed a pair of exceptionally 
large right and left veins, more or less symmetrical, which fused near 
the anterior end of the liver, and then emerged from it as a single 
vessel to enter the sinus venosus in a precisely similar manner to the 



THE VASCULAR SYSTEM 229 

normal post-caval. It would therefore appear that the post-caval vein 
is normally formed by a dorsal extension of the right hepatic vein 
anastomosing with the fused inter-renal portion of the post-cardinals. 
The dorsal extension of the right hepatic vein has been named by 
Hochstetter, who was the first to record the abnormality referred to, 
the 'Mittelstiick' of the post-caval. The post-caval vein is to be 
regarded therefore rather as a modification of existing vessels than 
as a new structure. 

B. The post-cardinal veins (v.card.p.). 

Venae vertebrales ....... Funk. 

Veines de I'oviducte ....... Rusconi. 

Venae azygoi ....... Hochstetter. 

In view of the above considerations the post-cardinal system should 
logically be regarded as arising from the posterior end of the kidneys 
as a median vessel, but since this section has already been included 
under the post-caval vein it will be described from the point where 
the post-cardinal veins separate from the latter vessel. 

As already mentioned, the post-cardinal system separates from the 
post-caval as a single vessel which almost immediately divides into 
two, and these pass anteriorly on either side of the dorsal aorta. One 
case has been found in which the right post-cardinal passed dorsal 
to the aorta to reach the right side, but this is abnormal, the veins 
usually remaining ventral to the artery. This division takes place at 
the anterior end of the kidneys, and occasionally a remnant of the 
primitive connexion between the post-cardinals and Jacobson's vein 
may persist. 

During their course alongside the aorta the post-cardinal veins re- 
ceive tributaries from the muscles of the back. These join the main 
veins at about the centre of each vertebra and appear at first sight 
to enter alternately from the right and left sides, but on closer inspec- 
tion each tributary is found to collect from both sides. The tributary 
from the side opposite to that of the main factor passes ventrally 
between the muscle and the vertebra, and then crosses over the middle 
of the vertebra and dips dorsalwards again between the vertebra and 
the muscle on the other side to join the main factor within the sub- 
stance of the muscle. A little dissection is therefore necessary to dis- 
play the relations. Besides these dorsal factors the post-cardinal veins 
also receive some three or four anterior oviducal veins from the an- 
terior portion of the oviduct connecting them with the longitudinal 
oviducal vtin (v.od.long.). This vessel runs along the ventral aspect 
of the anterior three-quarters of the oviduct (i.e. along the portion 



230 THE SALAMANDER 

anterior to that in which the embryos develop), draining the whole 
of the blood therefrom and delivering it, by means of the anterior 
oviducal veins above mentioned, into the post-cardinal veins. The 
anterior oviducal veins are somewhat variable in number and position 
and should be distinguished from the anterior oviducal arteries (see 
p. 209). 

Hochstetter (1888) figures the tight post-cardinal passing dorsal, 
and the left ventral to the corresponding subclavian artery. While 
this arrangement is not infrequent it would appear normal for both 
veins to pass dorsal to the artery. 

Just before the veins cross the arteries they receive an anastomosis 
from the post-axial vein of the arm. This vessel, usually quite small, 
runs parallel with, and posterior to, the subclavian artery. It would 
appear to be an enlarged parietal vessel (see also p. 234). 

Immediately anterior to the subclavian arteries the post-cardinal 
veins turn off laterally at right angles to loop around the oesophagus. 
At this point a number of fine vessels enter, including one of variable 
size which continues the direction of the post-cardinals along the 
remainder of the vertebral column as far as the occiput, where it 
usually anastomoses with the vessels of the pharyngeal plexus. It is 
also in communication dorsalwards with a factor of the internal jugular 
within the substance of the muscles (see p. 222). Osawa (1902) 
calls this vessel the anterior cardinal vein, but since the internal 
jugular must be regarded as the derivative of that sinus, the anterior 
prolongation of the post-cardinal vein is here called the vertebral vein. 

The origin and development of the post-cardinal veins, their rela- 
tion to the kidneys and to Jacobson's veins, were first described by 
Hochstetter (i 8 8 8) for Salamandra atra. The substance of his work 
is quite well known and there is therefore no need to repeat it here. 

C. V. suhclavia (v.scl.) or V. cutanea magna (partim) (Bethge). 
In addition to the blood from the fore-limb the subclavian vein also 
receives two important factors from the muscles and skin of the 
trunk, viz. the V. lateralis and the V. epigastrica anterior. The 
description of the subclavian vein thus falls naturally into three 
heads. 

{a) V. epigastrica anterior (v.e.a.). The anterior epigastric veins lie 
in the ventral body-wall alongside the arteries of the same name, at 
the lateral edges of the MM. recti abdominales superficiales. Each 
arises a little posterior to the caudal edges of the liver by factors from 
the neighbouring muscles, as well as by transverse tributaries from 
the median epigastric vein (see p. 237), and passes forwards to the 
posterior border of the coracoid, where it divides into two branches. 



THE VASCULAR SYSTEM 231 

The mesial branch of the lejt side continues directly forwards, in- 
cHned sHghtly mesiad, and enters the junction of the post-caval and 
sinus venosus. The mesial branch of the right side also passes for- 
wards to the pericardial margin and then turns abruptly to the left 
around the apex of the pericardium and joins its fellow on the left side 
just as it enters the post-caval. The lateral branches of the anterior 
epigastric veins are symmetrical, so that the description of one will 
serve for both. It accompanies the artery and turns sharply dorsal- 
wards to the region of the axilla, where it joins the V. lateralis and the 
V. brachialis medialis. On its way it receives two important factors : 

(i) V . pectoralis (v.pect.) from the muscle of the same name, as well 
as from the M. coraco-brachialis brevis. The factor from the latter 
can be traced right down the muscle into the arm, where it appears 
to form a direct continuation of the V. anastomotica brachii (see 
also p. 235). 

(ii) One or two VV. cutaneae from the skin of the axilla. 

The exact pattern of the junction of the vessel with the V. lateralis 
and V. brachialis medialis is somewhat variable, but usually the V. 
lateralis and anterior epigastric vein first join to form a short common 
trunk, a V. axillaris (v.ax.), and then receive the vein from the arm. 

The vessel just described is called the epigastric vein in Crypto- 
branchus (Osawa) and Rana (Gaupp).i In the Salamander it is norm- 
ally larger than the mesial branch of the anterior epigastric vein and 
therefore carries most of the blood from that vessel. Nevertheless 
it seems quite probable that it actually represents a 'V. coraco-pec- 
toralis' which should be regarded as a factor of the V. axillaris that 
has become secondarily associated with the V. epigastrica anterior 
by fusion with a factor of the latter vein, the association becoming 
increasingly stronger during phylogeny until it finally provides the 
only path of egress for the blood from the epigastric vein. Such an 
hypothesis is rendered the more probable in that it allows a simple 
explanation of the direct connexion between the anterior epigastric 
veins and the sinus venosus in Salamander, which would otherwise be 
somewhat puzzling. 

As is well known from the work of Goette, Hochstetter, and 
others, the abdominal vein in Amphibia develops by the fusion of 
a pair of lateral vessels connected anteriorly with the sinus venosus 
or ductus Cuvieri and posteriorly with the iliac veins. In the Frog 
the former connexion normally disappears while the latter develops 

' In these animals there is no connexion between the anterior epigastric vein and the 
sinus venosus, although it is possible that one may exist in Cryptobranchus but has been 
overlooked by Osawa. It is very easily destroyed in opening the animal. 



232 THE SALAMANDER 

as the pelvic veins. The Dipnoi also possess an abdominal vein, but 
in these animals the anterior connexion is also retained. Now in 
Salamandra the anterior epigastric veins are connected by a short 
common stem with the sinus venosus post-caval junction, and pos- 
teriorly with the abdominal vein, by means of the median epigastric. 
It seems therefore not unreasonable to suppose that they represent 
the primitive vessels connecting the abdominal vein with the sinus 
venosus, a connexion which is in the process of being replaced by the 
lateral connexions with the VV. axillares, while their anterior junction 
with the heart has become secondarily associated with the posterior 
end of the sinus venosus rather than with the ductus Cuvieri. 

(b) V, lateralis (v.l.) or V. cutanea magna (partim) (Bethge). The 
two synonyms for this vessel are fairly uniformly distributed through- 
out the literature, but the former has been chosen as it is not by any 
means a purely cutaneous vein, but receives numerous factors from 
the muscles of the body-wall also. It was first described by Bethge 
in 1898. 

The lateral vein arises at the side of the tail, a little posterior to the 
cloaca, immediately beneath the skin, and passes right along the body 
in the sulcus lateralis between the dorsal and ventral body-muscles to 
a point just anterior to the third myoseptum. On its way it drains both 
the skin and the muscles of the lateral body-wall. An important anas- 
tomosis with the V. iliaca transversa occurs in the pelvic region (see 
p. 243), and sometimes one with the V. pelvica also. It is in com- 
munication with the lymphatic system by means of the lateral lymph 
hearts which occur at each myoseptum (see also pp. 253, 259). The 
vessel has a tendency to sink into the sulcus lateralis and to become 
partly hidden by the muscles. At the anterior end it passes inwards, 
deep to the M. dorsalis humeralis, and then turns sharply ventralwards 
just anterior to the third rib, passing deep to the portion of the M. 
thoraci-scapularis arising therefrom. At this point a vessel is re- 
ceived from the M. thoraci-scapularis and the anterior body-muscles 
beneath the supra-scapula. This portion of the vein is called the 
cervical portion of the V. cutanea magna by Osawa, but since it has 
no connexion with the skin, and is a purely muscular vein, it would 
seem to be a misnomer. At the posterior border of the glenoid 
cavity it is joined by a cutaneous vein from the skin covering the M. 
dorsalis humeralis, the V. cutanea dorsalis scapulae -posterior (v.cut.d. 
sc.p.) and then almost immediately unites with the lateral branch of 
the V. epigastrica anterior to form the V. axillaris (v. ax.). At this 
point an anastomosis with the V. brachialis medialis may occur, but 
is not constant. At the posterior border of the scapula the V. dorsalis 



THE VASCULAR SYSTEM 233 

scapulae is received from the MM. dorsalis scapulae and dorsalis 
humeralis, and at about the same point a large branch from the V. 
brachialis medialis as well as the F. cutanea humeri -posterior from the 
M. anconaeus and the skin covering it. The vessel, which may now 
be called the V. circumflexa scapulae (v.circ.sc), passes lateral to the 
ventral end of the scapula, between this bone and the muscles cover- 
ing it, and dorsal to the head of the humerus. At the anterior border 
of the scapula the V. brachialis lateralis joins the vessel, as well as 
small factors from the MM. cucullaris and opercularis and also a 
larger cutaneous vein, the V. cutanea dorsalis scapulae anterior (v. cut. 
d.sc.a.) which latter drains the skin covering the M. dorsalis scapulae. 

The subclavian vein (v.scL), as it may now be called, continues 
directly forwards along the dorso-lateral margin of the procoracoid, 
and, near the anterior extremity thereof, receives the V. supracora- 
coidea (v.sup.cor.). This factor drains the muscles of the same name, 
and then penetrates the coracoid foramen and crosses the dorsal sur- 
face of the procoracoid almost at right angles to its long axis. After 
receiving the supracoracoid vein, the subclavian turns in mesially 
and joins the internal and external jugulars and post-cardinal to form 
the ductus Cuvieri. 

There is a small vessel joining the anterior aspect of the subclavian 
vein just before it enters the ductus Cuvieri which Bethge calls the 
V. cutanea parva (v.cut.pa.). It collects from the posterior margin of 
the thymus gland, and from the overlying skin, and it also receives 
some small twigs from the M. cucullaris. The hypoglossal nerve 
passes between this vein and the V. subclavia, and thus forms a useful 
landmark for identifying the vein. 

{c) Veins of the Fore-limb, These arise between the fingers as the 
FF. interdigitales which flow into a venous arc stretching across the 
dorsal surface of the hand at about the level of the centre of the meta- 
carpals — the Arcus venosus dorsalis manus (Osawa). On the radial side 
of the limb this arc flows into the F. superficialis radialis (Osawa), 
and at about the same point a large palmar cutaneous vein enters. 
The V. superficialis radialis continues along the fore-arm, just under 
the skin, between the flexor and extensor muscles of the radial side. 
It drains the muscles by several large twigs, and, at the elbow, it 
receives a large R. communicans from the V. interossea. From this 
point to the shoulder the vessel becomes the F. brachialis lateralis 
(Osawa). It receives, just proximal to the elbow, one or two large 
cutaneous factors, as well as several twigs from the adjoining muscles. 
At the proximal end of the arm, quite close to the shoulder, there is 
yet another deep plexus, lying on the flexor side of the humerus, 



234 THE SALAMANDER 

whereby communication is established between the lateral and medial 
brachial veins as well as with the V. anastomotica brachii. Directly 
anterior to the insertion of the M. procoraco-humeralis the V. brachi- 
alis lateralis joins the V. circumflexa scapulae to form the V. sub- 
clavia as described above. 

The V. interossea arises on the dorsum of the hand by several factors 
from the ulnar and middle regions of the arcus venosus dorsalis manus, 
and from the several extensor muscles of that region. These factors 
unite to form a single vessel at the proximal border of the carpus, just 
mesial to the distal end of the ulna, and pass along the fore-arm on 
the extensor side of the M. interosseus in company with the artery 
and nerves. At the proximal end of the muscle it passes to the flexor 
side of the limb and receives a large factor — the F. antibrachii ulnaris 
— draining the flexor muscles of the ulnar side of the limb, as well as 
the M. flexor primordialis communis, the MM. flex, accessorius 
medialis and lateralis, and the M. pronator profundus. It may also 
anastomose with the palmar factor of the V. super, radialis, but it is 
uncertain whether this is constant. 

At the flexor side of the elbow the combined vein divides into 
three. Of the two main limbs one fuses with the V. brachialis later- 
alis, the other becomes the V. brachialis jnedialis (v.br.m.) and passes 
along the ventral, or mesial side of the arm. The third branch, much 
smaller than the other two, is the V. anastomotica brachii. Not infre- 
quently the V. brachialis medialis and the V. anastomotica brachii 
arise as a single vessel and divide later. 

As the V. brachialis medialis becomes superficial it receives one 
or two large cutaneous veins — VF. cutaneae antibrachii mediales — from 
the skin covering the ulnar side of the fore-arm. It also drains both 
the extensor and flexor muscles of the arm. Near the shoulder it 
receives a fairly large factor whereby it communicates, through the 
plexus above mentioned, with the V. brachialis lateralis and V. anas- 
tomotica. It then turns somewhat posteriorly, and, after receiving 
another factor from the skin of the arm, it joins, or sends a large 
branch to, the V. axillaris. In the latter case the remaining portion 
passes antero-mesially beneath the scapula, ventral to the nerves of 
the brachial plexus, and, after receiving a small twig or two from the 
M. thoraci-scapularis, enters the post-cardinal vein close to the point 
where it bends around the oesophagus. If the whole of the V. brachi- 
alis medialis joins the axillary vein then a branch from the common 
vessel thus formed passes to the post-cardinal. The relations in this 
region are subject to a considerable amount of variation, and, in one 
case, the vessel passing to the post-cardinal was so fine as to be almost 



THE VASCULAR SYSTEM 235 

non-existent, but a larger vessel, leaving the V. axillaris at about the 
same point, passed anteriorly, ventral to the nerves, and entered the 
subclavian vein anterior to the shoulder. Grodziriski (1930) states 
that the vessels supplying the developing fore-limb in Amblystoma 
form a network just proximal to the glenoid region, from which the 
subclavian and other veins develop by the enlargementof some vessels 
at the expense of others, so that the network ultimately disappears. 
Assuming that the development of Salamandra follows similar lines, 
it is easy to understand the rather considerable variations in the 
connexions just described. 

The V. anastomotica hrachii (v.anas.br.) has already been mentioned 
several times. It is a curious little vessel which arises in the plexus 
just proximal to the elbow and passes up the flexor side of the arm 
between the two main veins. It passes right across the ventral side 
of the shoulder-joint and penetrates the M. coraco-brachialis brevis 
to join the V. pectoralis (cf. p. 231). It drains the muscles alongside 
which it passes, and communicates with the other two brachial veins 
at the plexus just distal to the shoulder, as already described. 

It should be noted that the vessel which Bethge calls the 'Vena 
cutanea magna' comprises those here described under V. lateralis, 
V. axillaris, V. circumflexa scapulae, and V. subclavia. 

Veins of the Lungs. 

The pulmonary veins arise on the mesial aspect of the lungs and 
pass transversely from the proximal ends of those organs across the 
ventral wall of the oesophagus, where they fuse, near the middle 
line, to form a single unpaired vessel which runs directly forwards 
and enters the left auricle of the heart. The junction of the right 
and left pulmonary veins occurs immediately dorsal to the apex of 
the ventricle. 

A few small factors from the ventral oesophageal wall also enter 
the pulmonary veins. As the pulmonary vein passes between the 
oesophagus and the sinus venosus it becomes embedded in the tissues 
of the dorsal wall of the latter. Its entry into the left auricle is not 
valved (see also p. 188). 

Veins of the Portal Systems 

The Hepatic Portal System (PI. XXII). 

A. F. ahdominalis (v.abd.) (Hochstetter) (see also PI. XVI). An 
abdominal vein is present as in the Frog, but instead of being formed 
largely by the fusion of the two pelvic veins, as in that animal, it 



236 THE SALAMANDER 

also receives the considerable addition of the following two median 

veins in the pelvic region. 

(a) The median cloacal vein (v.cl.m.) arises from the anterior and 
mesial regions of the cloaca by several fine factors and runs forwards 
in the middle line ventrally, along the pubo-ischiadic symphysis, 
immediately under the skin. At the ypsiloid cartilage it bends round, 
either to the right or left, and joins the abdominal vein in the middle 
line at its formation by the fusion of the right and left pelvic veins. 

(J)) T\\t posterior vesica/ vein (Fig. 63 ; v.ves.p.) enters the abdominal 
at the same point, but lies dorsa/ to the pubo-ischiadic symphysis. It 
drains the blood from a rich vascular network spread over the pos- 
terior region of the bladder, and receives a pair of veins from the 
dorsal side of the pubo-ischium. These small factors are readily 
broken when the pubo-ischiadic symphysis is split and pulled apart, 
so that they are easily overlooked. They receive a small tributary 
which passes through the obturator foramen from the ventral side. 

The abdominal vein passes forwards in the middle line from the 
anterior edge of the pubes to the anterior extremity of the bladder 
(about 14 mm.) through the subperitoneal epithelium, without re- 
ceiving any further factor of importance other than a small ypsiloid 
vein at the anterior end of the ypsiloid cartilage from the surrounding 
tissues and skin of that region. 

At the anterior end of the bladder it is joined by vessels from this 
organ. 

(c) One or more anterior vesical veins (Fig. (>y^ v.ves.a.) collecting 
the blood from the anterior, ventral, and lateral regions of the bladder. 

(^) The vesico-haemorrhoidal vein (v.ves.hm.) comprising two main 
factors. 

(i) A dorsal vesical vein formed by many factors from the dorsal 
bladder-wall. These usually unite to form a single vessel before join- 
ing the other factor, but may enter at several points. 

(ii) A ventral rectal vein. It is important to notice that the whole of 
this vessel lies ventral to the gut, i.e. between it and the bladder. 
The blood from the dorsal side of the rectum enters the portal vein 
by means of the haemorrhoidal vein, but there appear to be frequent 
fusions within the tissue of the rectum between the factors of these 
two vessels. 

The combined dorsal vesical and ventral rectal veins form the 
vesico-haemorrhoidal vein which enters the abdominal a little an- 
terior to the bladder. 

In one case the abdominal vein divided at this point to reunite a 
little farther on, forming a closed loop. Normally, however, it con- 



THE VASCULAR SYSTEM 237 

tinues undivided to the liver. It passes dorsal to this organ, and 
about 5 mm. from its edge, fuses with the anterior gastric branch of 
the portal vein, which, continuing along the dorsal side of the liver, 
appears to be a direct continuation of the abdominal vein. The sec- 
tion between the anterior end of the bladder and the posterior edge 
of the liver receives some two or three tributaries. 

{e) Several factors from the ventral body-wall and the rectus ab- 
dominis muscle. These factors frequently, if not invariably, anasto- 
mose with factors of the anterior epigastric veins. They often, instead 
of entering the abdominal vein direct, enter the median epigastric 
vein (see below) to which they probably belong. 

(J) A small factor from the proximal end of the duodenum may 
frequently be found entering the ventral side of the abdominal vein 
at the point where it crosses that section of the gut. 

(^) The median epigastric vein (y. cm.) is represented by one or two 
factors of the abdominal vein lying ventral to it and collecting from 
the ventral parietes. It corresponds very closely with the similarly 
named vein in Reptiles as described by Beddard (1906) and O'Dono- 
ghue (1921). 

The epigastric veins are notoriously difficult to investigate, and 
the Salamander is no exception. So far as could be determined the 
median epigastric vein extends anteriorly through the subperitoneal 
epithelium as far as the sternal cartilage. Whether it anastomoses 
with the sternal vein could not be determined with certainty, since 
the pressure of the cartilage squeezed all the injection fluid and blood 
from this section. In any case if such a connexion exists it is very 
small. The vein receives several lateral factors which in some cases 
appear to anastomose with corresponding factors of the anterior 
epigastric veins. They correspond roughly in number and position 
with the inscriptions in the rectus abdominis muscle. (For relations 
of the anterior epigastric veins see pp. 2 30, et seq.) A comparison with 
the work of Beddard and O'Donoghue will readily demonstrate the 
striking general agreement between the abdominal, pelvic, and epi- 
gastric veins of the Salamander and the reptiles concerned, particu- 
larly Sphenodon, a fact which has also been noted by O'Donoghue in 
his account of the vascular system of this animal. 

B. The F. porta (v.p.) arises dorsally at the level of the pylorus 
(i.e. at about the middle of the body cavity) by the junction of the 
lieno-gastric vein with the mesenteric veins. It runs in a dorso- 
ventral direction, slightly obliquely anteriorly, passing through the 
pancreas a little posterior to the hepatic artery. 

It should be noted that the factors and branches of the portal vein 



238 THE SALAMANDER 

pass through the mesentery on the left side, and the branches of the 

mesenteric arteries on the right. 

The factors of the -portal vein, 

{a) The V. gastrico-lienalis (v.g.lien.) runs posteriorly through the 
mesentery which suspends the spleen from the dorsal stomach-wall. It 
is constituted by a series of tributaries, on the one side from the gland, 
and on the other from the dorsal stomach-wall, and is the venous 
counterpart of the lieno-gastric artery which runs more or less 
parallel with the vein, between it and the stomach. 

Just before its junction with the mesenteric veins it gives off a 
small branch which may be termed the minor portal vein. This 
vessel passes along the postero-ventral side of the post-caval lobe of 
the liver. 

As was the case with the arterial system it is not easy to differenti- 
ate the anterior from the posterior mesenteric vessels. The following 
arrangement seems to be the most satisfactory and convenient 
generally. 

{U) The V. haemorrhoidalis (v.hm.) drains the blood from the dorsal 
side of the rectum, and the extreme posterior end of the intestine. 
A short distance before it unites with the lieno-gastric vein at the 
origin of the portal vein it is frequently joined by the posterior intes- 
tinal vein. 

{c) The V. intestinalis posterior (v.in.p.) is the largest of the three 
intestinal veins. It returns the blood from the posterior two-thirds 
of the intestine (excluding the rectum). It always enters the portal 
vein in very close association with the haemorrhoidal vein, and as 
already stated they frequently unite and enter as a single vessel. 

{d) and (<?) The middle and anterior intestinal veins. These two 
factors enter the portal vein at intervals of about a millimetre along 
its length. Their origin in the intestine is very restricted, and to- 
gether they return the blood from the remaining third of the intestine 
and the posterior end of the duodenum. There is much variation. 

(/) Rusconis vein (v.R.) (Hochstetter), or V. omphalo-mesenterica 
sinistra (Choronshitzky). This vessel arises from numerous factors 
in the duodenum and anterior part of the intestine, and passes cepha- 
lad, parallel with the gut, to about the middle of the duodenum, 
where it turns dorsalwards and enters the pancreas to join the pan- 
creatic vein, the common trunk thus formed finally uniting with the 
portal vein just before this vessel joins the abdominal vein. 

The vessel under consideration was first described by Rusconi, 
who referred to it as the longitudinal vein of the gut, while its onto- 



THE VASCULAR SYSTEM 239 

genetic history has been investigated by Choronshitzky (1900), who 
showed that it is really the persisting left vitelline vein of the embryo. 
It is therefore homologous with the V. subintestinalis of the fish. 
According to this author the right vitelline vein is never large and 
disappears early. Hochstetter (1908) confirms these views. 

{£) The anterior gastric vein.^ It would perhaps seem more correct 
to regard this vessel as a branch of the portal vein than as 2. factor^ but 
actually it is both. It passes along the dorsal surface of the liver and 
has the appearance of being the direct continuation of the abdominal 
vein. On the one side it sends numerous branches into the liver, and 
on the other it receives four or five factors from the ventral stomach- 
wall. These factors are always associated with corresponding arteries, 
and there is usually an artery on either side of the venous factor. 

It should be noted that the blood-system of the stomach is of a two- 
fold nature. It both receives its blood from, and returns it into, 
distinct vessels on both dorsal and ventral sides. 



The Renal Portal System (PI. XVI). 

A. The F. caudalis (v.cd.), as its name implies, arises near the 
extremity of the tail and runs forwards through the haemal arches ven- 
tral to the caudal artery. Almost immediately after it emerges from 
the haemal canal it divides symmetrically into two vessels, which pass 
respectively to the dorso-lateral border of each kidney. These veins 
are portal to the kidneys. Hochstetter's name for them is retained, 
namely, Jacobson's veins. 

VV. Jacobsoni (v.J.) (Hochstetter). Each Jacobson's vein, al- 
most immediately after its origin from the caudal vein, receives a 
small inguinal vein from the muscles of the inguinal region, and then 
passes to the dorso-lateral surface of the kidney, to which organ it 
gives a number of Venae advehentes. Just before reaching the level 
of the iliac artery Jacobson's vein bends round the edge of the kidney 
to the ventral side, passes ventral to the artery, and immediately 
anterior to it anastomoses with the common iliac vein. Jacobson's 
vein, thus reinforced, passes anteriorly along the lateral border of 
the kidney, giving off on the one side numerous Fenae advehentes 
to the kidney, and receiving on the other four or five dorso-lumbar 
veins from the muscles of the back, and also a number of posterior 

' This terminology is due to O'Donoghue (1921). Beddard (1905) regards it as the 
anterior continuation of the abdominal vein. As O'Donoghue rightly points out, only a 
study of its development can decide to w^hich it belongs. Reference should be made to 
O'Donoghue's paper. 



240 THE SALAMANDER 

oviducal veins from the posterior region of the oviduct. (For the 
entry of the pelvic vein see under this vessel.) If the injection is good 
it is usually possible to trace Jacobson's vein to the anterior extrem- 
ity of the kidney, but its size in this region varies very much. It 
may remain fairly large, and anastomose with the post-cardinal 
vein, but more usually it tapers to an almost indistinguishable 
thread. 

B. The Common Iliac Vein (v.il.com.). The blood from the hind- 
limbs has, in all Amphibia, the choice of two courses. It may go 
through the pelvic and abdominal veins to the liver, or it may enter 
the renal-portal system and pass through the kidney to enter the post- 
caval vein. It is to provide for this latter course that common iliac 
veins exist. Each lies on the dorsal side of the pelvis just posterior to 
the ilium and to N. spinalis 1 7, and dorsal to the common iliac artery 
of the corresponding side, and forms what appears to be the direct 
continuation of the V. ischiadica, but is marked off from this vessel 
by the origin of the V. iliaca transversa and the entrance of V. 
pudenda (see Figs. 62 and 6'^). The common iliac vein joins Jacob- 
son's vein a short distance from the posterior end of the kidney and 
usually a little to the ventral side of that organ. 

The veins of the hind-limb must^ of course, be regarded as commenc- 
ing at the distal extremities of the digits. The blood from the toes 
collects into the digital veins which lie along the sides of each digit, 
and, near the bases of the metatarsals, enter a transverse vessel lying 
across the dorsal side of the foot — the Arcus venosus dorsalis pedis. In 
addition to the above-mentioned veins from the toes this arc receives 
several tributaries from the extensor muscles of the foot and tarsus. 
In turn it delivers its blood into three main vessels. Of these two are 
superficial and pass along the tibial and fibular sides of the leg, 
corresponding respectively to the NN. dorsalis pedis tibialis and 
fibularis, while the third and largest vein is a deep vessel and courses 
in the interosseal space between the bones of the leg in company with 
the NN. dors, pedis intermedius and interosseus. It lies superficial to 
the A. interossea and deep to the above-named nerves. Just proximal 
to the knee these three veins unite to form a single large vein, which 
passes along the posterior aspect of the thigh together with the 
sciatic nerve. 

This is rather different from the condition found in the Frog, but 
bears a much closer resemblance to Osawa's account of Crypto- 
branchus. The difference may probably be accounted for by the 
specialized nature of the hind-limbs in the Anura. 

A comparison with the fore-limb and with Osawa's description 



THE VASCULAR SYSTEM 241 

of Cryptobranchus suggests that the two superficial vessels may be 
called V. lateralis cruris and V. medialis cruris on the tibial and 
fibular sides respectively, and the deep vessel the V. interossea. 

The main factors of the V. lateralis cruris. 

(a) A plantar vein from the skin of the sole of the foot as well as 
from the muscles of the same region. 

(b) A cutaneous factor from the plexus covering the extensor surface 
of the leg. Through this plexus communication with the V. medialis 
cruris is established. 

(c) A fairly large factor from the deeper muscles enters the V. 
lat. cruris at about the middle of the leg. This factor anastomoses 
with one from the V. interossea. 

(d) Just proximal to the knee one or two cutaneous factors from 
the flexor side of the leg are received. 

(e) At about the same level a small muscular factor from the inser- 
tion of the M. pub. isch. tib. and the tissues around the knee-capsule 
enters the vessel. 

At the knee the vein turns somewhat dorsalwards and passes into 
the thigh, dorsal to the M. pubo-tibialis, to join the F. circumflexa 
genu, which is itself a factor of the V. ischiadica. 

On the post-axial or fibular side of the limb the V. medialis cruris 
arises from the Arcus venosus dorsalis pedis and receives a large 
factor from the dorsal plexus of the leg, whereby communication 
with the V. lat. cruris is established. Just distal to the knee it may 
receive a factor which drains the extensor and flexor muscles and lies 
close to the external border of the fibula, or this factor may enter the 
V. poplitea separately. At about the same point it receives a cutan- 
eous factor which probably represents the V. cutanea lat. sup. 
(Gaupp) of the Frog. The main vein then turns into the thigh 
proximal to the knee and immediately dorsal to the MM. ischio- 
flexorius and flexor fibularis, between which muscles and the M. 
extensor ilio-tibialis it joins the V. poplitea. 

The F. interossea is the largest of the three crural veins, and, as 
already mentioned, passes along the interosseal space together with 
the corresponding artery and nerves. It arises from the middle part 
of the Arc. ven. dors. ped. by factors which cross the dorsal side of 
the tarsus, deep to the extensor muscles, and unite to form a single 
trunk at its proximal edge. During the course of the vein down the 
leg it receives several factors from the surrounding muscles. Within 
the bend of the knee a cutaneous factor from the flexor side of the 
leg is received, as well as the V. medialis cruris. The vein may now 

4038 R 



242 THE SALAMANDER 

be called F, poplitea^ and is soon joined by the V. circumflexa genu 

to form the F. ischiadka. 

The F. circumflexa genu is a short vessel joining the sciatic vein 
almost at right angles just proximal to the knee. As stated above it 
joins the V. lat. cruris, while it also receives twigs from the knee- 
capsule and a longitudinal recurrent vessel^ which lies in the thigh 
between the MM. pubo-tibialis and pubo-ischio-femoralis internus, 
i.e. along the anterior border of the limb. This vessel drains the 
adjoining muscles and also receives large cutaneous factors from the 
skin covering that region (possibly to be regarded as the VV. cutaneae 
femoris anteriores mediales). It is usually traceable towards the body 
as a very fine vessel which finally enters the pelvic vein, or anasto- 
moses with a small factor thereof. There is therefore a continuous 
but ill-defined venous channel passing along the whole length of the 
pre-axial margin of the hind-limb, corresponding with the V. brachialis 
lateralis of the fore-limb, which is doubtless the homologue of the 
V. femoralis lateralis (Osawa) of Cryptobranchus. The position of 
this channel and the fact that it is the only vein entering the pelvic 
loop from the hind-limb anterior to the ilio-femoral joint would sug- 
gest that it is to be regarded as the representative, in the Salamander, 
of the femoral vein, which otherwise is wanting. A second anasto- 
mosis with the V. ischiadica occurs at about the middle of the thigh 
by means of the V. circ. fem. med. (see below), and hence the vessel 
may be called F. collateralis femoris (v.col.fem.). 

The F. ischiadica (v.is.). After its origin from the union of the 
popliteal vein with the V. circ. genu, as above described, the sciatic 
vein continues along the posterior aspect of the thigh and receives 
the following factors. 

(a) Posteriorly, the F. profunda femoris posterior enters from the 
MM. ischio-flexorius and pubo-ischio-femoralis externus and from 
the overlying skin. The cutaneous factor is the F. cut. fem. medialis 
(Gaupp). ^ 

(b) Mesially a pair of FF. circumflexae femoris mediales are received, 
one passing dorsal and the other ventral to the femur. They are both 
muscular factors draining the thigh muscles. The ventral one anasto- 
moses with the V. collateralis femoris. 

At the level of the hip-joint, in what Appleton conveniently terms 
the 'posterior groin', the V. ischiadica unites with the F. iliaca trans- 
versa and F. pudenda to form the common iliac vein. The sciatic vein 
thus forms the main vein of the thigh in the Salamander and carries 
almost the whole of the blood from the limb towards the trunk, whereas 
in the Frog (and in man and mammals generally) this function is 



THE VASCULAR SYSTEM 243 

assumed by the femoral vein. In Cryptobranchus, judging from 
Osawa's account, the V. femoralis lateraHs (=femoral vein) and the V. 
femoralis medialis (=sciatic vein) are approximately equal in size and 
importance. Hence there seems to be a twofold departure from the 
Cryptobranchid type, one where the posl-axia/ vein becomes the im- 
portant vessel — e.g. Salamandra — and the other in which the pre-axia/ 
path carries most of the blood — e.g. the Frog and the Amniota in 
general. 

The F. pudenda (v. pud.) enters the proximal end of the V. ischia- 
dica at the point where it merges into the common iliac vein. It 
arises in the female from the lateral wall of the cloaca, and in the male 
from the lateral portion of the cloacal gland, and runs antero-dorsally 
between the MM. ischio-caudalis and caudali-pubo-ischio-tibialis, 
receiving factors from both these muscles. At the dorsal border of 
the former muscle it receives an anastomosis from the median cloacal 
vein, while just before its entry into the ischiadico-iliac junction it 
receives several small cutaneous factors from the skin at the root of 
the tail, and sometimes a rather large anastomosis from the caudal 
section of the lateral vein, which probably represents a portion of the 
inferior lateral cutaneous vein of Selachian fishes. ^ 

The F. iliaca transversa (Rusconi) (v.il.tr.). Leaving the junction 
of the sciatic and iliac veins from the point where the V. pudenda 
enters, and passing round the dorsal side of the hip-joint to the 
region of the groin immediately beneath the skin, is a large vein 
which has been called V. iliaca transversa. It corresponds with the 
similarly named vein in the Frog, but it is a very much larger and 
more important vessel in the Salamander than in that animal. At the 
anterior edge of the ilium it enters the pelvic vein — in fact the major 
portion of the blood flowing through the ventral section of the pelvic 
veins into the abdominal vein arrives there through the V. iliaca 
transversa, since, in almost all cases, it is decidedly larger than the 
dorsal portion of the pelvic vein (cf. below). 

Factors of the F. iliaca transversa. 

(i) The anastomosis above described as sometimes entering the 
pudendal vein may enter the V. iliaca transversa direct. 

(ii) The F, cutanea femoris posterior lateralis (Gaupp) from the skin 
covering the dorso-lateral aspect of the proximal portion of the thigh. 

(iii) A twig from the M. extensor ilio-tibialis. 

(iv) A cutaneous factor from the lateral parietes immediately an- 
terior to the limb. This factor not infrequently anastomoses with the 

' Cf. Daniell and Stoker 1927, and Daniell's Elasmobranch Fishes, California 1928. 



244 THE SALAMANDER 

abdominal portion of the V. lateralis, or with one of its factors, and 
enters the V. iliaca transversa close to the point where this vessel 
joins the pelvic vein. When the fusion between the factors of the 
lateral vein and the V. iliaca transversa occurs, the condition closely 
resembles that found in some reptiles, e.g. Sphenodon (O'Dono- 
ghue, 1 921) and Heloderma (Beddard, 1906). 

C. The Pelvic Veins (v.pel.), as in the Frog, offer an alternative 
course to the blood coming from the hind-limbs, and enable it to avoid 
the kidney. They arise normally on either side from Jacobson's vein a 
little anterior to the common iliac vein, and pass around the anterior 
rim of the pelvic basin, internal to all the muscles, to the mid-ventral 
line, where they fuse to form the abdominal vein. It is convenient to 
distinguish two sections in each pelvic vein, viz. a dorsal section from 
Jacobson's vein to the point where the V. iliaca transversa is received, 
and a ventral section from that point to the abdominal vein. The 
dorsal section varies considerably both as to its exact point of origin 
from Jacobson's vein and in size. It is never very large, and seems 
sometimes to be absent entirely. The ventral section is very much 
larger since it receives the considerable addition of blood from the 
V. iliaca transversa. It also receives the V. -parietalis lateralis (v. par. 1.) 
(lateral abdominal, of some authors) from the lateral abdominal 
muscles, and the small vein from the anterior aspect of the thigh 
joining the V, collateralis jemoris (v.col.fem.) which may represent 
the femoral vein. Small twigs from the origin of the M. pubo-ischio- 
femoralis internus and from the insertion of the rectus abdominis 
muscles are also factors of this section of the pelvic vein. 

Vessels of the Brain (PI. IX, figs. 51 and 52). 

A. Arteries. As already described the internal carotid artery 
passes into the cranial cavity through the carotid canal and the basi- 
cranial fenestra. In the cranial cavity it is known as the A. carotis 
cerebralis (a.car.ce.) (Scholb) which almost at once divides into a R. 
anterius and a R. posterius. 

(I) R. anterius (a.ce.a.) (Scholb) courses along the lateral border 
of the thalamencephalon to the level of the optic chiasma, where it 
divides into dorsal and ventral branches. 

{a) The ventral branch, R. cerebralis anterior ventralis (Osawa), 
Arteria lobi haemisphaerici inferior externa (Scholb), or more simply 
the ventral cerebral 2iTt&TY(2i.ct.Y.)y runs along the ventro-lateral edge 
of the cerebrum as far as the olfactory nerves. From it vessels arise 
which form a rich network in the primary vascular coat (see 'Mem- 



THE VASCULAR SYSTEM 245 

branes of the Brain', p. 123, et seq.). These do not arise periodically 
along the whole length of the vessel, but almost exclusively from 
near its origin, and at its anterior end. They are drained by the dorso- 
median vein of the cerebrum. 

(b) The dorsal branch, R. cerebralis anterior dorsalis (Osawa) 
(Arteria lobi haemisphaerici superior interna, Scholb) passes dorso- 
mesially round the thalamencephalon to the paraphysial region 
(nodus chorioideus. Rex), giving on its way a branch to the posterior 
and mesial regions of the cerebral hemisphere. 

(i) The mesial cerebral artery passes anteriorly along the mesial face 
of the hemisphere somewhat obliquely in a dorso-ventral direction. 
This vessel does not appear to have been previously described unless 
it corresponds with the 'posterior dorso-median vein of the fore- 
brain' of Rex, which otherwise has not been found. 

At the nodus chorioideus the dorsal cerebral artery sends a few 
small branches to the plexus and then turns ventralwards and divides 
into two branches. 

(ii) One branch supplies the lateral plexus of the first ventricle 
(pl.ch.ven.i). The artery enters the ventricle through the foramen 
of Monro, and within it forms a very beautiful coiled plexus. If this 
plexus is carefully flattened out it is found to be roughly triangular 
in shape, with the artery bordering the perpendicular and base, and 
the veins gathering towards the apex. 

(iii) The other branch supplies the inferior median plexus (Burck- 
hardt's plexus chorioideus inferior), a small, but rich plexus lying in 
the recessus praeopticus. 

The superior median plexus appears to be supplied usually by a 
branch from the right dorsal cerebral artery. This branch comes 
off just as the main vessel turns ventralwards. The plexus is long, 
tapered to a point at its posterior end, and flattened laterally. It lines 
in the dorsal part of the cavity of the diencephalon. 

It should be noted that the right and left dorsal cerebral arteries 
are not always equal in size. Either may develop at the expense of 
the other. 

(II) Ramus posterius (Scholb) or R. cerebralis posterior (Osawa). 
This branch passes directly mesialwards to a point on the ventro- 
lateral border of the midbrain, just at the posterior border of the 
infundibulum and anterior to the origin of the Ilird nerve. Here it 
breaks up into four branches. 

{a) R. opticus ^Vf\i\Q\s. ramifies fan-wise over the side of the optic lobe. 

(^) The R. pituitarius which passes round the infundibulum to its 
ventral surface and then turns posteriorlytosupply the pituitary body. 



246 THE SALAMANDER 

(c) R. mesencephali superior (Gaupp, Frog). This branch passes 
round between the optic lobes and the cerebellum. It supplies the 
latter, and finally terminates within the chorioid plexus of the fourth 
ventricle. 

((T) R. communicans continues the direction of the main stem 
mesially, and anastomoses with the corresponding branch from the 
other side. From the transverse vessel thus formed there arises, in 
the middle line, the basilar artery. 

The Arteria hasilaris (a.b.) (Gaupp, Frog) arises by two or three 
roots, which almost immediately fuse to form a median vessel cours- 
ing along the mid-ventral line of the medulla oblongata, and giving 
numerous fine branches to it on either side. The auditory artery is 
given off just anterior to nerve V. 

The Arteria auditiva (Gaupp, Frog) passes into the auditory cap- 
sule along with nerve VIII. 

The A. basilaris is continued right along the vertebral column, 
where it is called the A. spinalis ventralis (Gaupp, Frog). It is 
the artery of the spinal cord. 

B. Veins. It is not possible to give a connected account of veins 
of the brain since they converge on more than one point. The follow- 
ing description therefore commences at the anterior end of the brain 
and works backwards. 

The sinus sagittalis (s.sag.) (Rex) is a vein of variable size. It 
arises in the primary vascular coat anteriorly and passes backwards 
in the middle line to the nodus chorioideus (Rex) — a triangular 
venous plexus in the paraphysial region (nod.ch.). At about the 
middle of its length it is considerably strengthened by the addition of 
a dorso-median cerebral vQin (Rex, translated), bringing the blood from 
the hemispheres. The vein arises by the fusion of numerous factors 
from the ventral and mesial surfaces of the hemispheres, and, pass- 
ing up between them in the sagittal cleft through the subdural space, 
enters the sinus sagittalis. It is not always possible to trace the sinus 
anterior to the entry of this vein. In any case it is very small. Rex 
has described two dorso-median cerebral veins, but only one has been 
found. It is possible he may have confused the mesial cerebral artery 
with the venous system (see p. 245). The blood from the various 
plexuses within the ventricles of the brain (see under 'Arteries') is 
also returned to the nodus chorioideus by numerous small veins. 

The blood is carried from the nodus chorioideus by a pair of veins, 
one of which runs from each lateral angle of the triangular nodus. 

The oblique cranial veins (v.cr.ob.) (Venae craniales obliquae, 
Gaupp, Frog) pass obliquely from the nodus chorioideus between 



THE VASCULAR SYSTEM 247 

the posterior end of the cerebrum and the corpus opticum, and dis- 
charge into the venous plexus covering the sinus endolymphaticus 
(plexus lateralis, Scholb). About midway they receive lateral tribu- 
taries on either side. 

Each lateral cerebral vein collects from the dorsal and lateral 
regions of the fore-brain. 

The dura mater covering the diencephalon is fairly free from 
veins, and no vessel of any size occurs in it. The primary vascular 
coat, on the other hand, possesses a comparatively rich network of 
veins. This network collects into two dorsal longitudinal vessels 
(v.long.di.) near the middle line, which apparently correspond with 
the single, median, Vena longitudinalis diencephali (Gaupp) of the 
Frog. From the anterior end of these veins one or two vessels pass 
out through the subdural space and discharge into the plexus cover- 
ing the sinus endolymphaticus. 

The blood from the cerebellum collects into a vessel which enters 
the plexus covering the fourth ventricle. 

The lateral plexus (pl.l.), covering the sinus endolymphaticus 
(Scholb's name is retained for sake of convenience), is a coarse venous 
network forming a hollow pyramid with a triangular base on either 
side of the diencephalon, the base being directed inwards and the 
apex outwards. The blood gathers into a single large vessel at the 
apex of the pyramid and passes out of the cranial cavity, ventral to 
the R. oph. profundus V, and enters the Vena petrosa lateralis (q.v.). 
This is the Vena cranialis pro-otica (v.cr.pr-ot.) (Gaupp), and is the main 
venous exit from the cranial cavity. It corresponds with the Vena 
cerebralis anterior of van Gelderen. 

The anterior part of the fourth ventricle is roofed over by a very 
rich plexus, more or less rhomboidal in shape, and communicating 
at its anterior corners with the lateral plexuses. There is usually a 
fairly pronounced vessel running transversely across the anterior 
edge of this plexus. Entering the plexus dorsally are two longitudinal 
medullary veins which pass anteriorly along the borders of the pos- 
terior part of the fourth ventricle. They evidently correspond with 
the Venae craniales post-occipitales (Gaupp) of the Frog, but they 
are not nearly so pronounced in the Salamander, and do not com- 
municate, other than very indirectly, with the Vena spinalis superior. 

Rex has described for Triton several basal veins entering this 
plexus from the ventral side of the medulla oblongata, but, while the 
veins from the ventral side of the medulla of Salamandra certainly 
do communicate quite freely with the plexus, no vessels of sufficient 
distinction to warrant a name have been found. 



248 THE SALAMANDER 

There is, however, a Vena communicans forming an anastomosis 
between the Vena craniaHs pro-otica of either side. It passes between 
the roots of nerve V, dorsal to the pituitary body, and receives a pair 
of tributaries therefrom. 

The -pituitary veins (v.pit.), one on either side, arise from the pos- 
terior edge of the pituitary body. 

At the posterior corners of the plexus covering the fourth ventricle 
the network collects into a single vessel which also receives one or 
two basal veins from the ventral side, and then passes out of the 
cranial cavity together with the combined nerves IX and X. This 
is the sinus jugularis of Rex, but to avoid confusion with the jugular 
vein it may be called Vena cranialis post-otica (v.cr.po-ot.). It enters 
the occipital vein (q.v.) and is equivalent to the Vena cerebralis pos- 
terior of van Gelderen. In Triton it forms the principal venous exit, 
the Vena cranialis pro-otica being quite small (Rex). 

The region of the medulla behind this point and the anterior 
section of the spinal cord are surrounded by an irregular, wide- 
meshed, venous network which again collects into a vessel leaving 
the neural canal through a foramen in the first vertebra, the Vena 
post-occipitalis (v.po-oc.) (Gaupp, Frog). 

Almost immediately behind this point a definite dorsal longitu- 
dinal vein maybe traced right down the spinal cord. This is the Vena 
spinalis superior (Gaupp), and collects the blood from the Arteria 
spinalis centralis. It is connected with the Venae post-occipitales by 
a fairly definite branch. 

Summary of Vessels of the Brain. 

It will have been noticed that the main arteries are almost entirely 
confined to the ventral surface, while the larger venous vessels occur 
dorsally. Further, the arteries are more closely applied to the neural 
tissue than are the veins. The connexions between the arteries and the 
veins within the substance of the brain have been shown by Scholb to 
consist of simple loops, which may be spirally twisted but never 
branched. 

The vascular plexuses fall under three distinct heads : 

(i) Those to which the blood is carried by arteries and drained 
by veins. These are the internal plexuses, viz. the lateral plexuses of 
the first ventricle^ and the superior and inferior median plexuses of the 
diencephalon. 

(ii) Those to which the blood is carried by both arteries and 
veins, viz. the nodus chorioideus and the plexus covering the fourth 
ventricle. 



THE VASCULAR SYSTEM 249 

(iii) Those receiving their blood from veins only — the lateral 
plexuses covering the sinus endolymphaticus. 

Rex describes all the plexuses as venous, although it is quite easy 
to trace the arterial connexions. He does not seem to have differen- 
tiated clearly between arteries and veins. 

The roof of the diencephalon is peculiar in that it is comparatively 
free from blood-vessels, but is bordered by four large plexuses. It 
forms a sort of 'island'. 

The main venous exit is alongside the Vth nerve, anterior to the 
auditory capsule. Smaller veins can usually be found leaving the 
cranial cavity together with most of the other cranial nerves. 

LYMPHATIC SYSTEM 
I. Historical. 

Panizza (1833) was the first thoroughly to investigate the lym- 
phatics of the Salamander, and, while his account is somewhat 
incomplete, and in one or two instances not quite correct, it is 
nevertheless a classic piece of work. His use of mercury as a medium 
for injection has been very sharply criticized by some later workers, 
notably Rusconi and Meyer, on account of the weight of this 
medium, which tends to deform the larger spaces. While this criti- 
cism undoubtedly has some justification it resulted in Panizza's work 
being discredited to an extent by no means deserved. Some five years 
laterBazin(i 838— 9) published a resume of Panizza's work in French, 
but his extract of the portion dealing with Salamandra is very brief. 

The work of the investigators immediately following Panizza 
(Rusconi, Meyer, Calori) was almost entirely confined to the lymph- 
atics of the viscera and the sub-vertebral trunks (DD. thoracici). 
It is in this region that the distortion caused by mercury would be 
most evident, and hence attention was concentrated upon it. 

J. Miiller (1833) ^^^ ^^ ^^^t to discover lymph hearts in the 
Salamander, and he described four, an anterior pair just behind the 
scapula and a posterior pair immediately caudad of the ilium. Meyer 
(1845) confirms the existence of these hearts and describes and 
figures an additional pair posterior to the scapula, making a total of 
six pairs. In 1 884 Weliky showed that a whole row of segmentally 
arranged contractile bladders exists, lying along the sulcus lateralis 
between the dorsal and ventral body-muscles. He also investigated 
the histology of the bladders, and found that they were provided with 
muscles characteristic of lymph hearts. He says 'Die Muskeln der 
Lymphherzen sind bei beiden untersuchten Thieren' {Salamandra 



2 50 THE SALAMANDER 

maculosa and Siredon pisciformis) 'quergestreift, meistens spindel- 
formig, mit excentrischen grossen Kernen'. Finally Greil (1903) 
completed our knowledge of the contractile portion of the lymphatic 
system by describing the central lymph heart, within the truncus 
arteriosus, dorsal to the aortic arches. 

An excellent detailed study of the caudal lymph hearts and of the 
caudal lymph vessels was made by Favaro in 1906, while in 19 12 
Hoyer and Udziela undertook an investigation of the lymphatics of 
the larva. Their work is very thorough, and is based on direct obser- 
vation by dissection of injected larvae as well as from serial sections 
of both injected and uninjected specimens. The condition they 
describe for the larvae needs but little amendment to make it applic- 
able to the adult, the chief points of difference being the considerable 
extension of the jugular sinuses and the comparative diminution of 
the dorsal and ventral longitudinal vessels. 

2. Technique. 

The present investigation was carried out on animals that had 
been killed with chloroform and injected immediately after death 
with a semi-gelatinous mass made up as follows : 

An appropriate quantity of best white gelatine is allowed to soak 
in a good excess of water for about twelve hours (overnight). The 
excess water is then drained off and the gelatine melted in the water 
it has absorbed. The resulting mass is referred to below as 'stock 
gelatine'. The injection mass consists of — 
60 gm. stock gelatine (melted) 

6 gm. potassium iodide 
60 cc. pure glycerine 
240 cc. sat. soln. of prussian blue in water. 
This mass is liquid slightly above room temperature, but 'sets' to 
about the consistency of the white of an egg. It flows freely, and the 
prussian blue does not precipitate in the cannula, and, what is more 
important where the larger sinuses are concerned, is freely mobile 
without the injected mass breaking up or rupturing the walls of the 
vessel. The 'setting' is a delayed action so that cold apparatus may 
conveniently be used. It may be hastened by immersing the animal, 
after injection, in formalin. Gravity pressure of about 150 cm. of 
water was used for the injection, and, the point of the fine glass 
cannula having been inserted under the skin of the palm of one hand, 
the skin of the other palm and the soles of the feet were pricked with 
a needle, and the extreme tip of the tail was cut off. When the injec- 
tion mass dripped more or less freely from all these four points 



THE VASCULAR SYSTEM 251 

(10—15 mins.) the operation was stopped and all the points of exit 
sealed (with pyroxylin) so as to prevent leakage, and the animal 
immersed in 5-10 per cent, formalin overnight to assist in 'setting' 
the jelly. 

3. Description (Pis. XVIII and XIX). 

For convenience of description the lymphatic system may be 
divided into the following sections : 

A. Subcutaneous network. 

B. Lymphatics of the limbs. 

C. ,, „ „ pelvic region and tail. 

D. ,, „ ,, trunk and viscera. 

E. ,, ,5 55 pectoral region. 

F. „ „ „ head. 

G. Lymph hearts and their relation to the blood-vessels. 

A. Subcutaneous network. 

In contrast to the Frog there are no extensive lymph-sacs imme- 
diately under the skin, but, with the exception of a few more or 
less well-defined areas, the subcutaneous lymphatics form a rich 
reticulum with irregular meshes. The mesio-ventral portion of the 
abdomen is very sparsely supplied with lymphatics, which are 
almost confined to two small longitudinal vessels running along 
the mesial edges of the M. rectus abdominis superficialis, and re- 
ceiving lateral tributaries segmentallyat each myoseptum. Posteriorly 
these vessels join the sub-pelvic plexus, and anteriorly they enter 
the pectoral network. They represent the trunci lymphatici longitu- 
dinales parahdominales (Grodzinski in Amblystoma). Lymphatics 
are also nearly absent from the dorsum of the trunk and tail, but are 
rich along the sides of the body. 

The sub-pelvic plexus is a rich, subcutaneous network with 
irregular meshes, extending forwards from the cloaca on the ventral 
side of the body, between the legs, as far as the anterior end of the 
ypsiloid cartilage. It covers the pelvic sinuses (see p. 1^'^ and to 
some extent the inguinal sinuses also. It communicates with the 
former sinuses on their ventro-mesial aspects at about the level of the 
transverse axis of the limbs. 

The sub-pectoral plexus similarly underlies the pectoral girdle, and 
extends from the sternum to the gular fold. It becomes richer and 
sinusoidal laterally in the region of the arms. 

The sub-maxillary plexus covers the whole of the skin of the mouth- 
floor, and becomes sinusoidal in the region of the gular fold. In 



252 THE SALAMANDER 

appearance it closely resembles a sinus in Lacerta, figured by Hoyer 
(Anat.Anz., Bd. 73) and named by him 'Sinus thyroideus', but in Sala- 
mandra the glands lie much deeper than the plexus and are not 
associated with it. It therefore seems preferable to avoid this name. 
The lateral plexuses underlie the skin right along the sides of the 
trunk and the base of the tail. They lie mainly ventral to the sulci 
laterales, but extend dorsalwards over the scapula and ilium. They 
are particularly rich in the neighbourhood of the limbs. 

B. Lymphatics of the Limbs. 

These were first described by Panizza with considerable accuracy, 
and later Hoyer and Udziela gave a more detailed account of their 
course in the larva. There is no essential difference between the 
fore- and hind-limbs so that a general description may serve both. 

The whole limb is covered by a subcutaneous network which may 
be imagined as arising along the sides of the digits. At the bases of 
the metacarpals (or metatarsals) between the digits there are small 
lymphatic sacs which lead by short vessels into a palmar (or plantar) 
sac covering the whole ventral surface of the hand (or foot). This 
sac lies between the skin and the fascia of the superficial flexor 
muscles, and is penetrated by a number of fibrous threads which 
attach the fascia to the skin, so that the sac is broken up into a reti- 
culum with very small meshes. It is in free communication proxi- 
mally with the network covering the rest of the limb. In the region of 
the elbow (or knee) this network tends to drain into a pair of vessels 
passing along the pre-axial and post-axial sides of the proximal part 
of the limb, which latter in turn drain into the sacs lying around the 
shoulder and hip respectively. Small lymphatics also accompany the 
blood-vessels within the deeper parts of the limbs and communicate 
freely with the vessels just described. 

C. Lymphatics of the Pelvic Region and Tail (Fig. 69). 

On the ventral side of the pelvis is the sub-pelvic network above 
described. The greater part of this may be lifted as a sheet disclosing 
a series of three pairs of deeper sinuses, which, nevertheless, still lie 
superficial to the muscles. They may be called, from their positions, 
cloacal sinuses, pelvic sinuses, and inguinal sinuses respectively. 

The cloacal sinus (s.ly.cl.) differs somewhat in extent in the two 
sexes. In both it extends as a broad sheet across the ventral surface, 
on either side of the cloacal opening. In the female it also extends 
dorsalwards lateral to the walls of the cloaca itself, while in the male 
there is a lymphatic network ramifying among the tubules of the 



THE VASCULAR SYSTEM 253 

cloacal gland. The cloacal sinus has two communications with the 
vesicular sinus surrounding the base of the bladder, namely, one 
mesial, close to the middle line, and one lateral, passing in company 
with the pudendal vessels and nerve, which may accordingly be 
termed the -pudendal trunk. Anteriorly the cloacal sinus is continued 
as the pelvic sinuses. 

The pelvic sinuses (s.ly.pel.) lie on either side of the middle line 
beneath the pelvis, and consist of a very coarse and irregular network 
with small meshes. They drain the sub-pelvic network, and, in part, 
the cloacal sinuses, and finally enter the inguinal sinuses at their 
antero-mesial angles. A small vessel may usually be traced leaving 
the anterior extremity of the pelvic sinuses and passing along the 
lateral edges of the ypsiloid cartilage to join the para-epigastric 
trunks. The sinuses of either side communicate with one another 
by several small vessels. 

The inguinal sinuses (s.ly.ing.) (sinus inguinales, of the German 
authors) lie lateral to the sinuses just described within the 'groin', and 
are really the ventral portion of a continuous sinus surrounding the 
base of the limb, the dorsal portion being known as the iliac sinus. 
After joining with the pelvic sinus anteriorly each inguinal sinus 
sends a trunk around the anterior and dorsal aspects of the limb 
which accompanies the V. iliaca transversa. This trunk receives the 
pre-axial trunk from the leg, and communicates freely with the sub- 
cutaneous network covering the ilium as well as with the iliac sinus. 

The iliac sinus (s. ly.il.) is a rather irregular sac lying at the posterior 
aspect of the leg immediately behind the ilium. It stands at the 
junction of several large sinuses which meet in that region, namely, 
the post-axial trunk from the hind-limb, the inguinal sinus, and the 
vesicular sinus from the bladder. It also communicates freely with 
the subcutaneous network overlying it, and by means of this network 
a fair proportion of the lymph from the iliac region enters the lateral 
lymph hearts of the tail and is discharged by them into the V. 
lateralis (see p. 232). The rest of the lymph passes through the 
truncus iliacus accompanying the iliac artery and vein, to enter the 
subvertebral trunk. 

All sinuses so far described are superficial to the muscles and 
skeleton. Immediately dorsal to the pubo-ischium, between it and 
the bladder, lies the vesicular sinus (s.ly.ves.). It is hammock-shaped 
and covers the ventral side of the neck of the bladder, and, as already 
mentioned, it communicates ventrally with the cloacal sinus, laterally 
with the iliac sinuses, while dorsally near its posterior margin it 
enters the peri-anal sinus. 



2 54 THE SALAMANDER 

The peri-ana/ sinus (s.ly.pan.), as its name suggests, surrounds the 
posterior end of the rectum, and lies between this portion of the gut 
and a sheath of fibrous tissue. It communicates ventrally with the 
vesicular sinus, dorsally with the subvertebral sinus, and laterally 
with the renal sinuses. 

Panizza saw a part of the cloacal vessels and described some of 
the connexions between the lymphatics of the anus, cloaca, and 
bladder. Hoyer and Udziela commenting on his account state 
that these connexions are very much smaller and less important in 
the larva. 

The caudal lymphatics have been described in considerable detail 
by Favaro for Salamandra atra. In the adult the main lymphatic 
trunks of the tail are the subvertebral vessels which run in the 
haemal canal and form a ladder-like network around the caudal 
artery. (According to Favaro there is only a single subvertebral 
trunk in the tail of Triton.) They are continuous with the sub- 
vertebral sinus of the abdomen. In the larva there is also a pair of 
lymphatic vessels just beneath the skin mid-dorsally and mid-ven- 
trally — the TT. lymphatici longitudinales dorsales et ventrales, but 
in the adult only the latter persist, and that only as a very fine vessel 
passing along the mid-ventral line in the groove between the lateral 
myomeres. 

D. Lymphatics oj the Trunk and Viscera (see also PI. XVI). 

These vessels have received considerable attention from the 
earlier workers and have been largely used to discredit Panizza's work 
by drawing attention to the distortion caused by his use of mercury. 

The lymphatics of the trunk comprise four pairs of longitudinal 
vessels. 

(i) The Trunci lymphatici longitudinales parabdominales (t.ly.l.pab.) 
(Grodzinski, in Amblystoma) have already been mentioned. They 
pass along the mesial edges of the M. rect. abd. superficialis. 

(ii) The TT. lymph, long, parepigastrici (t.ly.l.pe.) (Grodzinski) 
form a network around the epigastric arteries and veins. They are in 
communication posteriorly with the iliac and inguinal sinuses and 
anteriorly with the subscapular sinuses. 

(iii) The TT. lymph, long, laterales (t.ly.1.1.) (Hoyer and Udziela) 
lie immediately under the skin in the sulci laterales. 

(iv) The TT. lymph, long, subvertebrales (Hoyer and Udziela) are 
the largest of all and lie immediately ventral to the vertebral column 
(Fig. 6y, t.ly.l.sv.). 



THE VASCULAR SYSTEM 255 

The TT. lymph, long, laterales extend right along either side of the 
body from a short distance behind the sacrum to the posterior border 
of the scapula. They lie in the sulci laterales. In the larva they form 
fairly straight and well-defined vessels, but in the adult they assume 
a more zigzag course and are only distinguishable as larger vessels 
among the general lateral network. They are associated with the 
lateral lymph hearts, and through them discharge their lymph into 
the lateral veins. They are in communication posteriorly with the 
iliac sinuses by means of the subcutaneous network, while anteriorly 
they enter the subscapular sinuses. The relations between these 
trunks and the lymph hearts is further discussed on p. 260. 

The TT. lymph, long, suhvertehrales. Panizza supposed that the 
subvertebral lymphatic stem consisted of but a single trunk. Rus- 
coni assented to this and said that the dorsal aorta and the vessels 
arising therefrom were totally surrounded by a lymph space as a 
finger is by a glove, and that the arterial walls were attached to the 
walls of the lymphatic vessel by fibrous bands. Meyer was of the 
same opinion. The dorsal aorta was thus supposed to be freely sus- 
pended in a lymph sinus. These earlier authors had not seen the 
caudal section, which was discovered later by Favaro. 

By studying the larva Hoyer and Udziela showed that the sub- 
vertebral space is really to be regarded as a double trunk lying on 
either side of the aorta, the two elements of which are in frequent 
communication with one another by means of bridging vessels which 
pass both dorsal and ventral to the aorta. In the section between the 
kidneys the latter become so extensive as to give the appearance de- 
scribed by Rusconi. At about the level of the posterior extremity of 
the stomach, i.e. at the level of the pylorus, there is a large lymph sac 
extending ventrally between the mesenteric laminae — the cisterna 
linfatica (Panizza), while one limb of it extends into the ligamentum 
gastro-lienale. Into this sac the lymph from the whole gut drains, 
being carried thither by a number of small lymphatic vessels which 
run on either side of the blood-vessels. The lymph capillaries within 
the gut-wall are definitely external to the blood-vessels and they are 
arranged in different patterns, each characteristic of some particular 
region of the gut. On the stomach they form a fairly coarse, irregular, 
brick-like meshwork; on the duodenum and intestine they are 
arranged in a number of more or less parallel rows connected by 
bridge pieces so as to give a ladder-like appearance, while on the 
rectum the pattern assumes that of a rich small-meshed network. 

From the subvertebral stem there are extensive lateral communi- 
cations with what may be described as the renal sinuses. 



256 THE SALAMANDER 

The renal sinuses (s.ly.ren.) are extensive sacs occupying the space 
between the mesenteric laminae suspending the oviducts in the 
female, or the Wolffian duct in the male. They are particularly 
extensive in the posterior region and are interrupted only by the blood- 
vessels or ureters, which also traverse the mesentery. There are also 
smaller extensions into the mesovarium. The walls of the oviduct 
are richly supplied with lymphatic capillaries forming a network. 

The pair of subvertebral trunks emerge again from the cisterna 
lymphatica anteriorly, and, although they are still extensively con- 
nected by bridges, their double nature is more apparent. At the 
level of the subclavian arteries a pair of transverse trunks pass out, 
parallel with these vessels, to the subscapular sinuses. The subverte- 
bral trunks themselves follow the curve of the systemic arches, and, 
after joining with the axillary and jugular sinuses, enter the central 
lymph heart (see p. 260). It is convenient to call this loop truncus 
peri-pharyngeus (Fig. 70; t.ly.p-ph.). The section of the subvertebral 
trunks just described, i.e. the region anterior to the cisterna lym- 
phatica, is the 'ductus thoracicus' of Panizza, who erroneously sup- 
posed that there was a single trunk only which split into two at the 
systemic arches. He also believed that the ductus thoracicus dis- 
charged into the subclavian veins. Meyer thought that the anterior 
bifurcation was an artificial production due to Panizza's use of 
mercury, and that the ductus thoracicus actually ended blindly in 
the region of the heart. This view is, of course, equally wrong. 

Entering the subvertebral trunks dorsally are a number of seg- 
mental vessels from the dorsal muscles. Hoyer and Udziela have 
shown that, in the larva, these vessels anastomose with the T. lymph, 
long, dorsalis and send branches to the lateral lymph hearts, while 
in the anterior or 'thoracic' region they also fuse with one another 
to form a longitudinal vessel running close beside the vertebral 
column on either side. They call this vessel the truncus lymph, long, 
paravertehralis. In the adult it is not possible to trace the connexions 
with the dorsal vessel, which itself tends to disappear and is distorted 
by the development of the cutaneous glands, but the connexion with 
the lymph hearts exists, and it is possible to find some traces of the 
paravertebral vessel, although it shows signs of degenerating. 

E. Lymphatics oj the Pectoral Region (PI. XVIII, fig. 68). 

These are very extensive and paired in the adult, but the actual 
arrangement does not differ markedly from that of the larva. The 
largest and most important sinus in the pectoral region is the sub- 
scapular sinus (plexus axillares, Panizza). In the adult this sinus 



THE VASCULAR SYSTEM 257 

becomes divided into two more or less distinct sections, namely, a 
pars dorsalis lying between the nerves of the brachial plexus and the 
scapula and forming the subscapular sinus proper, and a pars ven- 
tralis which, although in frequent communication with the former 
sinus, is separated from it by the nerves of the brachial plexus and 
by the subclavian artery. Its anterior extremity lies ventral to the 
aortic arches and is separated by them from the peri-pharyngeal trunk. 

The sinus subscapularis^ pars dorsalis (s.ly.sc") receives the T. 
lymph, long, lateralis, a portion of the T. lymph, long, parepigas- 
tricus, the post-axial trunk from the fore-limb and the transverse 
trunk from the subvertebral stem already mentioned. Its lymph 
flows into the pars ventralis by two main tracks, one, a direct path, 
just anterior to the third spinal nerve, and the other by a circumflex 
route, the T, lymph, circumflexus scapulae (t.ly.circ.sc), around the 
lateral aspect of the scapula, between the bone and the MM. dorsalis 
humeralis and dorsalis scapulae. This trunk receives the pre-axial 
trunk from the limb just before joining the pars ventralis, and also 
sends a branch directly forwards to enter the jugular sinus. 

The pars ventralis (s.ly.sc.') is a large sac-like structure closely 
surrounding the roots of the main veins. In addition to the con- 
nexions with the pars dorsalis, already mentioned, the pars ventralis 
receives the remainder of the T. lymph, long, parepigastricus, i.e. 
the portion entwining the anterior epigastric veins (t.ly.l.pe.) and also 
2. sinus anastomoticus (s.ly.anas.) from the pars ventralis of the opposite 
side. This sinus passes round through the fold between the peri- 
cardium and the peritoneum, ventral to the oesophagus, and some- 
what dorsal to the posterior end of the ventricle. Entering the pars 
ventralis anteriorly are the sinus thyroideus (s.ly.th.), an elongated sac 
lying lateral to the thyroid gland, and a communication with the 
subcutaneous sinuses covering the M. interhyoideus posterior. The 
latter connexion penetrates the muscle and passes a short distance 
laterally over the dorsal side thereof before entering the pars ventralis. 

The pars ventralis, in turn, discharges into the truncus peri- 
pharyngeus by means of several vessels that pass in mesialwards 
between the aortic arches. 

Panizza supposed that the contents of this sinus flowed into the 
subclavian vein, which is undoubtedly incorrect, although, in the larva, 
Hoyer and Udziela show several fine connexions between it and the 
ductus Cuvieri and the anterior cardinal. The appearance in the adult, 
however, renders it extremely doubtful whether these communica- 
tions persist. There are a number of fine lymph capillaries adhering 
very firmly to the bases of the internal jugular and subclavian veins, 

4038 c 



2 58 THE SALAMANDER 

but they may be scraped off without leaving any apparent trace, and 
while there may be some entry of lymph at these points it is very un- 
likely in view of the free passage for this fluid into the veins by means 
of the central lymph heart in the immediate vicinity. 

F. Lymphatics oj the Head (see also PL XIX, fig. 70). 

These may be conveniently distinguished as 'dorsal' or 'ventral* 
according to their position relative to the mouth. 

The dorsal lymphatics oj the head (Fig. 70) are almost entirely con- 
fined to a pair of Y-shaped sinuses beneath the eyes — the peri-orbital 
sinuses (s.ly.p-o.). They were first described by Tretjakoff (1930). 
Each sinus lies in the posterior half of the orbit, between the M. 
levator bulbi and the skull, so that the bulbus oculi rests between the 
limbs of the 'Y', while the stem passes across the palato-quadrate and 
enters the peri-pharyngeal trunk. A small vessel from the peri- 
orbital sinus passes through the antrum petrosum laterale and enters 
the jugular sinus. 

As Tretjakoff points out, in Urodeles lymph has replaced blood 
as a 'cushion' for the eye, enabling it to turn freely, since in fish that 
function is performed by the peri-orbital blood sinus. 

Lymph capillaries are rich on the roof of the mouth and pharynx, 
and they also ultimately join the truncus peri-pharyngeus. 

The ventral lymphatics 0} the head(¥\g. 68) are much more numer- 
ous and extensive than the dorsal sinuses. On removing the skin 
there may be seen, in addition to the subcutaneous network already 
described, a lymphatic sinus running around the lower jaw — the 
mandibular sinus (s.ly.man.). At the posterior margin of the M. inter- 
mandibularis this sinus unites with the submaxillary network to 
form a common stem that passes dorsalwards between the above- 
named muscle and the M. interhyoideus and enters the sinus mandi- 
bularis profundus. 

The sinus mandibularis profundus (s.ly.m.pf.) is a short sinus 
situated mesial to the angle of the lower jaw. 

If the submaxillary network be removed, and the MM. inter- 
hyoideus and intermandibularis cut along the linea alba and turned 
back, the deeper sinuses may be traced. They probably arise from 
the tongue as the central organ, although the whole floor of the 
mouth and pharynx is rich in lymph capillaries. The lymphatics 
of the tongue drain into a saccus copularis (sac. cop.), an irregular 
sac lying on the dorsal side of the copula, and the lymph flows there- 
from by several alternative routes. In injected specimens the saccus 
copularis may be seen extending anteriorly from the copula as a small 



THE VASCULAR SYSTEM 259 

bladder from which pass two pairs of small vessels. Each pair 
accompanies the hypoglossal nerve of one side and enters the sinus 
thyroideus of that side. Between the radials on either side of the 
copula further bladder-like extensions of the saccus occur, which are 
drained by a pair of sinuses lying dorsal to the cerato-hyals. From 
their position they may be named the hyoidean sinuses (s.ly.hy.). 
Each joins the sinus mandibularis profundus of its side and enters 
the jugular sinus. On the dorsal side of the branchial cartilages on 
either side, and filling the space between them with the exception of 
that occupied by the M. rectus cervicis, is the branchial sinus (s. ly.br.). 
This also arises from the saccus copularis, but flows into the T. peri- 
pharyngeus. Its course between the posterior end of the branchial 
cartilages and the peri-pharyngeal trunk is sinuous and follows the 
lingual vein. 

Tht jugular sinus (s.ly.j.) lies directly behind the corner of the 
mouth. Into it drain (i) the combined SS. mandib. prof, and hyoi- 
deus, (ii) the direct continuation of the S. circum. scap., and (iii) 
numerous small vessels from the occipital region, the thymus gland, 
and the antrum petrosum laterale. It discharges directly into the peri- 
pharyngeal trunk. 

It must be noted here that the large space into which the posterior 
cornu of the hyoid projects is labelled by Driiner 'Sinus lymphaticus'. 
It is, however, not a true lymph sinus. Of the several injections that 
have been made in the course of the present investigation, in none 
has the injection mass used entered this space, neither has it been 
possible to find any opening therefrom. It therefore seems to be a 
closed sac of connective tissue whose sole function is to allow the free 
extension of the hyobranchial cartilages. 

Extending posteriorly from the copula in the middle line is a median 
sinus connecting the saccus copularis with the Truncus peri-pharyn- 
geus. It also receives many tributaries from the floor of the mouth. 

G. Lymph hearts and their Relation to the Blood-vessels (PI. XVII). 

As previously mentioned there is a series of contractile vesicles along 
each side of the body in the sulcus lateralis, which place the lymphatic 
system in communication with the V. lateralis. There are normally 
fifteen pairs of these hearts. Four pairs are post-sacral, i.e. caudal, and 
the remaining eleven pairs lie between the sacrum and the posterior 
border of the scapula. There is some variation in size — those lying 
in the caudal and sacral regions being the largest, while those in the 
middle of the trunk are the smallest. Each heart lies embedded in 
the muscles directly behind the myoseptum, at the lateral extremity 



26o THE SALAMANDER 

of a rib. It is usually necessary to dissect away some muscle-fibres 
in order to expose them fully. Particularly is this so in the trunk 
region, but behind the sacrum they lie more superficially. 

Lymph is received by each heart from two sources — first, from the 
T. lymph, long, lateralis, i.e. from the lateral subcutaneous network, 
and secondly from the parietal branches of the subvertebral trunks. 
The openings both into and from the hearts are guarded by valves. 
Figs. 65, 66^ and 67, taken from Hoyer and Udziela's paper, show 
these relations clearly. The chief points in which the adult differs 
from the larva are, first that the hearts are more deeply embedded in 
the muscles, second that the T. lymph, long, lateralis pursues a more 
zigzag course, and is not so clearly defined, and third that there tends 
to be formed between each heart and the V. lateralis a short efferent 
stem, so that the heart is further removed from the vein. 

Zacwilichowski (19 17) has studied the development of these 
lateral hearts in Molge vulgaris (presumably Triton taeniatus\ and 
found that they develop from the wall of the vein, and later become 
secondarily associated with the lymphatic system. 

There remains yet the central lymph heart (ct.ly.h.) to be con- 
sidered. Its location in the truncus arteriosus has already been 
described (p. 1 90). To observe it the bulbus cordis must be severed 
near the ventricle and turned over anteriorly — then, if the muscles 
on the dorsal side of the truncus arteriosus be carefully removed, 
the lymph heart is revealed. The lymph enters from the T. peri- 
pharyngeus between the carotid and systemic arches and is dis- 
charged, posterior to the pulmonary arch, into the proximal portion 
of the lingual vein (V. pharyngea). Regurgitation at both entry and 
exit is prevented by valves. Greil's figures are a little misleading in 
that he shows the afferent vessels pointing directly forwards. Accord- 
ing to Hoyer and Udziela they form a more oblique angle even in 
the larva, while in the adult their direction is almost transverse. The 
shape of the central lymph heart is roughly pyramidal, the base being 
directed forwards, while the apex is continued posteriorly as a fine 
vessel draining lymph from the bulbus cordis. The central lymph 
heart really lies dorsal to the truncus arteriosus, but, during develop- 
ment, it becomes enclosed by a strap-like band of cardiac muscle 
from the bulbus cordis that grows completely round between the 
right and left carotid arches and forms a continuous strip connecting 
the dorsal and ventral sides of the truncus arteriosus. 

It is uncertain whether there is an intrinsic rhythm in the heart 
itself or whether the contraction is brought about entirely by the 
extrinsic muscles of its neighbourhood. 



THE VASCULAR SYSTEM 261 

BIBLIOGRAPHY OF SECTION VI 

26, 28, 29, 39, 40, 49, 59, 60, 61, 86, 97, 108, 130, 131, 179, 198, 237, 239, 
272, 277, 278, 279, 280, 284, 288, 314, 315, 316, 317, 318, 319, 328, 331, 
337^ 340, 341, 343. 353. 354. 389. 39i, 4o8, 410, 418, 444, 452, 474, 475, 
476, 477, 478, 480, 502, 503, 504, 508, 520, 528, 529, 530, 531, 532, 542, 
552, 553, 563, 596, 597, 617, 625, 626, 638, 639, 640, 642, 643, 644, 671, 
672, 701, 736, 747, 751, 756, 763, 770, 771, 774, 775, 830, 836, 839. 



VII 

THE ALIMENTARY TRACT AND ITS 

APPENDAGES, TOGETHER WITH THE SPLEEN 

AND THE MESENTERIES 

THE ALIMENTARY TRACT (PL XXII) 

1. General. 

The alimentary tract and the associated structures of the Salamander 
show a general similarity to those of the Frog, but there are several 
noteworthy differences. Of these the following may be mentioned : 
(i) the presence of teeth on ^o//^ jaws, and the great backward exten- 
sion of the pre-vomerine teeth ; (ii) the relatively immobile tongue ; 
(iii) the absence of Eustachian tubes opening into the pharynx; 
(iv) the fairly simple undivided liver ; (v) the size and shape of the 
spleen. 

2. The Mouth Cavity. 

Owing to the fact that the outer skin is reflexed some distance 
dorsalwards at the angles of the mouth over the coronoid process 
of the lower jaw, there is a difference between the actual gape and 
the apparent gape. Thus the former extends to about the level of the 
posterior boundary of the eye, but the latter is continued a short dis- 
tance behind this. There is only a slight suspicion of free 'lips', but 
the outer skin covering the jaws is glandular and contains the so- 
called labial glands. 

The teeth are pleurodont and are directed inwards. They do not 
terminate in a single point but are bifid at the tips and sometimes 
tricuspid. There are slight differences in shape between the teeth in 
the various regions of the mouth, but for practical purposes the 
dentition may be termed homodont. The teeth are small and occur 
on both jaws as well as on the pre-vomers. The dentigerous pro- 
cesses of the latter extend far caudalwards to the back of the mouth, 
while their S-like shape is characteristic of true Salamanders, and, 
coupled with the short thick tongue, are definitive of the genus. 

A detailed study of the teeth of Amphibia was undertaken by 
Hertwig (i 874). He distinguishes two sections of each tooth — the 
crown which is visible above the gum, and the socket embedded in the 



THE ALIMENTARY TRACT, SPLEEN, AND MESENTERIES 263 
gum. The junction between these two portions is clearly marked. 
Three tissues combine to form each tooth, namely, dentine, enamel, 
and cement. These features may all be clearly seen by pulling out 
a tooth — which is quite easy — and staining it in a solution of 
alizarin in absolute alcohol and mounting it in balsam on a 
slide. 

The mandibular and maxillary teeth serve merely to retain the 
food in the mouth, but the pre-vomerine teeth on the palate assist 
the tongue to work the food back to the pharynx so that it may be 
swallowed. 

The tongue of the Salamander is a much less highly developed 
organ than that of the Frog, and consists simply of an oval pad firmly 
affixed to the floor of the mouth and with it to the copula of the 
hyobranchial apparatus. It is free laterally and to some extent pos- 
teriorly also, but nothing like to the same extent as the Frog's tongue. 
As Kallius (1901) has shown, the adult structure consists of a secon- 
dary tongue superimposed on a larval ■primary tongue. The secondary 
tongue forms practically the whole organ in the adult, and its epithe- 
lium, which is highly glandular, is thrown into deep folds arranged 
in a somewhat radial manner, so that the superficial area of the mucous 
epithelium is enormous. The primary tongue is just visible under the 
posterior edge of the secondary tongue. It is non-glandular. There 
is only a slight tendency for the secondary tongue to be bifid 
posteriorly. The ontogeny of the tongue as described by Kallius 
(1901) shows that the tongue of Salamandra exhibits a close 
recapitulation of the changes which Gegenbaur (1894) suggested 
had taken place during the evolution of this organ in the vertebrate 
series. 

The mechanism whereby the tongue is extruded from the mouth 
in the act of seizing food is described on p. 61. 

The epithelium of the mouth and pharynx is glandular and highly 
vascular, and is of considerable importance in respiration (see also 
p. 274, etseq.). In addition to this general glandulartissuethereare two 
specialized sets of glands opening on to the roof of the mouth. These 
have been described in detail quite recently by Seifert (1932). The 
first and most important has long been known. It is the glandula 
intermaxillaris (or internasalis)^ and is situated between the nasal 
capsules. It discharges by two or three openings into a small median 
depression lying on the roof of the mouth immediately beneath the 
gland, and anterior to the pre-vomerine teeth. The other set of 
specialized glands are the 'Gaumendriise' (Seifert) or glandula pala- 
tina, which open into a series of grooves on the bases of the 



264 THE SALAMANDER 

prevomerine teeth. These grooves and the median pit are easily- 
discernible with a lens, but the actual openings of the glands must 
be sought in transverse sections under a microscope. 

Kingsbury (i 9 1 2) in Salamandra atra and various other Amphibia 
finds, in as many as twelve regions of the mouth and pharynx, 
aggregations of cells of a lymphoid character which he regards as 
analogous to the tonsils of Amniota. A detailed discussion of this 
question lies outside the scope of the present work, but the position 
of the following four patches, which Kingsbury finds to be definite 
and constant, may be mentioned. 

(i) The choanal in the roof of the mouth immediately posterior to 
the internal narial opening. 

(ii) The lateral^ situated laterally in the pharynx in the general 
region of the articulation of the jaw. 

(iii) The sublingual lying lateral to the tongue over the expanded 
cerato-hyal. 

(iv) The preglottideal on the floor of the mouth just in front of the 
glottis. 

It must be pointed out that Kingsbury's results are of a general 
nature and have not been checked specifically for Salamandra 
maculosa. 

As in Amphibia generally the eyeballs are not separated from the 
palate by any skeletal element, and their form is therefore plainly 
visible as a pair of convex bulges on the roof of the mouth. Just in 
front of these may be seen a pair of relatively large oval openings, 
the choanae or internal nares. 

3. The Pharynx and Oesophagus. 

The pharynx leads directly from the mouth as a wide tube. It is 
richly vascular and unmarked by any Eustachian tubes and passes 
imperceptibly into a wide oesophagus (oes.) which in turn leads, also 
without any sharp line of demarcation, into the stomach. The external 
walls of both oesophagus and stomach, and in fact of the whole gut 
also, are smooth, but the internal walls are variously corrugated, 
each section of the gut having its own particular pattern. Thus the 
distinction between the oesophagus and stomach is clearer internally 
than externally, the lining of the former being thrown into deep 
longitudinal folds, while the folds of the stomach are marked by a 
relatively shallow, wide-meshed, irregular network, of which the 
longitudinal members are the more prominent. The mucous epithe- 
lium is velvety, and when scraped off the blood capillaries are found 
to be fiat and not raised into loops. 



THE ALIMENTARY TRACT, SPLEEN, AND MESENTERIES 265 

Jacobshagen (19 14) has made a comparative study of the relief 
pattern of the gut lining in Amphibia. He denies the presence of 
a pyloric valve in Salamandra — a conclusion which may be justly 
criticized. 

4. The Stomach. 

The stomach (stom.) is conical and lies almost longitudinally along 
the left side of the body. It narrows considerably at its posterior end, 
and there is always a well-defined constriction between this end of 
the stomach and the duodenum. The constriction is specially notice- 
able when the gut is full, and a slight thickening of the wall is dis- 
cernible in longitudinal section, so that it seems impossible to avoid 
the conclusion that this constriction is indeed the pylorus (py.)> 
although it must be admitted that it is less perfectly formed than in 
many animals, and it is doubtful whether the lumen is ever completely 
closed. 

5. The Duodenum. 

The duodenum (duo.) is short and is sharply reflexed along the 
mesial aspect of the stomach, but follows a more transverse direction 
relative to the body than does that organ. It is indistinguishable 
from the rest of the intestine except that it receives the pancreatic 
and bile ducts. 

The dorsal pancreatic duct (d.p.d.), or rather the duct of the dorsal 
pancreas, enters the proximal end of the duodenum some 2 mm. or 
so from the pylorus. The ventral pancreatic ducts (d.p.v.), which 
belong to the two ventral pancreata, discharge into the common bile- 
duct which, in turn, enters the distal end of the duodenum, some 
4—5 mm. from the dorsal pancreatic duct (see also p. 267). 

6. The Intestine. 

The intestine (int.) is a coiled tube of a regular diameter, approxi- 
mately equal to, or slightly smaller than, that of the duodenum. Its 
length is about one-half that of the whole gut, measured from the 
pharynx to the cloaca. The pattern of the internal relief of the duo- 
denum and intestine, considered as a whole, consists of a series of 
sinuous longitudinal ridges. The ridges are thick proximally — that 
is, at the duodenal end — but tend to become thinner and straighter 
towards the hinder end of the gut, until they are almost knife-like. 
If the mucous epithelium is scraped or brushed off it is seen that 
the underlying vascular network is also raised into ridges, and that 



266 THE SALAMANDER 

the capillary loops follow the same wavy outline. The network is 
much richer at the anterior end of the gut. 

7. The Rectum. 

The rectum (rect.), like that of the Frog, is an expanded flask- 
shaped structure arising quite suddenly from the posterior end of 
the intestine. It is much thinner walled than the rest of the gut. 
Crofts (1925) has described 'a small lateral "cul-de-sac" lying to the 
left side' of the rectum and containing lymphoid tissue, which she 
considers to be the homologue of the rectal gland of Elasmobranchs. 
It is supplied by the haemorrhoidal artery. It is often difficult to 
determine the exact location of this gland by direct observation, but 
is more easily seen when the rectum is full than when it is empty, as 
then it shows up against the dark faecal matter. The internal surface of 
the rectum is also traversed by longitudinal ridges, set fairly close 
together and thicker than in the intestine, but not so wide as in the 
stomach. It differs from the latter also in that the vascular network 
extends into the ridges. The rectum and urino-genital organs dis- 
charge their contents into a common cloacal chamber, but since 
the cloaca has a closer functional association with the latter organs it 
is described together with them (see p. 286). 



THE GLANDS ASSOCIATED WITH THE 
ALIMENTARY TRACT 

The glands associated with the gut, in addition to those already 
mentioned, are the liver^ the pancreas, and the spleen. The association 
of the last-named with the stomach is topographical only, since it has 
neither functional nor developmental connexion with the gut. 

I. The Liver. 

The liver (liv.) is large, and only very slightly lobed. It lies ventral 
and to the right of the stomach, and, in the fresh condition, is dark 
red in colour. The lejt side of the gland is prolonged posteriorly into 
a thin leaf-like lobe. The backward extension of the right side is not 
so well defined nor so thin, but from this side of the liver there is a 
pronounced dorsal extension which encircles the post-caval vein, and 
which may hence be termed the post-caval lobe. It extends dorsally 
right to the point where the post-caval separates from the post- 
cardinal veins. The liver is completely suspended by mesenteries 



THE ALIMENTARY TRACT, SPLEEN, AND MESENTERIES 267 
(q.v.), and has a slight median cleft anteriorly where the ventral, or 
sub-hepatic, mesentery joins the lig. hepato-entericum. 

There is a relatively large gall-bladder (gl.b.) lying just dorsal to 
the right lobe of the liver. In order to distinguish the details of the 
bile-ducts it is necessary to dissect away some portion of the liver 
tissue since they are completely embedded therein, with the excep- 
tion of the common bile-duct itself which is embedded in the 
ventral pancreas. 

Three main bile-ducts emerge from the liver (see Fig. 76). On 
the extreme right — left as viewed — is the hepato-cystic duct (d. h-c.) 
which bears the gall-bladder as a lateral expansion of its walls, and 
afterwards becomes the cystic duct (d.cy.), which is joined almost 
immediately by the Jirst hepatic duct (d.h.'). The latter is separated 
from the former by the ventral hepatic artery. The second hepatic duct 
(d.h.") enters farther down, within the pancreas, so as to produce 
a very short common bile-duct (d.c.) into which the ducts of the ventral 
pancreas also discharge. 

2. The Pancreas. 

^he pancreas (pan.) is roughly triangular in shape, and lies rather 
to the dorsal side of the duodenum, between it and the stomach. As 
Goppert (1891) and Choronshitsky (1900) have shown, the adult 
pancreas is a composite gland, representing three outgrowths from 
the embryonic gut, namely, two ventral and one dorsal. As already 
stated, the two ventral pancreata discharge their secretion into the 
common bile-duct, while the dorsal pancreas discharges by a separate 
duct directly into the duodenum. The pancreatic ducts are by no 
means easy to see by direct observation, but may be readily demon- 
strated by cutting a series of microtome sections of the duodenum. 
The three lobes of the embryonic pancreas become indistinguishably 
fused in the adult into a single mass. 

THE SPLEEN 

The spleen (sp.) is not strictly an appendage of the gut, but, since 
it is suspended from the left dorso-lateral wall of the stomach by the 
lig. gastro-lienale, and in part shares its blood-supply, it may con- 
veniently be considered here. It is a comparatively thin, elongated 
ovoid body, some 10-15 mm. long by 4-4^ mm. broad. It is deep 
red in colour in the fresh condition. Choronshitsky holds that it is 
purely mesodermal in origin, while functionally it is of course a 
blood gland. 



268 THE SALAMANDER 

THE MESENTERIES 

The mesenteries of the Salamander exhibit many interesting 
features. They have been investigated by Klaatsch (i 893), Mathes 
(1896), and Brachet (1896). These authors are at some variance — 
in fact each seeks to correct his immediate predecessor, but the causes 
of their dissension lie in the mode of origin of the mesenteries 
rather than in their final relationships, so that it is unnecessary to 
enter into the long and rather involved discussion here. The following 
description therefore applies simply to the relations as they may be 
observed by the dissection of the adult. 

1. The Dorsal Mesentery and its Derivatives (see also Pis. XX, 
XXI, and XXIII). 

The dorsal mesentery (l.d.) attaches the gut to the dorsal body-wall. 
It is not quite a continuous sheet of membrane but is interrupted at 
the pyloric end of the stomach by a large circular hole, the posterior 
rim of which is bounded by the pancreas. Anteriorly the mesentery 
extends as far as the base of the lungs, i.e. approximately to the level 
of the anterior end of the ventricle of the heart. 

The gonads are suspended by a pair of lateral diverticula from the 
dorsal mesentery which form the mesovaria (l.ms.) in the female and 
the mesorchia (l.mch.) in the male. 

The kidneys are not suspended by a definite mesentery but lie fairly 
close to the body-wall ; there is, however, a mesentery attaching the 
oviduct in the female, and the Wolffian duct in the male, to the outer 
edges of the kidneys. 

The spleen is suspended from the left side of the stomach also by 
a reduplication of the dorsal mesentery. This fold is continued from 
both ends of the spleen as a 'tuck' in the mesentery. The anterior 
prolongation reaches to the stomach-wall and is the plica gastro- 
lienalis (Klaatsch), while the posterior extension is much longer and 
reaches to the rectum, and is the plica recto-Uenalis (Klaatsch). It is 
important to notice that this latter fold does not follow the convolu- 
tions of the intestine, but stretches straight across the comparatively 
short distance between the posterior end of the spleen and the anterior 
end of the rectum, so that the whole of the convoluted part of the 
mesentery lies ventral to it. 

2. The Ligamentum Hepato-entericum (l.h-e.). 

Between the ventral side of the stomach and the dorsal surface of 
the liver there is a mesentery which carries the factors of the anterior 



THE ALIMENTARY TRACT, SPLEEN, AND MESENTERIES 269 

gastric vein and the branches of the dorsal hepatic artery. It is the 
lig. hepato-entericum. Its posterior margin ends freely at about 
the level of the bile-duct, while anteriorly it becomes attached to the 
post-caval vein, the pericardium, and the posterior wall of the 
trachea, where it ends. 

3. The Ventral Mesentery. 

The Ventral Mesentery^ or lig. suspensorium hepatis (Rabl), as its 
name implies attaches the liver to the ventral body-wall. It continues 
posteriorly along the abdominal vein as far as the pelvic girdle, while 
anteriorly it ends at the pericardium. 

4. The Ligamentum Hepato-cavo-pulmonale. 

The lungs are not symmetrically related to the mesenteries. On the 
left the lung is supported only from the dorsal side by a reduplication 
of the mesentery supporting the stomach — the pleural ligament 
(l.pl.) — the posterior third of the lung (in the collapsed condition) 
being entirely free. The right lung on the other hand, in addition to 
being so supported, is attached to the liver by the lig. hepato-cavo- 
pulmonale (l.h.c.p.'). This mesentery attaches the ventral surface of 
the lung to the lateral edge of the liver, and, as already stated, occurs 
on the right side only. 

There yet remain to be mentioned a pair of mesenteric folds of 
considerable interest and importance. They are the lig. coronaria 
hepatis (l.c.h.) (Rabl) and extend from the abdominal openings of 
Miiller's ducts — the ostia tubae — to the liver. The free edge of each 
fold forms the ventral lip of the funnel leading into the ostium, the 
dorsal lip of the funnel being formed by the lig. tubae (l.t.) (Rabl), 
which is a smaller fold joining the larger towards the liver. A similar 
ligament is also present in the male, although of course it is much 
smaller, and Rabl (1904), who has studied the development of this 
structure, states that, in the male, the normally single ligament may 
represent either the lig. tubae or lig. coronarium hepatis, the other 
having atrophied. The same author suggests that the lig. coronarium 
hepatis is homologous with the pleuro-peritoneal membrane of the 
mammalian embryo, which had been previously regarded as the an- 
lage of the diaphragm (vide infra). 

The peritoneum is a thin, loose membrane lining the abdominal 
cavity. It is only very loosely attached to the body-wall so that it 
is easily stripped therefrom. It is pigmented, but not heavily, the 
chromatophores being arachnoid. 



270 



THE SALAMANDER 



5. The Diaphragm. 

Although there is no thorax in Urodela and the lungs retain their 
primitive position in the abdomen, nevertheless the coelomic cavity 
is bounded anteriorly by precisely the same structures as it is in the 
Mammalia. The comparison between the Amphibia and Mammalia, 
and also with Birds and Reptiles, has been discussed in detail by 
Keith ( 1 905), who regards the pleural coelom of Mammals as having 
been cut off from the abdominal cavity by the evagination of the 
lungs through the diaphragm, the points at which this occurred 
being marked by the pleuro-peritoneal openings of the embryo. 
Keith agrees with Rabl in regarding the pleuro-peritoneal membrane 
as 'merely a cresentic fold of peritoneum forming part of the 
genital mesentery'. 

The three structures entering into the composition of the dia- 
phragm in Salamandra are : (i) ventrally, a portion of the superficial 
stratum of the M. rectus abdominis which is closely attached to the 
ventral wall of the pericardium, while a few fibres are actually in- 
serted into it; (ii) a central portion formed by the septum trans- 
versum^ i.e. by the membrane dividing the pericardial coelom from 
the main body cavity; and (iii) a dorsal part formed by those fibres 
of the transversalis muscle-sheet which are inserted into the dorsal 
pericardial wall and around the tracheal chamber (cf. Fig. 77; m.t.). 

Regarding the function of the diaphragm, Keith declares that it 
is primarily concerned with assisting the venous circulation of the 
abdominal cavity and filling the heart with blood by compressing 
the viscera, and that its respiratory functions are secondary. The 
latter function is expiratory in character in Salamandra, but becomes 
inspiratory in Mammalia, where the lungs are anterior to the dia- 
phragm. 

BIBLIOGRAPHY OF SECTION VII 

435 44, M, 1% 108, 121, 189, 218, 234, 255, 298, 309, 315, 350, 356, 368, 
375, 3795 390j 418, 43O5 44^, 484, 505, 507, 51O5 606, 615, 627, 654, 694, 
706, 786. 



VIII 

THE RESPIRATORY SYSTEM, INCLUDING 
THE LARYNX 

THE RESPIRATORY SYSTEM (PI. XXIII, %. 77) 

Under this head it is proposed to consider those structures usually- 
regarded as directly associated with the physiological process of 
respiration, namely the glottis, the laryngo-tracheal chamber, the 
bronchi and the lungs, and finally, to review the physiological aspect 
and to discuss the probable means by which re-oxygenation of the 
blood is brought about in the adult Salamander. 

As is well known, the Caducibranchiate Urodeles employ the 
'buccal-force-pump' method of breathing similar to that of the 
familiar Frog. This involves the presence of mechanisms for closing 
the nares, and for raising and depressing the floor of the mouth, 
but, as such structures are only secondarily subservient to respira- 
tion, they are described elsewhere in their appropriate places (see 
pp. S5 and 303). 

1. The Glottis. 

The glottis (gts.) is a small slit-like aperture lying longitudinally 
on the floor of the pharynx. It is raised only very slightly above the 
level of the surrounding tissues, and leads directly into the laryngo- 
tracheal chamber. 

2. The Laryngo-tracheal chamber. 

The laryngo-tracheal chamber (tr.ch.) is a more or less triangular 
cavity. The glottis opens into the apex, antero-dorsally, while the 
posterior basal angles merge into the bronchi. The whole structure 
lies immediately dorsal to the truncus arteriosus, which thus forms 
a useful landmark for dissection purposes. The sides of the chamber 
are supported by a series of semicircular cartilages, the whole series 
forming the cartilage lateralis (Edgeworth),^ while the anterior pair 
are distinguished as the pars laryngea s. arytenoid (Edgeworth), and 
the remainder as the pars trachealis (Edgeworth). The arytenoid 
cartilages (c.ary.) bound the sides of the glottis and are approximated 

' The synonyms of other authors are sufficiently obvious to render citation un- 
necessary. 



272 THE SALAMANDER 

by the M. constrictor laryngis and pulled apart by the M. dilatator 

laryngis. These muscles are described on p. 64. 

3. The Bronchi. 

The bronchi are short, wide tubes leading directly from the pos- 
terior angles of the laryngo-tracheal chamber to the lungs. Their 
epithelium is ciliated (Suchard). 

4. The Lungs. 

A very good account of the lungs has been given by Suchard ( 1 903), 
who deals with their minute, as well as their gross structure. They 
consist of a pair of small, heavily pigmented, conical sacs, pointed 
posteriorly, and presenting a sacculated appearance externally. Their 
cavities are not simple but are interrupted by trabeculae. According 
to Suchard, two main longitudinal compartments may be recognized, 
containing respectively the artery and the vein, together with sub- 
sidiary longitudinal compartments formed around the arterioles and 
venules, while the transverse septa are merely folds in the pulmonary 
membrane. The same author states that smooth muscle elements 
are present, but apart from these the lungs would be satisfactorily 
compressed by the muscles of the abdomen. The lungs are not sus- 
pended freely in the pleuro-peritoneal cavity as are those of the Frog, 
but are firmly anchored dorsally to the body-wall by the pleural 
mesenteries, and ventrally to the liver by the lig. hepato-cavo-pul- 
monale (see also p. 269). 

Homologies of the Urodele Larynx. 

A great deal of work has been published on this question and 
various authors have advanced different theories in an attempt to 
demonstrate that the arytenoid cartilages and the laryngeal muscles 
represent the remains of a transformed visceral arch. 

The discussion began in 1892, when Wilder and Gegenbaur 
independently advanced similar theories. Wilder suggested that the 
■pars laryngea cartilaginis lateralis really represented the cartilage 
of the fifth branchial arch of Selachians, and that the M. dilatator 
laryngis was homologous with the dorsal segment of the muscles of 
that arch. He also thought that the ring of muscle surrounding the 
larynx — M. constrictor laryngis — was a continuation of the circular 
muscle of the alimentary canal, and that X^sx&'pars trachealis cartilaginis 
lateralis arose later than, and independently of, the arytenoids from 



THE RESPIRATORY SYSTEM, INCLUDING THE LARYNX 273 

the surrounding tissue. In other words he claimed that the cartilago 
lateralis had a dual origin. 

Gegenbaur's theory was very similar except that he thought that 
the whole cart, lateralis was derived from the fifth branchial arch, and 
that the M.. transversus ventralis was homologous with the constrictores 
arcuum of that arch. Goppert ( 1 8 94) accepted Gegenbaur's theory and 
said that the M. dilatator laryngis represented M. levator arcuum F, 
and was therefore the homologue of the levatores of the branchial 
arches. He opposed Wilder's theory as to the derivation of M. con- 
strictor laryngis on the ground that it is a striped muscle, while the 
circular muscle of the alimentary canal is of course composed of 
smooth elements. 

In 1896 Wilder returned to the subject. He renounced his former 
theories concerning the dual origin of the cart, lateralis and deriva- 
tion of the ring of muscle round the larynx from the muscles of the gut, 
and accepted the views of Gegenbaur and Goppert. Goppert (1898) 
also abandoned his former theory that the M. laryngeus ventralis is 
derived from the transverse muscle of the fourth branchial arch and 
said that it was really a serial homologue of this and represented 
the M. transversus ventralis V. 

In 1 90 1 Druner published his classic work on the visceral skeleton 
and its muscles in Salamandra and Triton in both larval and adult 
stages. He declared a general acceptance of the current theories but, 
on the ground of innervation, suggested that the laryngeal skeleton 
and its muscles were not derived from the fifth branchial segment but 
from the sixth or some more posterior arch. 

Wiedersheim (1904) was the first to put forward the suggestion 
that perhaps the cart, lateralis of Amphibia had no connexion with 
the visceral arches after all, but was rather a supporting structure 
produced independently in response to muscular pressure. This 
suggestion was based on his study of the Ganoid and Dipnoan 
fishes [Zool. Jahrb. Suppl. vii, pp. 1—66). 

Edgeworth (1920) turned Wiedersheim's suggestion into an 
established fact. After giving an anatomical description of the de- 
velopment of the laryngeal structures in various Amphibia, he 
reviews the work of previous investigators and then summarizes 
his own conclusions in the following words : 

'The arytenoid s. pars laryngea cartilaginis lateralis of Amphibia is developed 
within the constrictor laryngis, or within this and the laryngei, and like them is 
differentiated from cells from the splanchnic layer of the coelomic epithelium. 
It does not therefore represent a fifth, or a more posterior branchial bar. Its 
development is probably, as Wiedersheim suggested, dependent on muscular 
4038 T 



274 THE SALAMANDER 

action. It is possibly related to the new development of the dilatator laryngis 

in Amphibia. The tracheal skeleton s. pars trachealis cartilaginis lateralis is 

a backward prolongation of the arytenoid and related to the development of the 

trachea. 

It must be noticed that an excellent description of both the 
muscles and the skeleton of the larynx of Salamandra was given as 
early as 1 839 by Henle, but he made no suggestions as to the homo- 
logies of the several parts. 

Physiological Considerations. 

It is well known that the common Frog is capable of respiring to 
a large extent through the skin as well as through the lungs. That 
Caducibranchiate Urodeles are also not entirely dependent on pul- 
monary respiration is obvious from the fact that many are devoid of 
lungs altogether, e.g. Eurycea (Spelerpes), Salamandrina, &c., while 
the lungs of other forms, e.g. Triton, are mere hollow sacs with 
smooth walls which present only a relatively small surface to the 
contained air, and are probably hydrostatic rather than respiratory 
in function. In support of this latter statement Lonnberg (1899) 
points out that all aquatic Urodeles devoid of lungs craw/ r^ither than 
swim. Numerous experiments have been carried out, by Camerano 
and others, with the object of discovering to what extent the skin 
acts as a respiratory medium. These have been well summarized by 
Bethge (1898), and it is not proposed to discuss them here. They 
are none of them free from the objection that they do not ensure the 
absence of secondary effects being introduced which may affect the 
general health of the animals being treated. Bethge himself draws 
attention to the interesting fact that the cutaneous capillaries of 
Salamandra — where the lungs have an undoubted respiratory func- 
tion — have only about one-half the diameter of those of Eurycea 
(Spelerpes), an animal entirely devoid of lungs, while in Triton — 
whose lungs are largely hydrostatic in function — the capillaries of 
the skin are about intermediate between the other two. It therefore 
seems probable that the skin has some influence in respiration but 
only to a slight extent in Salamandra. 

In 1906 Whipple made a careful study of the respiratory move- 
ments of pulmonate Urodeles, and, while she did not use Salamandra 
itself, her conclusions are quite applicable to that animal. She finds 
that there are two forms of respiratory movements in Urodeles 
possessing lungs. The first consists of a shallow oscillation of the 
floor of the mouth which takes place rapidly and continuously, and 



THE RESPIRATORY SYSTEM, INCLUDING THE LARYNX 275 

involves only the air contained in the mouth and pharynx. This is 
known as bucco-pharyngeal respiration. The second is more compli- 
cated and involves the lungs, so that it is conveniently referred to as 
pulmonary respiration. 

More recently Willem (1923 and 1924) has carried out experi- 
ments on the respiratory movements of the mouth-floor of a large 
number of Amphibia, both Urodele and Anuran, including Sala- 
mandra, and has taken tracings of these movements by means 
of a stirrup placed under the head and attached to a pointer. 
(Text-figs. I, 2, and 3 are taken from his work.) Text-figure i is one 




Text-Fig. i. A tracing of the movements of the floor of the mouth, taken by means 
of a stirrup passing under the head, reading from right to left. The ordinates are magni- 
fied by 5 and the speed is 4 mm. per sec. The change in the level of the tracing is due to 
a movement of the head. (After Willem.) 



Text-Fig. 2. A tracing of the periodic variations in the amplitude of the pulmonary 
expirations. Read from right to left. (After Willem.) 

such tracing in which the ordinates, representing the amplitude of 
the oscillations of the mouth-floor, are magnified by five, the ribbon 
at the same time passing under the needle at the rate of 4 mm. per 
second. The movements involved in pulmonary respiration are 
represented towards the right-hand side of the figure as a deep de- 
pression in the otherwise more or less regular succession of bucco- 
pharyngeal movements. Text-figure 2 shows that there is a rhythmic 
variation in the amplitude of the pulmonary respirations, while text- 
figure 3 is an enlarged detail of one series of movements involved in 
filling and emptying the lungs. According to Willem the bucco- 
pharyngeal respirations occur at the rate of 1 20 per minute and have 
an amplitude of nearly 2 mm. The interval between successive pul- 
monary respirations depends on circumstances. When the animal 
is in a state of repose it is about fifteen minutes, but if the animal is 




276 THE SALAMANDER 

excited, or moving, or is fatigued through being handled, the interval 

becomes much shorter. 

The complicated movements carried out to effect the filling and 
emptying of the lungs are as follows. The depression of the mouth- 
floor is exaggerated and prolonged. The 
external nares are open during the first 
part of this act and the air is drawn into 
the mouth. This process is known as 
aspiration. Just as the mouth-floor is ap- 
proaching its lowest point the nares are 
closed, or nearly so, so that the residue of 
air is drawn from the lungs and mixed 
with the pure air already in the mouth. 
This process is called expiration. The 
mouth-floor then rises and, the nares re- 
maining closed, the mixed air is forced 
into the lungs. This constitutes the act 
£)iagram of the ^^ inspiration. The nares then open again 
respiratory movements of the and there follows a longer or shorter 
Salamander, comprising the clo- period of bucco-pharyngeal respiration 
sure of the nares (upper line), the before this more complicated process of 
rTTracinS'^'n^The'^T P^^^'^^U respiration is repeated. 
moTrVpreTure\w^ To summarize: it seems that in all 

(After Willem.) Urodeles devoid of gills, the bucco- 

I. Commencement of pharynx is the region which is chiefly 
aspiration. 2. Opening of concerned in the gaseous interchange 
the glottis and the commence- ^^tween the blood and the air, to which 
ment of the closure ot the , , • 11 /• 1 r 1 • 1 

nares. 3. The nares com- the skm and lungs (m those forms which 
pletely closed. 4. Closure possess them) are subsidiary. In support 
of the glottis. 5. Com- of this it should be noted that the whole 
mencement of the opening of of the mouth and throat is richly vascular, 
while Bethge has observed that the capil- 
laries of this region approach very close 
to the surface, actually pushing into the epithelial layer, between 
the cells. 



2 4 



Text-Fig. 3. 



the nares. 6. 

completely open. 



The 



BIBLIOGRAPHY OF SECTION VIII 
90, 91, 98, 99, 143, 144, 145, 146, 218, 233, 256, 259, 275, 293, 299, 391, 
392, 437> 438, 452, 4975 498, 520, 554, 555, 737, 79^, 797, 801. 



IX 

THE URINO-GENITAL SYSTEM 

General and Historical. 

This is perhaps the most widely and most accurately known system 
in the entire animal. The majority of the older authors made 
more or less accurate observations so far as the gross anatomy of the 
kidneys, gonads and their ducts is concerned (see Historical Intro- 
duction). Apart from such accounts the earliest author to study the 
system in any detail was Leydig (1853), who added many histological 
observations to his account. He observed and figured the recepta- 
culum seminis^ but failed to realize its function. He also recognized 
and correctly described the adrenal bodies, and he further figures 
and describes a curious pear-shaped epithelial bladder attached to 
the vestigial Miiller's duct in a male specimen, which he thought 
was a vestige of an anterior part of the kidney (he of course did not 
realize the true significance of Miiller's duct but thought it was 
connected with the ureters). Spengel (1876), the next worker of 
note, was unable to find anything which would correspond with the 
structure described by Leydig. Spengel's account of the urino- 
genital system of Amphibia is very detailed and accurate, and em- 
braces all three classes (Gymnophiona, Urodela, and Anura) involv- 
ing a number of genera in each class, and includes Salamandra. It 
is mainly owing to his labours that the anatomy of this system is so 
well known, and he deals not only with the macroscopic structures 
but also with histological details of the kidney tubules, &c. There 
has been no subsequent paper worthy of note dealing with the entire 
system, although a number of authors have treated particular sections 
in more detail. Their papers will be referred to later in the appropriate 
places. A possible exception may be made with regard to Fiir- 
bringer's paper (1878), which deals with the development of the 
organs concerned, while Gray's (1932) work on the development of 
the mesonephros in Triton throws much light on the general con- 
dition of this organ in Urodeles. 

The urino-genital system of Salamandra generally, and of the 
male particularly, is very interesting, since it shows an even more 
primitive condition than is found in the Selachians in that the anterior 
part of the kidney is less specialized for a purely genital function, and 
retains clear evidence of its segmental nature. 



278 THE SALAMANDER 

The kidneys in both sexes are paired and symmetrical and are 
very much more elongated than they are in the Frog. Each is narrow 
and ribbon-like anteriorly and becomes wider and thicker posteriorly. 
The former portion, called the 'sexual kidney' on account of its 
association with the gonad in the male (vide infra\ is a true mesone- 
phros. The recent work of Gray referred to above shows that in 
Triton the 'sexual kidney' is sharply defined from the posterior 
portion in its developmental history. He calls the latter the 'defini- 
tive kidney' and says that in this portion the primary definitive units 
arise from a set of vesicles distinct from those of the sexual part, and 
that these primary units eventually serve as collecting ducts for the 
secondary definitive units which bud off from them. There is thus 
not only a differentiation of function but also of fundamental struc- 
ture in the two portions of the kidney. 

It is of course unsafe to assume that the developmental history of 
the Salamander's kidney is identical with that of Triton, but there is 
every reason to suppose that it is similar^ particularly as far as the 
'definitive kidney' is concerned. The 'sexual kidney' is quite different 
in macroscopic appearance, and hence developmental differences are 
more likely to be present in this region if anywhere. Ciliated nephro- 
stomes are present, but show a degenerative tendency in the sexual 
kidney of the male (Spengel). The glomeruli are easily observed 
with the aid of a dissecting microscope, either directly in a freshly 
killed specimen or after injection with Prussian blue. The Mal- 
pighian bodies of the 'sexual kidney' are very regularly — but not 
segmentally — arranged in a linear row, while in the 'definitive 
kidney' the arrangement is necessarily more complex. 

Adrenal bodies are present and appear as small orange patches on 
the ventral surface of the kidneys in both sexes. They are not, how- 
ever, confined to the kidneys as they are in the Frog, but may be 
found anterior to them, as far as the subclavian artery, associated 
with the sympathetic ganglia. 

In both sexes there is to be found lying in the dorsal mesentery, 
ventral to the aorta, about 7 mm. posterior to the subclavian artery, 
a pair of small pear-shaped bodies, looking, at first sight, rather like 
large sympathetic ganglia, although they vary considerably in size 
and appearance. They seem to represent the structure described 
and figured by Leydig (see p. 277), but he found it on the right 
side and in the male only, whereas that described here normally 
occurs on both sides and in both sexes. Leydig thought that the 
structure was connected with Muller's duct, which is certainly not 
the case. It is, however, richly vascular and is connected with the 



THE URINO-GENITAL SYSTEM 279 

dorsal aorta and the post-cardinal veins. A microscopic examination 
of a series of transverse sections reveals its epithelial nature, and, 
while a few nerve-cells may be present, it is definitely not a ganglion, 
and their presence must be regarded as incidental and consequent 
on its close proximity to a true ganglion of the sympathetic chain. 
While, as already stated, the appearance of these bodies varies con- 
siderably, they not infrequently consist of a hollow epithelial vesicle 
with a knot of cells towards one side — in other words, they are 
strongly reminiscent of an abortive Malpighian body. At other 
times they are more dense and do not exhibit this appearance. Al- 
though an investigation of their embryological history would be 
necessary in order completely to elucidate their identity, it is sug- 
gested that they are probably of nephric origin and may possibly 
represent the vestige of a pronephros. 

Since there are definite sexual differences in the shape of the 
kidneys as well as in the arrangement of the ureters it is necessary 
to consider each sex separately. 

THE FEMALE SYSTEM (PI. XX) 
I. The Kidneys and Ureters. 

The two sections of the kidney as outlined above are distinguish- 
able, although the 'sexual kidney' is such in name only since it has 
no connexion with the female gonad. It also merges more gradually 
into the definitive kidney, which in turn is somewhat smaller in the 
female than in the male. The ureters (utr.) or Wolffian ducts are very 
different in the two sexes. In the female the duct of the 'sexual 
kidney' is a very fine longitudinal canal lying close alongside 
its lateral edge. This duct is joined at intervals by about eight 
transverse tributaries from the tubules of the kidney. As it 
approaches the 'definitive kidney' the Wolffian duct diverges some- 
what from its edge and passes backwards via the mesentery 
supporting the oviduct and, like this duct, becomes folded over to 
the ventral side of the kidney just before entering the cloaca. It 
receives tributaries from the kidney all the way along, and becomes 
correspondingly thicker. The fact that these tributaries enter the 
Wolffian duct approximately opposite the point where they leave 
the kidney is in strong contrast to the condition in the male. The 
duct opens into the cloaca at the apex of a small papilla just dorsal to 
the opening of Miiller's duct (oviduct). It remains entirely separate 
from Miiller's duct throughout its length, and does not discharge 
into it as some of the early workers supposed. 



28o THE SALAMANDER 

2. The Ovaries. 

The ovaries (ov.) are paired and are either symmetrically placed 
or else the right ovary is slightly anterior to the left, but the asym- 
metry is not so marked as in the case of the male gonad. The actual 
appearance and size of the ovary naturally depends on the season at 
which the animal was killed, but, assuming it to be approaching 
ripeness, it will measure about 30 to 35 mm. long and will contain 
some 7 to 20 large eggs about 3 mm. in diameter surrounded by 
numerous smaller ones. The eggs are very heavily laden with yolk 
and have no black pigment but are deep yellow in colour. The ovaries 
are supported by the mesovaria from the mesial side of the kidney, 
and resemble those of the Frog in structure in so far as they are 
enclosed in an ovisac which must be ruptured to allow the ripe eggs 
to escape into the coelom. The internal openings of the oviducts 
into which the eggs make their way, probably assisted by ciliary 
action, are situated near the bases of the lungs as in the Frog. In 
many, if not in all, females some vestiges of vasa efferentia and of 
Bidder s duct (which are only functional in the male, see p. 286) may 
be found. In any case these structures are hardly visible to the naked 
eye and are best sought for along the mesial edge of the kidney 
where the mesovarium is attached. 

3. The Fat-body. 

Th.G, fat-body (ft.b.) is markedly different from that of the Frog, 
both in position and shape. In the female it is suspended from the 
mesial aspect of the mesovarium and is a narrow ribbon-like structure, 
golden-yellow in colour, containing numerous fat globules which 
give it a spangled appearance. The seasonal variation in size is quite 
inconsiderable, and according to Funke (i 899), it can have but small 
influence as a somatic reserve. Apparently this function is mainly 
undertaken by the liver, which, in Salamandra, is characterized by 
a high fat content. Considerable fat reserves are also to be found in 
the cavities of the bones, around the spinal ganglia, and in the con- 
nective tissue between the muscles. 

4. The Oviducts (MuUer's Ducts). 

The oviducts (Mullers ducts) (od.) are a pair of white convoluted 
tubes passing the whole length of the abdominal cavity. They are 
never pigmented in the female. They open into the coelom at the 
ostia tuhae (ost.od.i.) situated just lateral to the bases of the lungs, 
and the ostia in turn expand into large funnels bounded by the 



THE URINO-GENITAL SYSTEM 281 

ligamenta tubarum (l.t.) and the ligamenta coronaria hepatis (l.c.h.) 
— see also p. 269. The oviducts open separately into the cloaca 
(ost.od.e.). Their lips protrude somewhat and are incised so as to 
give the openings a star-shaped appearance. The actual appearance of 
the oviduct within the abdomen will depend on whether the animal 
is pregnant or not, but in either case two regions are distinguishable, 
namely, an anterior convoluted portion which is dead opaque 
white with thick glandular walls, and a posterior 'uterine' portion 
in which the larvae develop. The latter part is thinner walled 
and is not convoluted, and it is this portion which alters very 
considerably in size and appearance during pregnancy. The figure 
(Fig. 72) was drawn from a female devoid of embryos and gives a 
good impression of the relations of the oviduct under such conditions. 
When the animal is pregnant, however, the 'uterine' portion (i.e. 
the portion posterior to the ij^ in the figure) elongates very consider- 
ably and becomes doubled on itself, protruding far forwards into 
the abdomen. In other words the junction between the glandular 
'oviducal' portion and the 'uterine' portion remains fixed near the 
posterior end of the abdomen, so that the enlargement and elonga- 
tion of the latter part must necessarily produce a duplicated loop 
as above described. The result is very striking in the early stages of 
the development of the eggs, when the whole structure has the 
appearance of a string of beads enclosed in a semi-transparent en- 
velope. It will also be noticed that the arrangement of the veins is 
different in the two portions. The blood from the glandular part 
drains into the longitudinal oviducal vein (v.od.long.) which in turn 
discharges into the post-cardinal vein, while the 'uterine' portion 
drains by numerous tributaries (v.od.p.) directly into Jacobson's 
vein. Stiive (1889) incorrectly describes the vascularization of the 
oviduct since he thought that there was actual extravasation of blood 
from the capillaries, which he compared with the menstruation 
of Mammals. A year later Wiedersheim committed the same error 
with regard to S. atra, but Schwalbe (1897) and Noble (1927) have 
corrected these mistakes and have shown clearly that the blood is 
normally confined to the capillaries. Nevertheless the whole 'uterine' 
region is very vascular, the capillaries forming a rich network 
immediately beneath the epithelial lining. 

Much has been written with regard to the development of the 
Miillerian ducts in Amphibia and only a very brief summary of 
this work can be given here. Fiirbringer (1878) believed that they 
split off from the Wolffian ducts, while on the other hand Gregg 
Wilson (1896— 8) found that the Miillerian ducts arose from the 



282 THE SALAMANDER 

peritoneal epithelium independently of the Wolffian ducts. Gemmill 
(1897) took a view midway between these two extremes and held 
that the anterior part of the oviduct arose as Wilson described from 
a peritoneal evagination, but that the posterior portion was formed 
by splitting off from the Wolffian duct. Finally Rabl (1903-4) 
affirms that the funnel is formed from the metamorphosed 'second' 
nephrostome of the pronephros, and that the part of the duct immedi- 
ately posterior to this, forming the anterior part of the duct, is 
formed by a thickening of the coelomic epithelium which then grows 
backwards independently to form the posterior part of the duct. 
Hall (i 904) confirms this mode of origin for Amblystoma, and hence 
it may be accepted as a final verdict on this vexed question. 

5. The Receptaculum Seminis and the Cloaca. 

The cloaca of the female is a simple chamber into which open the 
ureters, the oviducts, the rectum, and the urinary bladder. As men- 
tioned above, the openings of the ureters are situated immediately 
dorsal to the oviducal openings, which, in turn, lie on either side of 
the rectum, while the opening of the bladder lies directly beneath 
them ventral to the anus. The external opening of the cloaca is a 
longitudinal slit which, in the female, lies flush with the surrounding 
tissues and is not possessed of tumid or swollen lips. The lateral and 
dorsal walls of the cloaca are thrown into deep furrows, while on the 
dorsal wall, just behind the oviducal openings, may be seen a pair 
of pigmented patches. These mark the position of the receptaculum 
seminis (rec.sem.) which is usually considered to be the homologue 
of the pelvic gland of the male (see p. 287). 

The receptaculum seminis was noticed and described by Leydig 
(1853), but he did not observe the sperms within it, and did not 
realize its significance. This omission was supplied by von Siebold, 
who is usually credited with the discovery some five years later 
(1858). The organ consists of a number of small finger-like pockets 
protruding dorsally from the roof of the cloaca, which imbibe the 
sperms liberated from the spermatophore after it has been taken up 
into the cloacal chamber of the female (see also p. 5). The sperms 
are apparently able to, and normally do, live for a considerable time 
within the receptaculum. 

6. The Urinary Bladder. 

The urinary bladder (Fig. Gi^^ bl.) is a large bilobed sac with very 
thin walls lying ventrally in the posterior abdomen between the 
rectum and the pelvic girdle. It extends anteriorly almost as far as 



THE URINO-GENITAL SYSTEM 283 

the posterior margin of the Hver, closely adherent to the abdominal 
vein. Its walls, though thin, are well supplied with blood-vessels. 
There is no direct connexion between the bladder and the ureters, 
so that the urine can only gain access to it through the cloacal 
chamber (see also p. 287). 

THE MALE SYSTEM (PI. XXI) 
I. The Kidneys and Ureters. 

The chief differences to be observed between the kidney of the 
male and that of the female are the sharper distinction between 
the 'sexual kidney' and the 'definitive kidney', and the greater 
relative size of the latter. The 'sexual kidney' in this sex is directly 
associated with the gonad and serves to connect the vasa efferentia 
with the Wolffian duct, while the sperms actually pass through its 
tubules. Since Malpighian bodies are also present in the 'sexual 
kidney' it is presumably able to secrete urine, and hence its duct 
must be looked upon as a true urino-genital (Wolffian) duct. A 
much more striking difference between the sexes is noticeable in the 
ureters (utr.). In the first place although they are only small as they 
leave the kidney, they expand very rapidly and become much larger 
in the male and thicker walled (= appendices penis glandulosae, 
Funk), and secondly the urino-genital duct draining the 'sexual 
kidney' is not joined by the purely urinary ducts from the 'defini- 
tive kidney' until quite close to their common entry into the cloaca. 
Thus while the urino-genital (Wolffian) duct differs only in size 
(and function) from the corresponding portion in the female, the 
ureters proper consist of a series of about twelve C-shaped tubes of 
gradually decreasing length pursuing a parallel course round the 
ventro-lateral aspect of the kidney, until they finally enter a very 
short common duct and pass through the cloacal wall. A com- 
parison between Fig. 72 and Fig. 73 will make this clear. An 
interesting comparative series (mainly of Japanese Urodeles) has 
been worked out by Yamagiva (1924) which shows a progressive 
dissociation of the true ureters from the urino-genital duct during 
the course of evolution. 

The urino-genital duct appears at first sight to be continued for- 
wards beyond the anterior end of the kidney, and in fact this was 
believed by the older authors (Rathke, Bidder, &c.) to be the case. 
Leydig (1853) demonstrated the tubular nature of this anterior 
prolongation, and revealed it as the homologue of the Mullerian 
duct of the female, although he does not actually call it such. 



284 THE SALAMANDER 

He however made the mistake of imagining that in both the male 
and the female the ureters entered Miiller's duct some distance 
anterior to the cloaca. Spengel and Schneider in the year 1876 de- 
monstrated independently that the two ducts remain quite distinct 
throughout their course, and enter the cloaca by separate orifices. 
The former author made a careful study of the Miillerian duct 
in the male, and found that it was traceable, by means of sections, 
the whole way along the ventral side of the urino-genital duct to the 
cloaca, where it ended blindly. It normally extends forwards to the 
point on the posterior wall of the 'diaphragm', where the ostium 
tubae is situated in the female. It is bound up in a common connec- 
tive tissue capsule with the urino-genital duct so as to be indistin- 
guishable from it except in sections. It is noteworthy that Miiller's 
duct, and its associated urino-genital duct in the male, are always 
pigmented, while they never are in the female. The ureters may also 
bear pigment in the male, but it is usually absent or only very slight. 
There are no vesiculae seminales. 

2. The Testes. 

The testes (tes.) are paired and somewhat asymmetrically placed 
in the abdomen, the right side being slightly anterior to the left. 
They are suspended by the mesorchia from the mesial edges of 
the 'sexual kidneys'. Each testis consists of one, two, or three main 
lobes (usually two), and each lobe is further subdivided by constric- 
tions into two or three zones of different texture and colour, depend- 
ing on the state of development attained by the sperms within. The 
lobes of the testis are connected by a narrow strand of genital tissue, 
while a flagelliform projection of a similar nature extends anteriorly 
and posteriorly from the corresponding terminal lobes. Meves (1896), 
as the result of a cytological investigation, distinguishes the following 
zones in a testis lobe during July or August, i.e. when fully 'ripe'. 

(i) The anterior (or posterior) 'flagellum' and the connecting 
strand, greyish in colour, containing spermatogonia embedded in 
connective tissue. 

(ii) The anterior zone — forming the bulk of the lobe — also grey, 
containing spermatocytes, and 

(iii) a more opaque zone (or sometimes two zones), yellowish or 
whitish in colour, lying immediately posterior to the spermatocyte 
zone, containing ripe sperms. 

More recently (1922) Humphrey investigated the seasonal de- 
velopment of the testis in an American Urodele and obtained some 
interesting results which may well be recapitulated here, since they 



THE URINO-GENITAL SYSTEM 285 

are doubtless applicable to Salamandra or to any other Urodele 
possessing a multiple testis. The following is a summary of Hum- 
phrey's conclusions. 

In the young mature male the testis consists of but a single lobe, 
the multiple lobes being found only in older animals. The residual 
spermatogonia in the portion of the testis which is emptied in any 
given year, i.e. zone (iii) above, do not regenerate immediately, but 
remain dormant for several months, while the functional testis for 
the ensuing season (zone ii above) develops anterior to the emptied 
zone. Then supposing the testis at this stage to consist of but a 
single lobe, the spermatogonia of the posterior 'flagellum' become 
active and give rise to spermatocytes, thus forming a secondary lobe 
posterior to the first with the emptied dormant zone forming the 
connecting strand. This second lobe shifts forwards each season in 
a similar manner to the first, and eventually a third lobe may be 
formed posterior to it, but when each lobe reaches the anterior end 
of the genital tissue it just disappears and leaves the anterior 'flagel- 
lum' of dormant spermatogonia. Thus a continual spermatogenic 
wave moves slowly forwards along the genital strand, and the lobu- 
lated appearance is simply the result of this movement combined 
with the delayed regeneration of the emptied zones. 

Spermatogenesis has been investigated by a number of workers, 
notably von La Vallette St. George (1875), Flemming (1888), 
Nicolas(i892), von Rath (1893), Rawitz (1895), Meves (1895-6), 
and Champy (19 12). The haploid number of chromosomes is 12 
and the diploid 24. 

The sperm itself has been the subject of investigations by Czermak 
(1850), von Siebold (1850), and Retzius (1906). Although rela- 
tively short when compared with some other Urodeles, e.g. Triton, 
the sperm of Salamandra is long and the nucleus is filiform, while 
the acrosome is hooked. The middle piece is quite short and of the 
same diameter as the head. The flagellum bears on one side a crenate 
membrane along which a sinuous wave is continuously passing, so 
that it has been called the undulating membrane. The tail terminates 
in a short filiform end piece. The undulating membrane of course 
provides the means of locomotion. 

One hermaphrodite specimen has been described by Feistmantel 
(1902) in which eggs were found in the testis. 

3. The Vasa Efferentia, &c. 

The sperms leave the testis through a series (4 or 5) of fine ducts 
which pass through the mesorchium from the testis to the sexual 



286 THE SALAMANDER 

kidney. These are the vasa efferentia (va.eff.). They do not connect 
directly with the kidney tubules but enter a fine longitudinal duct 
which passes right along the mesial edge of the sexual kidney, and 
very close to it. This duct is in turn connected by a number of very 
short transverse tubes with the kidney tubules. The longitudinal 
collecting duct was first described by Bidder (1846) and is conse- 
quently called Bidder s duct (d.Bd.). According to Spengel the 
vasa efferentia are not ciliated, neither is Bidder's duct nor the con- 
nexions with the kidney tubules. The connecting ducts join the 
Bowman's capsules of the kidney at the pole opposite to that from 
which the uriniferous tubule leaves. As mentioned above some traces 
of this system may be found in the female, where of course it is 
functionless. 

4. The Fat-body. 

Th.Q fat-bodies (ft.b.) present no essential difference from those of 
the female {q-'V.). They are suspended from the mesorchia meso- 
dorsal to the testes. 

5. The Cloaca and its Glands. 

The cloaca of the male is very different from that of the female, and 
serves as a ready means for identifying the sexes by external char- 
acters, since, in the male, the cloaca is surrounded by a large tubular 
gland which gives its lips a tumid and swollen appearance, while in 
the female, where the gland is absent, the opening of the cloaca lies 
flush with the surrounding skin. The histology of the gland in Triton 
has been well described by Heidenhain (1890), but it is not easy to 
correlate his results with the condition in Salamandra, except in a 
very general way. The cavity of the cloaca is not quite a simple 
chamber, but is subdivided by infoldings of the wall. Of the diverti- 
cula thus produced the most pronounced are a pair of lateral grooves 
which run in an antero-posterior direction inclined slightly dorso- 
ventrally. They are somewhat L-shaped in transverse section, and 
their anterior extremities are expanded into small chambers in which 
the urino-genital papillae lie. There is further a small ridge-like pro- 
jection from the antero-ventral wall of the cloacal chamber which may 
be called the cloacal papilla (pap.cl.) (it has sometimes been called 
the penis^ but this is a misnomer). It may easily be seen, together 
with the lateral grooves, by pulling apart the lips of the external open- 
ing. The external lips of the cloaca are furrowed by a series (about 
1 5) of elongated pits, while the whole surface of the lateral walls of 
the chamber, especially ventral to the lateral groove, is thrown into 



THE URINO-GENITAL SYSTEM 287 

a series of deep knife-like ridges. These ridges are really composed 
of a number of long fine papillae amalgamated together, at the apices 
of which are the openings of the cloacal gland. The ridges are radi- 
ally arranged with the anterior extremity of the cloaca at the approxi- 
mate centre. Finally there is a slight median ridge passing along the 
dorsal wall of the cloaca, which apparently corresponds with the 
structure labelled /)«?;//j in Heidenhain's figures of Triton, but it is 
very much less developed in Salamandra. 

Histologically the glandular mass is divisible into three portions, 
called by Heidenhain the pelvic gland, the abdominal gland, and the 
cloacal gland. Of these the cloacal gland forms by far the largest 
portion and lies ventrally, i.e. ventral to the lateral diverticula above- 
mentioned, the abdominal gland lies postero-dorsally, and the pelvic 
gland antero-dorsally. The abdominal gland is very small and is not 
easy to distinguish from the pelvic gland, thus contrasting strongly 
with Triton, where the abdominal gland is well developed. 

On removing the skin the cloacal gland {^ = glandulae ani. Funk), 
appears as a heart-shaped structure (gl.cl.') surrounding the cloacal 
opening, and is seen to consist of a mass of coarse tubules. This ex- 
panded ventral portion of the gland is separated from a corresponding 
dorsal portion (gl.cl.") by the M. caudali-pubo-ischio-tibialis and M. 
ischio-caudalis, the two parts being connected by a mesial portion 
alongside the cloacal wall. It discharges its secretion by numerous 
fine pores situated at the extremities of the long papillae arranged in 
the cloacal walls as already described, as well as on the cloacal papilla. 
The abdominal and -pelvic glands open on the dorsal cloacal wall. 
Their tubules are smaller than those of the cloacal gland, and their 
openings are either flush with the surface or on very low papillae. 
The most anterior portion of the cloaca, i.e. the portion which ad- 
joins the rectum, consists of a wide tube with folded walls, and its 
epithelium is ciliated. In addition to the most anterior tubules of the 
pelvic gland it receives the rectum, the urino-genital ducts, and the 
bladder. The rectum merges directly into it, while the Wolffian ducts 
enter it at the apices of a pair of small urino-genital papillae on its 
dorsal wall (pap.u-g.), situated on either side of the middle line at the 
anterior end. These openings are relatively farther forwards in the 
male than in the female, and apparently correspond with the penis 
divisus of Funk. The opening of the bladder is not, as might be 
expected, at the extreme posterior end of this organ but a short dis- 
tance anterior to it, and consists of a longitudinal slit in its dorsal 
wall which opens ventrally into the cloaca immediately opposite the 
urino-genital papillae (Fig. 71). Thus, although there is no direct 



288 THE SALAMANDER 

connexion between the ureters and the bladder, the actual distance 
separating their openings is very small, and the urine, assisted by 
gravity, would quite naturally tend to flow into the relaxed bladder. 
The neck of the bladder, the posterior end of the rectum, and the 
anterior end of the cloaca are, in both sexes, surrounded by a strong 
fibrous sheath (fbr.sh.), containing smooth muscle elements, which 
is attached dorsally to the vertebral column and ventrally to the pubo- 
ischiadic symphysis. 

The function of the glands surrounding the cloaca is to secrete 
mucoid spermatophores which envelop the sperms when they are 
shed by the male in copulation (see also p. 5). The older anato- 
mists referred to the glands as the 'penis', but they are neither mor- 
phologically nor functionally to be compared with this organ. Leydig 
(1892) and others have sought to homologize the glands with the 
prostate of Mammals. Certainly there is some functional similarity, 
but an actual homology is doubtful. The cloacal opening, and indeed 
the whole gland, is surrounded by smooth muscles so that the tubules 
may be adequately compressed when their secretion is required. 

6. The Urinary Bladder. 

The urinary bladder presents no essential difference from that of 
the female (p. 282). 

BIBLIOGRAPHY OF SECTION IX 

51, 107, 127, 128, 136, 153, 157, 158, 159. 180, 182, 183, 184, 190, 209, 
210, 238, 268, 290, 296, 321, 334, 358, 373, 412, 416, 419, 420, 424, 469, 
470, 471, 472, 473, 513, 514, 526, 586, 587, 588, 592, 594, 595, 598, 602, 
614, 616, 669, 687, 702, 704, 709, 735, 753, 754, 802, 803, 819, 826. 



X 

THE DUCTLESS GLANDS 

General. 

Apart from histological considerations, and regarded from a purely 
anatomical point of view, there is not a great deal to be said relevant 
to the ductless glands of Salamandra. They were first subjected to 
a detailed investigation in 1888 by Maurer, who studied not only 
their character and histology in the adult, but also their origin and 
development. Bolau (1899) undertook a fresh investigation of the 
histology of the thyroid and thymus of a large number of amphibian 
types, including Salamandra, while Wilder (1929) discussed the signi- 
ficance of the ultimo-branchial body in Urodeles. This practically 
exhausts the anatomical literature of note, but papers dealing with 
the physiology of the organs concerned are much more numerous. 

I. The Thyroid (Thyreoid) Gland (Figs. 37 and 38, gl.thy.). 

According to Maurer, the thyroid develops as an unpaired struc- 
ture which very soon divides into two. It makes its appearance very 
early in the developing embryo. The thyroid in the adult consists of 
a pair of elongated ovoid glands about ^'S ^^- long by i mm. broad, 
lying on the floor of the throat at about the level of the os triangulare, 
immediately in front of the arterial arches, lateral to the MM. genio- 
hyoideus and rectus cervicis superficialis, and posterior to the M. 
interhyoideus. Each gland is enclosed in a tough connective tissue 
capsule into which a few fibres from the M. genio-hyoideus are in- 
serted. The glands are nourished by the thyroid arteries, branches 
of the external carotids, the blood being returned to the heart by the 
thyroid veins (p. 226). The hypoglossal nerve crosses the ventral 
aspect of each gland. Even with the naked eye, or at any rate with 
a hand lens, each gland is seen to consist of about 20 translucent 
follicles. The microscopic examination of transverse sections shows 
that each follicle consists of a single layer of cells which stain very 
deeply with iron-haematoxylin, and is filled with a clear colloidal 
fluid, the exact size and appearance of the follicles depending of 
course on the seasonal and physiological conditions prevailing at 
the time of fixation. 

As is well known, the secretion from the thyroid has a very 



igo 



THE SALAMANDER 



important Influence over the inception and course of metamorphosis 
in Amphibia. 

2. The Thymus Glands. 

The thymus glands are paired from the first, and develop from the 
dorsal ends of the third, fourth, and fifth gill slits (Maurer). Each 
gland has therefore a triple origin and frequently presents a trilobed 
structure in the adult. They are quite large ovoid glands, about 
4 mm. long by 3 mm. broad, and are situated laterally on either side 
of the 'neck' immediately beneath the skin, and about level with the 
posterior margin of the paratoid cutaneous glands. They are of the 
'solid' type of gland, and their cells, which are small, stain intensely 
with iron-haematoxylin. They are supplied by the cutaneous branch 
of the systemic arch (p. 201) and drained chiefly by the common 
facial vein (p. 225). The function of the thymus in Amphibia does 
not yet seem to be fully understood, but according to Noble (193 i), 
'the thymus functions in producing lymphocytes, granulocytes, and 
also erythrocytes to a certain extent'. It is apparently stimulated by 
thyroid feeding. 

3. The Parathyroids (Epithelial Bodies) (Fig. 38, ep.). 

These bodies are also derivatives of the visceral clefts, notably the 
third and fourth, but of their ventral ends. They are easily seen in 
dissecting, lying lateral to the arterial arches immediately ventral to 
the thymus, and consist of a pair of small spherical bodies on either 
side. They are highly vascular and are usually supplied by small 
twigs direct from the arterial arches. They appear late in larval life 
and apparently have the same function in Amphibia as in Mammals, 
namely to control the concentration of calcium salts in the blood 
(Waggener, 1929). They are widely separated from, and have no 
connexion with, the thyroids, although there is experimental evi- 
dence to show that the removal of the thyroids in toads causes a 
corresponding hypertrophy of the parathyroids. 

4. The Ultimo-branchial Body. 

This structure has also been variously termed post-hranchial hody^ 
supra-pericardial body^ and ultimo-branchial body. The last name has 
been adopted since it best indicates the origin of the structure from 
the sixth gill-slit. In the adult Salamander it normally occurs on the 
left side only, although in the Anura it is a bilateral structure. It is 
quite small and is best observed in sections, where it may be seen 
lying ventral to the M. cephalo-dorso-subpharyngeus and mesial to 



THE DUCTLESS GLANDS 291 

the M. rectus cervicis profundus, in the angle formed by these 
two muscles, at the level of the truncus arteriosus. It presents, when 
stained with iron-haematoxylin, a very similar appearance to the thy- 
roid gland, but consists of only about two or three small follicles. 
From the study of this body in a large series of Urodeles, Wilder 
(1929) came to the conclusion that, owing to its extreme variability, 
it could have little or no physiological importance. 

5. Other Glands. 

Adrenal bodies are present and may be seen as a series of orange 
patches along the ventro-mesial borders of the kidneys. They are 
also to be found anterior to these organs, even as far forwards as the 
subclavian artery, and they are then associated with the sympathetic 
ganglia (see also pp. 179, 182, and 278). 

The -pineal organ and -pituitary body are described in the section 
dealing with the brain (pp. 129 and 130). The spleen^ although a 
ductless gland, is described on p. 267, together with the alimen- 
tary canal. 'X\\q pancreas^ liver, and gonads \\2iVC2i\so endocrine func- 
tions although they can hardly be considered as 'ductless glands'. 

BIBLIOGRAPHY OF SECTION X 
70, 273, 274, 285, 286, 375, 419, 420, 452, 453, 527, 612, 742, 800. 



XI 

THE SKIN AND SENSE ORGANS 

THE SKIN 

1. General. 

The outer skin of the Salamander is moist and devoid of scales. It 
is highly glandular, however, and the glands are of two kinds — the 
mucus glands^ whose function is to keep the skin moist, and the venom 
glands. In this respect the skin of Salamandra is less slimy than that 
of the Frog, a feature probably correlated with the diminished respi- 
ratory function of the skin in the former animal. The mucus glands are 
distributed over the whole surface, and do not disclose their presence 
by any external excrescence or other sign visible to the naked eye. 

2. Venom Glands. 

The second kind of gland is the poison or venom gland. These are 
much larger than the mucus variety, and usually reveal their location 
by a slight mammilliform protuberance, while the central pore is quite 
visible to the naked eye, and where the glands occur beneath the 
yellow pigment — as is frequently the case — the opening is often 
marked by a black dot. Although the venom glands may occur 
almost anywhere on the body, they are more numerous over the dorsal 
surface, and are more highly developed in certain definite areas 
as in the paratoid glands and along the mid-dorsal line from the 
nape to the tip of the tail. There are also, strangely enough, several 
medium-sized glands on the feet between the toes. The paratoid 
glands consist of some twenty-five to thirty flask-shaped glands, 
the bases of which may readily be seen on removing the skin. A 
double alternating row of such glands extends right along the back for 
the whole length of the vertebral column. The remaining scattered 
venom glands are definitely smaller and less conspicuous. The 
venom glands are distinguishable from the mucus variety, particu- 
larly in preserved specimens, by their more granular contents and 
opaque appearance, the mucus glands being translucent or pearly. 

With regard to the histology and development of the glands much 
has been published which is outside the scope of the present work. 
For these details the papers of Leydig (1876), Pfitzner (1880), 
Drasch (1894), Ancel (1900-2), Schuberg (1908), Nierenstein 



THE SKIN AND SENSE ORGANS 293 

(1908), and Theis (1932) may be consulted. Ancel showed that the 
poison glands were of ectodermic origin, while Nierenstein held that 
they arose secondarily from degenerating mucus glands, a view with 
which Esterly — for Plethodon — agrees. Recently, however, Theis 
has reinvestigated the whole question and concludes that there is no 
transition from mucus to poison glands. 

When a Salamander is irritated, e.g. by chloroform, a white milky 
fluid is seen to exude from the venom glands. This substance is a fairly 
potent irritant poison and eventually causes paralysis in the victim. 
Although there is no mechanism whereby the animal can inject the 
poison into a victim, and it must therefore remain a purely defensive 
weapon, nevertheless the Salamander can, on occasion, squirt or spray 
the fluid to a surprising distance, quite sufficient to get into the eyes 
and nose of a too inquisitive marauder. Probably this action is 
efl'ected by means of the smooth muscle elements which invest the 
glands (cf. also Esterly, 1904). 

3. Venom. 

A fairly considerable amount of work has been carried out in an 
endeavour to elucidate the chemical nature of the poison, and two 
alkaloids have been isolated from it in a pure state. The first worker 
in this field was Zalesky (i 866) who isolated the chlorhydrate of one 
of them, which he called Samandarin^ and suggested for it the formula 
CggHeeOioNa. His choice of name was due to a suggestion by Prof. 
Roth of Tubingen that the Greek word araXafjidvSpa comes from 
the Persian Samandar, a similar word occurring in both Arabic 
and Hindustani. Zalesky also investigated its physiological eff"ects. 
The next worker was Faust (1898 and 1900) who isolated the sul- 
phate of both alkaloids. To Samandarin sulphate he gave the formula 
C52H80N4O2+H2SO4 or (C26H4oN20)2+H2S04. The second base 
he named Samandaridin and suggested the formula (C2oH3iNO)2+ 
H2SO4 for its sulphate. He supposed that they were both built 
around the methyl pyridin group C5H5(CH3)N and that Samandarin 
has more of these groups than Samandaridin. He also found that the 
pharmacological strength of the former was 7-8 times greater 
than that of the latter. Gessner (1926), and Gessner and Craemer 
(1930) have recently made a fresh analysis of the Salamander venom 
and have isolated both substances in the pure state. Their analysis of 
Samandaridin does not quite agree with that of Faust, having one 
atom of carbon less and one atom of oxygen more than his, namely, 
C19H31 O2N, with a freezing point of 187° C. 

It is not proposed to catalogue the numerous pharmacological and 



294 THE SALAMANDER 

physiological experiments which have been carried out with these 

substances. They are fully described in the papers mentioned. 



THE SENSE ORGANS 
I. THE EAR. 

1. General. 

The auditory apparatus of Urodeles consists, as in all higher verte- 
brates, of a membranous labyrinth of ectodermal origin for the com- 
bined functions of balancing the body and the perception of sounds, 
together with certain skeletal elements modified for the purpose 
of conveying the auditory stimuli from the external medium 
to the labyrinth. These two distinct portions have each been the 
subject of separate investigations by the various workers dealing 
with the organ of hearing, and accordingly they will also be dealt 
with separately here. 

2. Skeletal Structures. 

The skeletal portion is the more highly specialized and is very 
characteristic of the group. 

The earlier authors dealing with the ear of the Salamander, as well 
as those investigating the structure of the skull in general, recognized 
only one skeletal element within the fenestra vestibuli (ovalis), namely 
the operculum. The first investigator to recognize the homologies 
of all the skeletal parts concerned and to give an adequate account 
of their relations was Gaupp (1898), but their full significance and 
mode of operation were not completely understood until the investi- 
gations of Kingsbury and Reed (1902—20). 

It is perhaps almost unnecessary to point out that the familiar 
tympanum and tympanic cavity of the Frog are wanting in Sala- 
mandra, or for that matter in all Urodeles. Nevertheless a columella 
is -present^ but it remains quite vestigial and does not reach the outer 
surface, and is a.ppa.rently fuficlionless in terrestrial adults. There is a 
considerable amount of variation among the several families of Uro- 
deles, and these differences have been used for classificatory purposes 
by Reed (1909 and 1920), although Dunn (1923) questions their 
value in this respect. It is unnecessary to go into the question in 
detail here, but the reader may refer, either to the original papers 
quoted, or to the summary given by Goodrich (1930, p. 480). 

In the Salamander there are two structures filling the fenestra 
vestibuli (ovalis) of the auditory capsule — (i) the columella^ which is 
extra-otic in origin, and (ii) the operculum^ a cartilaginous plate cut 



THE SKIN AND SENSE ORGANS 295 

out from the walls of the ear capsule. The former is the functional 
element in the larva, and the latter in the adult. 

The columella develops in a ligament, the suspensor-stapedial liga- 
ment^ stretching from the membrane of the fenestra to the proximal 
end of the quadrate and squamosal. It consists of two portions — first 
a short rod-like stylus connecting the suspensorium (squamosal and 
quadrate) with the second portion — a flat plate-like expansion filling 
the anterior portion of the fenestra. In the adult Salamander the 
columella is ossified and fuses with the wall of the auditory capsule 
so as to be scarcely distinguishable from it. 

The operculum (fig. 2, cop.) fills the remainder of the fenestra 
vestibuli. It is a plate of cartilage, approximately circular in plan, and 
shaped like a very thick crescent in transverse section. It is attached 
to the auditory capsule at its edges by the membrane of the fenestra, 
and thus lies entirely free from other skeletal parts, being supported 
by an elastic membrane only. As already mentioned it develops from 
the wall of the ear capsule, but it is perhaps questionable whether it 
is 'cut out', as it were, by absorption of the tissue round it, or whether 
it arises by an extension of cartilage into the membrane closing the 
fenestra. It appears probable that the latter is the chief process 
at work. The operculum is attached to the supra-scapula by a strap- 
like muscle derived from the M. levator scapulae, and called by Gaupp 
M. opercularis. The presence of this muscle is at first sight rather 
extraordinary, and it is absent in those forms, e.g. Necturus, which 
lead an entirely aquatic existence in both larval and adult phases. In 
such forms also the columella \s free from the auditory capsule but 
fused with the operculum, and it is supposed that in aquatic forms, 
both larvae and adults, the auditory stimuli reach the ear from the 
external medium — water — by means of the following path : — jaws, 
suspensorium, columella, fenestra. Such forms, particularly when 
adult, are much given to resting the whole of the body, including the 
jaws, on the bottom, and the lower jaw is thus in a favourable position 
for receiving vibrations directly from the ground. Now in terrestrial 
forms, e.g. Salamandra, the head is always kept well clear of the 
ground, and, since it is manifestly impossible for the vibrations of a 
rare medium like air to affect so heavy a skeletal structure as the 
jaws, the auditory stimulus must travel by some alternative route in 
these animals. This route is by means of the fore-limbs to the scapula 
and suprascapula, and from thence to the operculum via the M. oper- 
cularis. In this connexion it is interesting to note that Dunn (1922) 
gives a description of a primitive Urodele — Hynobius leechi — in which 
the columella is free from the auditory capsule, and there is in 



296 THE SALAMANDER 

addition a M. opercularis joining the operculum to the shoulder, 
whereas in almost all higher forms either one or other alone is present. 
The 'hearing' of Urodeles must obviously be confined to the per- 
ception of a very small range of low-frequency vibrations of wide 
amplitude, but it is possible that the M. opercularis may have the 
effect of increasing the range somewhat, since, being a striped muscle, 
its tension is capable of adjustment by the animal. 

3. The Membranous Labyrinth (PI. XXIV, figs. 80 and 81). 

Apart from the earlier workers (e.g. Scarpa, Pohl, Huschke, Win- 
dischmann, &c.) the first investigator to make a really detailed study 
of the membranous labyrinth of Amphibia was Hasse (1873). He 
was succeeded by Kuhn (1880), whose paper was, in turn, closely 
followed in the next year (i 88 i) by the well-known work of Retzius. 
All these authors give excellent descriptions of the ear of Salamandra 
and are, in the main, correct. They all fall into the same error in 
imagining that the ductus perilymphaticus communicates with the epi- 
cerebral space — a mistake which was corrected later by O'Neill, 
Sterzi, Harrison, &c. Both Hasse and Kuhn imagined that t\\G.pars 
neglecta (their /)<2rj tnitialis) was the anlage of the cochlea of mammals, 
but Retzius did not share this view, and to him is due the name -pars 
neglecta. Both the earlier authors describe a pars hasilaris supported 
by a 'cartilaginous' frame (Knorpelrahmen). Retzius denies the exis- 
tence of both a pars hasilaris and a 'Knorpelrahmen'. In this, how- 
ever, he is himself in error (as Harrison (1902) showed), and the 
earlier authors were right. Both Kuhn and Retzius deal with the 
microscopic structure of the maculae, &c. 

The membranous labyrinth of Salamandra is fairly typical of Uro- 
deles in general, but differs in several points from that found in 
Anura, e.g. the Frog. After clearing away the muscles arising from 
the auditory capsule the ridges over the semicircular canals may be 
clearly seen. In order to remove the labyrinth from the capsule — 
not a particularly difficult operation — it is advisable to decalcify the 
skull by immersing it in sulphurous acid. The roof of the capsule may 
then easily be cut away. Care must be taken with the anterior septum 
semicirculare (a bony pillar joining the roof to the ventro-mesial wall 
of the capsule), which lies just posterior to the anterior semicircular 
canal, and serves to separate this canal from the main cavity. The 
other two septa, related in the same way to the other semicircular 
canals, are rarely complete, so they do not usually give trouble. 

Compared with the size of the skull as a whole the ear is large and 
flat, and, in most places, is well separated from the bony capsule. 



THE SKIN AND SENSE ORGANS 297 

The older authors (Hasse, Kuhn, and Retzius) imagined that this 
space contained perilymph, and that the labyrinth simply floated free 
in this fluid supported by a few strands of connective tissue. Harri- 
son (1902) showed that this was not the case and that the perilymph 
was restricted to certain definite regions (see below), while the re- 
maining space between the labyrinth and the capsule was occupied 
by a tissue which he termed perilymphatic tissue. This is a connective 
tissue which becomes specially dense, 'almost cartilaginous', around 
the labyrinth itself. 

The following may be termed the 'intrinsic' portions of the ear — 
thrtt semicircular canals with their ampullae^ the utriculus^ the sacculus^ 
the lagena^ pars basilaris and pars neglecta^ and the ductus endolym- 
phaticus. The 'extrinsic' portions are the perilymph system and the 
saccus endolymphaticus. 

The semicircular canals are not all equal in extent or in curvature. 
The shortest, and at the same time the most nearly circular, is the 
posterior canal (can.s.p.). It lies in a vertical plane pointing in a 
postero-lateral direction relative to the axis of the body. It may be 
supposed to arise from the pars superior utriculi just behind the an- 
terior canal, and, after completing almost an entire circle, it re-enters 
the utriculus at t\\^ pars posterior ]\x^\. below the origin of the horizontal 
canal. Its ampulla is situated at the ventral end (amp. p.). 

The horizontal canal (can.s.h.) (or canalis externus, Kuhn) arises 
from the pars posterior utriculi between the two ends of the posterior 
canal. The curvature is not quite even throughout its length. The 
posterior portion describes roughly the quadrant of a circle and is 
then succeeded by a more flattened mesial portion. Finally the 
anterior end bends round more sharply again and enters the recessus 
utriculi close to the anterior canal. It does not lie in a truly hori- 
zontal plane relative to the other canals, but is slightly depressed 
anteriorly. The ampulla is at the anterior end (amp.h.). 

The anterior canal (can.s.a.) arises from the sinus superior utriculi 
close to the posterior canal, and lies in a vertical plane pointing in an 
antero-lateral direction. It remains almost flat for about two-thirds 
of its length and then bends sharply ventralwards to enter the dorsal 
aspect of the recessus utriculi. The ampulla is at the anterior end 
(amp. a.). 

The utriculus is a wide tube, more or less saddle-shaped, sitting 
over the mesial part of the sacculus. Its most dorsal portion, the 
broad sinus superior (s.s.utric), gives rise to the anterior and posterior 
canals, while its anterior end dips ventralwards to the recessus utriculi 
(rec.utric), which is the expanded portion into which the ampullae 



298 THE SALAMANDER 

of the anterior and horizontal canals open. Posteriorly the utriculus 
also turns ventrally into the sinus posterior^v^hicWis continued into the 
ampulla of the posterior canal. The utriculus further communicates, 
by means of an oval opening, with the sacculus. This opening, the 
canalis utriculo-saccularis (can.utric-sac), is situated at the base of the 
utriculus below the sinus superior, and from its mesial wall the pars 
neglecta (p.neg.) (Retzius) is evaginated. 

The sacculus (sac.) is a lens-shaped sac, compressed in a ventro- 
mesial plane. It contains otolith granules and communicates with 
the saccus endolymphaticus by means of the endolymphatic duct (d.el.). 
It also opens into the utriculus^ as already noted, and into the lagena 
and pars basilaris. The lateral wall is extremely thin and separates 
the sacculus from the main perilymphatic space — the spatium saccu- 
lare. The ductus endolymphaticus (d.el.) arises from the dorso-mesial 
aspect of the sacculus close to the canalis utriculo-saccularis and passes 
directly dorsalwards, mesial to the utriculus and close to its wall, 
and directly between the origins of the two vertical canals. At about 
the level of the dorsal margin of the sinus superior utriculi the duct 
passes through the foramen endolymphaticum into the cranial cavity, 
where it expands into the saccus endolymphaticus (see 'Membranes of 
the Brain', p. 121 et seq.). 

The lagena (lag.) (or lagena cochleae, Retzius) is a somewhat 
pear-shaped pocket opening from the mesial aspect of the sacculus 
near its posterior margin. It contains an otolith. 

The pars basilaris (p.bas.). Although both Hasse and Kuhn had 
correctly described the pars basilaris in Salamandra, yet Retzius 
(1881) denied its existence as a separate recess, and supposed it to 
be represented by a small oval nerve-ending on the inner wall of the 
upper end of the lagena. Harrison (1902) corrected this error in 
favour of the older authors — a result which is here confirmed. The 
pars basilaris is a small evagination from the dorso-mesial wall of the 
lagena facing the opening of the latter into the sacculus. 

4. Nerve-endings. 

The ear is supplied by the eighth cranial nerve. The fibres of this 
nerve terminate on plate-like areas of sensory cells, called maculae, 
which occur at the following positions — at the ampullae of the semicir- 
cular canals, here called cristae by Retzius, on the floor of the recessus 
utriculi, the sacculus, xh^ pars neglecta, pars basilaris, and the lagena. 

The cristae acusticae ampullorum (Retzius) are bi-concave areas 
lying on the ventral surfaces of the posterior and horizontal ampullae, 
and on the anterior face of the anterior vertical ampulla. 



THE SKIN AND SENSE ORGANS 299 

The macula acustica recessus utriculi (Retzius) lies on the floor of 

the recessus utricuh and, together with the two anterior (horizontal 

and vertical) ampullae, it is innervated by the ramus anterior of N. 

acusticus. 

The macula acustica sacculi (Retzius) is the largest of all the sensory 
plates and covers the mesial wall of the sacculus and is supplied by 
the ramus medianus^ N. acusticas. 

The macula acustica neglecta lies on the dorsal wall of the pars 
neglecta^ while the sensory area of the pars basilaris practically sur- 
rounds that part of the labyrinth, and that of the lagena is confined 
to its mesial wall. The last three maculae mentioned, together with 
the crista acustica of the posterior ampulla, are supplied by the 
ramus posterior of the auditory nerve. The histology of the maculae, 
&c., has been well described by Kuhn and Retzius from osmic acid 
preparations. Briefly the maculae consist of a superficial layer of 
cylindrical cells with sensory hair-like processes, supported by round 
basal cells. 

5. The Perilymphatic System. 

The correct elucidation of the extent and relations of this system 
is due to Harrison (1902). His terminology is therefore used here. 
Within the ear capsule the largest space containing perilymph is the 
spatium sacculare. This large sac lies lateral to the sacculus and mainly 
ventral to the horizontal canal, and fills the whole lateral half of the 
capsule. It lies in close apposition to the membrane closing the 
fenestra vestibulae, and to the operculum. The wall separating this 
space from the sacculus is exceedingly thin. 

Leaving the spatium sacculare posteriorly, on its dorso-lateral 
aspect is the ductus perilymphaticus (d.pl.), a fairly wide tube only 
slightly smaller in diameter than the semi-circular canals. The 
ductus passes mesialwards around the posterior end of the labyrinth 
and lies ventral and lateral to the posterior end of the horizontal 
canal, mesial to the posterior canal and the sinus posterior of the 
utriculus, finally emerging on the mesial aspect of the labyrinth, 
dorsal to the lagena and pars basilaris, and ventral to the pars neg- 
lecta. It then dips ventralwards and enters the cranial cavity through 
the apertura ductus perilymphatici (O'Neill) (the foramen rotundum of 
Hasse), in the mesial wall of the otic capsule. Within the cranium 
the duct expands into a comparatively small sac — the saccus peri- 
lymphaticus (or spatium meningeale^ Harrison). See also p. 126. 

There are two small outgrowths from the ductus perilymphaticus 
within the ear capsule. These occur just proximal to the point where 



300 THE SALAMANDER 

the duct enters the cranium. The first has been called the recessus 
partis neglecta by Harrison who first described it. It is a short wide 
diverticulum which comes into intimate contact with the ventral wall 
of the pars neglecta, the separating wall being exceedingly thin. The 
second diverticulum is smaller and is the recessus partis hasilaris of 
Harrison. It bears a similar relation to the pars basilaris. 

These three areas, at the pars neglecta, pars basilaris, and sacculus, 
where the membrane separating the perilymph from the endolymph 
is extremely thin, are spoken of by Harrison as 'tympanal areas', and 
it is, he supposes, through these areas, and through them alone, that 
the vibrations imparted to the perilymph by the operculum are trans- 
mitted to the endolymph. By these vibrations the otolith crystals in 
the sacculus and lagena are set in motion, and these in turn affect the 
sensory 'hairs' on the surface of the cells covering the maculae acus- 
ticae, and thus the auditory nerve is stimulated. The function of the 
saccus endolymphaticus within the cranial cavity is presumably similar 
to that of the foramen rotundum of the mammalian ear, namely to 
act as a buffer for the vibrations of the perilymph. 

II. THE EYE 
I. General. 

The eye of the Salamander is relatively large and stands up promin- 
ently from the surface of the head. The bulhus oculi is nearly spherical, 
and, in the larva, is protected by a scleral ring of cartilage which, 
however, disappears at metamorphosis so as to leave no trace in the 
adult. The lens is also nearly, but not quite, spherical. According 
to Beer (1899) accommodation is not effected by a change in the 
curvature of the lens but by an alteration in its position, since he 
finds that it moves towards the iris on stimulation. This movement 
is brought about by a ciliary muscle which constricts the back 
chamber and so forces the lens out, while the iris is said to play a 
small but unimportant part. 

Klingelhoffer (19 10- 11) shows that the refractive index of the 
Amphibian eye in general is such that it is a little long-sighted in 
water, and that the lens, being less truly spherical than that of the 
fish, and possessing some slight capacity for accommodation, is truly 
intermediate between an aquatic and an aerial structure. 

Johnson (1926) during the course of an ophthalmological investi- 
gation of Amphibian and Reptilian eyes found that 'Salamandra 
maculosa exhibits the simplest and apparently the most primitive 
condition of the disc.^ It is quite round and very small, being only 
' The point of entry of the optic nerve, E.T.B.F. 



THE SKIN AND SENSE ORGANS 301 

one-third or one-quarter that of the majority of the Anura. It is dull 
white in colour and is not covered by anything, nor is there pigment 
nor differentiation of any kind, nor the slightest trace of any vessels, 
either hyaloidean or discoidal.' In this last feature Salamandra differs 
markedly from the Frog. Another difference between these two 
animals lies in the absence, in the Salamander, of any trace of the 
vertical pigmented stripe which extends from the lower edge of the 
pupil to the margin of the iris in the Anura. Johnson supposes that 
this stripe, which is always associated with a groove and which some- 
times extends to the upper portion of the iris also, is the result of the 
mechanical strain of the iris muscles, and is phyletically the oldest 
condition, and that the round pupil of the Salamander represents the 
highest stage of development. The same author finds considerable 
accommodating powers, but no movement of the eye itself other than 
that of retraction. This last feature is not surprising in view of the 
relatively weak development of the ocular muscles other than M. 
retractor bulhi. 

Quite recently Dr. Ida C. Mann (1931) has made an exhaustive 
comparative study of the iris pattern in vertebrates, and notes that 
in the Salamander the arteries enter the iris from below and on the 
temporal side, while in some specimens the inferior iris artery is a 
branch of the temporal and does not enter separately. This latter she 
thinks is the more typical Urodele arrangement. The arteries break 
up irregularly round the pupil and the blood drains away by a few 
radial veins, not definitely superficial to the arteries and not arranged 
with anything like the regularity found in Reptiles. The pigmenta- 
tion of the iris is entirely brown. 

2. Glands of the Orbit. 

Apart from histological details there is not much that can be added 
concerning the structure of the eye itself. It is supplied by a pair of 
ciliary nerves, a superior (p. 140) and an inferior (p. 140), while the 
bulbus oculi is attached to the wall of the skull by a strong fibrous 
sheath surrounding the optic nerve^ the whole forming the opticpeduncle. 
Within the orbit, beneath the bulbus oculi, is a mass of glandular 
tissue which shows a tendency to become differentiated into two 
sections. The anterior region is the Harderian gland^ and the pos- 
terior the lacrimal gland. In Salamandra the separation of the Har- 
derian portion in the anterior angle of the orbit from the remainder 
of the gland may be partial or complete. In Triton there is no such 
separation (Sardemann, 1887). Hence this animal may be regarded 
as exemplifying the primitive condition, while the Salamander shows, 



302 THE SALAMANDER 

by comparison, that the Harderian and lacrimal glands have arisen 
by differentiation from a common glandular mass. 

The naso-lacrimal duct runs from the anterior corner of the eye to 
the nasal cavity passing beneath the pre-frontal bone in so doing. 

3. The Eyelids. 

The upper eyelid is the larger and is movable, as it is in the whole 
family Salamandridae. The lower lid, according to Johnson, is prob- 
ably homologous with the memhrana nictitans of higher vertebrates, 
the true lower lid of these animals arising as a cutaneous fold external 
to it, only a slight trace of this fold being detectable in the Sala- 
mander. Both upper and lower lids are moved by means of a tendon 
attached to the M.. retractor bulhi and they are therefore closed 
synchronously with the retraction of the eye-ball (cf. p. 51). In 
addition to this mechanism there are a few smooth muscle elements 
in the upper eyelid so that it is apparently capable of some independent 
movement. 

III. THE NOSE 

1. General. 

The olfactory passages of the Salamander are relatively large 
chambers situated within the nasal capsules. With the exception of 
a small area around the external openings they are completely pro- 
tected by the skeleton. Two distinct regions are distinguishable in 
each olfactory chamber, namely, the main chamber, or cavum nasi, 
which is shaped like a flattened ovoid, from which protrudes a lateral 
gutter-like diverticulum — the sinus lateralis nasi. Into this lateral 
gutter the ductus naso-lacrimalis opens, while the external narial 
opening is situated just mesial to its anterior end. The sinus lateralis 
nasi is lined by ciliated epithelium, but not the main cavum nasi. 

2. Jacobson's Organ. 

In 1895 Seydel suggested that Jacobsons organ occurred in the 
sinus lateralis, and named the gland lying just mesial to it, beneath 
the cavum nasi, Jacobsons gland. In 1898 Mihalkovics opposed this 
view on the ground that, in all other vertebrates which possess a 
Jacobson's organ (including the Frog), this structure is found mesial 
to the posterior nares and not lateral to them. He therefore concludes 
that a true Jacobson's organ is absent in Salamandra, but agrees that 
the sinus lateralis nasi — his recessus maxillaris — has a similar function, 
namely that of testing the expired current, and he therefore regards 



THE SKIN AND SENSE ORGANS 303 

it as analogous to but not homologous with a true Jacobson's organ. 
Three years later, in 1901, Hinsberg studied the development of the 
nasal capsule in Triton and came to the conclusion that, in that 
animal, a patch of sensory epithelium, homologous with Jacobson's 
organ, arises mesially and shifts to a position lateral to the internal 
nares hejore this opening develops. Thus he maintains that the 
lateral position is a specialization of the Urodela, but that neverthe- 
less the homology is true. Zuckerkandl (19 10) agrees in the main 
but concludes, on the grounds of its nerve supply, that the Jacobson's 
organ of Urodeles cannot be homologous with that of the Amniota. 
Bruner (19 14) agrees with Seydel as to the location of Jacobson's 
organ, while Herrick (19 14), also on neurological grounds, doubts 
the existence of this structure in the Urodela, but says that in Anura 
it has assumed a definitive form as a diverticulum from the mesial side 
of the nasal sac, with its characteristic innervation by the vomero-nasal 
nerve. Von Navratil (1926) does not agree with Seydel's identifica- 
tion of the sinus lateralis nasi with the organ of Jacobson on morpho- 
logical grounds, but thinks that Seydel's sensory epithelium may 
become the sinus maxillaris. In 1927 Kurepina investigated the 
development of the primary choanae in Urodeles (Triton) and 
arrived at conclusions diametrically opposed to those of Hinsberg. 
He claims the existence of a primary oro-nasal groove between the 
nose and mouth, and hence there can be no lateral shifting of Jacob- 
son's organ such as Hinsberg describes. 

Thus, while it still remains a somewhat doubtful question, there 
is evidently a growing consensus of opinion among morphological 
embryologists against Seydel's view, and in favour of the absence of 
Jacobson's organ in Urodela. The absence of a vomero-nasal nerve 
may perhaps be taken as the deciding factor. It is generally agreed 
however that the sinus lateralis nasi is t}\e functional equivalent o^ the 
Amniote organ of Jacobson, i.e. that it is used for testing the ex- 
halent current. 

3. Opening and Closing Mechanism. 

Another region of special interest in the nasal organ is the 
apparatus for opening and closing the external nares. This was dis- 
covered and described by Bruner (1896 and 1901) and consists of 
a set of three smooth muscles, one constrictor and two dilatators. 
There is no cartilaginous support to the edge of the narial opening 
as there is in the Frog. The opening in the cartilaginous nasal cap- 
sule, in which each external naris lies, is very much larger than the 
actual cutaneous opening, and houses the muscular apparatus as well. 



304 THE SALAMANDER 

It is the fenestra narina (Stadtmiiller) or fenestra rostro-nasalis 
(Bruner) (PI XXIV, fig. 82). 

TheM. constrictor naris (m.c.n.) (Bruner) bounds the posterior edge 
of the narial opening for rather more than a semicircle, its ends being 
inserted on the anterior wall of the fenestra on the cart, cupullaris. 

The M. dilatator naris (m.d.n.) (Bruner) arises from the posterior 
border of the fenestra narina, i.e. from the cart, retro-narina, and 
passes directly forwards to the posterior border of the external naris, 
deep to the constrictor muscle. It is inserted on the cutaneous wall 
of the narial opening. 

M. dilatator naris accessorius (m.d.n'.) (Bruner) is an obliquely 
directed muscle. It arises from the lateral border of the fenestra 
narina, partly from the cartilage and partly from the maxilla. The 
muscle passes obliquely antero-mesially and is inserted in the postero- 
lateral margin of the narial opening, deep to, and amongst, the lateral 
fibres of the M. constrictor naris. 

In his later paper (1901) Bruner further showed that the relation 
of the nasal muscles to the nasal gland, the glandula nasalis externa^ 
is such as to force the secretion from the gland simultaneously with 
the closing of the nasal opening. He also showed that they arise in 
situ from the mesoderm and are not derived from any other muscles. 

The internal nares are large and unprotected by valves. 

The glands of the nasal capsule may be reckoned as consisting of 
two masses, one external and one internal. The glandula nasalis 
externa has already been mentioned. It is situated above the sinus 
lateralis nasi within the fenestra narina in close proximity to the 
external naris, and to the opening of the ductus naso-lacrimalis into 
the nasal capsule (d.nas-lc). As described above it is compressed by 
the M. constrictor naris, so that its function would seem to be that 
of keeping the edges of the external narial opening moist. The 
glandula nasalis interna practically envelops the cavum nasi, and lies 
entirely within the nasal capsule. Its function is to keep the olfactory 
epithelium moist. As mentioned above Seydel distinguishes the 
ventral portion as Jacobsons ^ 



BIBLIOGRAPHY OF SECTION XI 

2,5.6,13,14,15,18,25,34,87,88,89,91,134,139,140,152,174,177,178, 
216, 223, 243, 244, 245, 251, 287, 294, 295, 297, 310, 311, 313, 338, 339, 
345, 352, 372, 374, 376, 377. 3^©, 397> 40i, 405, 409, 4^5, 422, 423, 439, 
443. 455, 482, 483, 485, 490, 491, 509, 515, 533> 534, 535, 53^, 573. 574. 
575, 576, 577. 582, 604, 605, 613, 650, 680, 685, 699, 700, 733, 741, 745, 
772,795,804,831,838. 



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See also Zool. Anz., 1890, Bd. xiii, pp. 589-607. 

732. 1895. 'The Cranial Nerves of Amphibia. A Contribution to the Morpho- 
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BIBLIOGRAPHY 339 

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INDEX 



Note. All blood-vessels, nerves, and muscles have been indexed under their Latin 
names regardless of whether they have been referred to in the text by the Latin or 
English terminology. Thus, all references to the peroneal nerve vpill be found under 
Ne-rvus peroneus, and those to the vertebral vein under Fena ^ertebralis, while certain 
of the more familiar terms have been listed under both systems, so that, for example, the 
subclavian artery will be found under Arteria subclanjia and also by its English name 
under the letter 'S'. 



Abdomen, muscles of, 94-8. 
Abdominal gland, 287. 

— vein, 220, 231-2, 235-7^ anterior, xxi, 
xxii. 

— viscera, xv, xviii, 265-7, 272, 278-86, 
Abducent nerve, xxx, 50, 51, 130, 138. 
Acetabulum, 44, 46. 

Acustico-facialis ganglion, 139, 144, 149. 
root, 126, 129, 130. 

Adrenal bodies, xv, xxiii, 277,278, 291; asso- 
ciation with sympathetic system, 179, 181. 

Age, in captivity, 8. 

Ahl, 8. 

Albertus Magnus, xiv. 

Albinism, occurrence of, 15. 

Albrecht, 17. 

Aldrovandus, xiii, XIV. 

Alimentary canal, 262—6; arteries of, 206-8; 
compared with Frog, 262; glands of, 266- 
7; relief pattern of, 264-5. ^^^ "-^^^ Gut. 

Alizarin, use of, 16, 263. 

AUantois, xxi, xxii. 

Alps, xiv, 13. 

Alveolar canal, 144. 

Amblystoma, loi, 124, 134, 135, 137, 145, 
148, 159, 235, 251, 254, 282. 

Amphibia, characters of, i. 

Amphiumidae , i. 

Amplexus, 5, 6. See also Copulation, Coition. 

Ampullae (of semicircular canals), xxi, 297. 

Anal gland, xxiii. See also Cloacal gland. 

Ancel, 292, 293. 

Andersson, 179, 180. 

Anterior (semicircular) canal, 297; ampulla 
of, 149. 

— cardinal vein, 230. 

— commissure, 131. 

— epigastric veins, 228, 230-1, 232, 237. 

— subclavian ganglion, 180. 

— visceral (sympathetic) nerve, 181. 
Antrum petrosum laterale, 26, 142, 144, 145; 

artery of, 199; vein of, 222; lymphatics of, 
258. 



Anus, xxvii; fibrous sheath of, 288. 5"^^ also 

p. 266 and p. 287. 
Aortic trunks, xxv, 190. 
Apertura ductus perilymphatici, 126, 299. 

See also Foramen perilymphaticum. 

— nasalis externa, 29. See also External 
nares. 

Aplectana brevicaudata, 10. 

Aponeurosis lingualis, 66. 

Apparent gape, 11, 262. 

Appendicular skeleton, 37-47. 

Appleton, 99, 102-9, 114, 157, 173, 175, 

176. 
Arachnoid, 124, 125, 128. 
Arcifery, 39. 
Arcus dorsalis, 213. 

— venosus dorsalis manus, 233, 234. 

pedis, 240, 241. 

Aristotle, xiv, xvii, xix. 
Arteriae abdominales, 215. 
Arteria alveolaris, 34, 200. 

— articularis genu fibularis, 216; tibialis, 
216. 

— auditiva, 246. 

— basilaris, 246. 

— brachialis, 211, 212. 

— carotis cerebralis, 199, 244; r. anterior, 
244; r. posterior, 245. 

externa, 197-8. 

interna, 30, 138, 150, 198-201; 

pharyngeal branch of, 200. 

— cerebralis anterior dorsalis, 245. 
medialis, 245. 

ventralis, 244. 

— circumflexa scapulae, 211. 

— cloacae medialis, 209. 

— coeliaco-mesenterica, 206. 

— collateralis radialis, 213. 

ulnaris, 213. 

Arteriae costales, 205. 

j Arteria cutanea antibrachii lateralis, 212; 
I medialis, 212. 
cruralis lateralis, 217; medialis, 217. 



346 



Arteria cutanea femoris lateralis; 
terior, 216. 

genu medialis superior, 216. 

magna, 211. 

— cystica, 207. 
Arteriae digitales, 218, 
Arteria dorsalis pedis, 218. 
Arteriae dorso-lumba]es, 214. 
Arteria duodeno-hepatica, 206. 

— duodeno-pancreatica, 207. 

— epigastrica anterior, 211; posterior, 211, 
215. 

— femoralis, 214, 215, 216. 

— gastrica anterior, 206; posterior, 206. 

— gastrico-Iienalis, 206, 238. 

— glutea, 214, 218; r. cutaneus cruralis 
posterior, 214. 

— hyomandibularis, 202. 

— iliaca communis, 214-18. 

externa, 214-15. 

interna, 214. 

Arteriae intercostales, 205. 

Arteria interossea (of arm), 212, 213; (of leg), 

217, 218. 
externa (of arm), 213; r. recurrens 

posterior, 213. 

— -(of leg), 217-18; 1 

— intestinalis, 206. 

— ischiadica, 214, 215, 216 

— labialis, 200. 

— laryngis, 218. 

— lingualis, 198. 

— mandibularis, 199. 

externa, 202; interna, 202. 

— mandibulo-jugularis, 201. 

— maxillaris anterior, 204; externa, 201, 
202 ; lateralis, 204; medialis, 204. 

— maxillo-palatina, 203. 

— mediana, 212. 

— mesencephali superior, 246. 

— mesenterica anterior, 207; posterior, 207, 
208; prima, 206, 207. 

— nasalis lateralis, 204; medialis, 204; pos- 
terior, 204. 

— obturatoria, 215. 

— occipitalis (of Frog), 204. 

— oesophagea dorsalis, 218; ventralis, 218. 

— ophthalmica, 200, 204. 

superior, 199, 201; r. anterior, 201; 

r. posterior, 201. 

— orbitalis anterior, 204. 

inferior, 199-200; superior, 199. 

— orbito-nasalis, 203-4; compared with A. 
occipitalis of Frog, 204. 

Arteriae oviducales, 208-10; anteriores, 209; 
mediales, 209; posteriores, 208. 



INDEX 
pos- 



recurrens, 2iJ 



217, 218. 



Arteria palatina anterior, 203; posterior, 203. 

— palato-nasalis, 145; 203. 

— pectoralis, 211. 

— pelvica, 215. 

— perforans carpi, 44, 84, 213. 
cruris, 2x7. 

inferior, 218; superior, 213, 217. 

tarsi, 47, 218. 

— petrosa lateralis, 26, 198-9; abnormalities 
of, 200. 

— pharyngea, 202. 

ascendens, 200, 201; possible homology 

with r. auricularis (A. cutanea magna) of 
Frog, 202. 

— plantaris, 217. 

lateralis, 217; medialis, 217. 

— procoraco-humeralis, 212. 

— profunda brachii, 211; r. collateralis ra- 
dialis, 213; r. cutaneus brachii posterior, 211. 

femoris, 216. 

— pterygoidea, 203. 

— pudenda, 214, 215. 

— radialis, 212. 

— recurrens radialis, 212. 

— renalis, 208. 

superficialis, 208-9. 

— spinalis ventralis, 132, 246. 

— subclavia, 210-13; relation to post- 
cardinal veins, 230. 

— sublingualis, 198, 202. 

— subscapulars, 211. 

— supracoracoidea, 211. 

— temporalis, 199, 200. 

— thoracica, 210. 

— thyroidea, 198. 

— tibialis medialis, 217. 

— trapezia, 212. 

— ulnaris, 212, 213; r. recurrens, 213. 

— vertebralis, 204-5. 
collateralis, 194, 204. 

— vesicalis, 214, 215. 

— volaris, 213. 

Arterial arches, 187, 195, 196. 

Arteries, spermatic, 208. 

Articular, 33, 34. 

Arytenoid cartilages, 271-3. 

Asbestos, xiii. 

Aspiration, 276. 

Atlas vertebra, 16, 17, 159; relation to skull, 

17- 
Atria (or Atrium), xxv, xxix. See also Auricle. 
Atwell, 123, 130, 131. 
Auditory capsule, 24-6, 125, 138, 149, 294, 

295. See also Otic capsule, Ear capsule. 

— nerve, xxi, xxx, 25, 149. See also N. 
acusticus. 



INDEX 



347 



Auditory organ, xv, xviii, xxi, xxiv, xxvii, 
71, 126, 294-300; artery of, 246. 

— ossicle (s), xxiv, xxv; absence of, xvii, 

— stimuli, path of, 295, 300. 

Auricle (Auricles), xv, xvi, xviii, xix, xxi, 
xxii, xxiv, xxv, xxvii, xxix, 187-8; left, 
187, 191, 193; right, 187, 191, 192; re- 
cesses of, 188. 

Auricular septum, xxv, 185, 187, 188. See 
also Septum atriorum. 

Auriculo-ventricular opening, 186, 188, 189; 
valves of, 188. 

Axial skeleton, 16-37. 

Back, muscles of, 92-3; arteries to, 214; 

lymphatics of, 256; veins from, 229, 239. 
Balfour, 147. 

Basalia (of carpus), 42-4; of tarsus, 47. 
Basi branchial, 35. 
Basicranial fontanelle, 130. 
Batrachia, xx, xx\'iii. 
Baur, 43. 
Bazin, 249. 

Beddard, 237, 239, 244. 
Bedriaga, 13, 14. 
Beer, 300. 
Beer, de, 156. 
Belon, xiv. 
Bendz, 132. 
Benecke, 5. 
Benninghof, 189. 
Berger, 127. 
Bethge, 194, 200-1, 204, 206, 211, 225-6, 

230, 232-3, 235, 274, 276. 
Biceps muscle, 77, 81. 
Bidder, xxx, 283, 286. 
Bidder's duct, xxx, 286; vestige in female, 

280. 
Bile duct, xv, 265, 267, 
BiscHOFF, xxvii. 
Bladder (gaU), xv, xvi, 267, artery to, 

207. 

— (urinary), xvi, xix, xxi, 282, 288; arteries 
to, 215; lymphatics of, 253-4; opening of, 
282, 287; veins from, 236. 

Blainville, xxiv. 
Blanchard, xxiii. 

Blood, separation of, 185, 189, 191-3. 
Blood-vessels, general features of, 195-6; his- 
torical, xxv, xxix, xxx, 194-5. 
Blumenbach, xviii. 
Boas, 185. 
Bochenek, 123. 
Body, length of, 11, 12. 
BOLAU, 289. 
Bolkay, 27. 



Boulenger, E. G., 13, 14. 
— , G. A., 2, 14. 
Brachet, 268. 

Brachial nerve, 78, 167. See also, N. brachia- 
lis. 

— plexus, 157, 159, 163. 
Brachycoelium salamandrae , 10. 

Brain, anatomy of, 128-31; arteries to, 244-6; 
commissures of, 131; epithelial portions of, 
131; historical, xxv, xxvii, 123; membranes 
of, 123-8; size of, 128; supply of pure 
blood to, 193; veins of, 246-8; ventricles 
of, 131. 

Brain-case, 22, 29, 32. See also Cranium. 

Branchial arches, 7, 34-5, 36-7. See also 
Visceral arches. 

— (lymphatic) sinus, 259. 
Braus, 43. 

Breathing, see Respiration. 
Breeding habits, xx, xxvi, 4-7. 
Bremer, 22, 23. 
Bronchi, 271, 272. 

BrONGNIART, XX. 

Broom, 24, 30. 
Brucke, 185-6, 191. 
Bruner, 29, 303, 304. 
Buccal-force-pump, xvii, xxvii, 271. 

— ganglion, 146. 

— respiration, xvii, xix, xxi, 192, 274-6. 
Bucco-pharynx, blood capillaries of, 276; 

importance in respiration, 192, 274-6. 
Buckland, xviii. 
Bulbus cordis, 185, 186, 189, 191; hyaline 

cartilage in, 189; origin of, 190; valves of, 

189. 

— oculi, 300, 301. See also Eye. 
burckhardt, 123. 
Burkard, 49, 55. 

Calori, 249. 
Camerano, 274. 
Canalis utriculo-saccularis, 298. 
Capitulum (of rib), attachment of, 18. 
Cardiac nerves, 155, 156. 
Carotid arch, xxv, 195, 196; branches of, 
197-201. 

— canal, 30, 199. 

— gland, xxi, xxii, xxix, 192, 197. 
Carpus, 40, 42, 43, 44, 47. 
Cartilage, intervertebral, 17. 
Cartilago antorbitalis, 22, 23, 50, 51. 

— cupullaris, 23, 304. 

— ectochoanalis, 22, 23. 

— lateralis, 64, 271, 272, 273; pars laryngea, 
271-3; pars trachealis, 271-3. 

— obliqua, 23. 



348 INDEX 

Cartilage pre-hallucis, 47, iii, 112, 113. 

— pre-pollicis, 43, 91. 

— retro-narina, 23, 304. 

— ypsiloidea, xx, 44, 45, 95, 96; homology 
of, 45; function of, 45. 

Carus, xxi, xxii, 48, 54, 57, 59, 60, 65, 68, 
69, 71-5, 79, 81-8, 92, 94, 99, 100, 102-3, 
105, 107, 109-12, 114. 

Caucasian mountains, 2. 

Caudal artery, 20. 

— lymphatics, 250, 254. 

— muscles, 98, 102. 

— nerves, 179. 

— vein, 20, 220, 239. 

— vertebrae, 16. 
Caudata, characters of, i. 
Caudo-crural muscle (Humphry), 102, 105. 
Cavum aortico-caroticum, 190. 

— cranii, 24, 25, 30, 136. See also Cranial 
cavity. 

— epicerebrale, 126. 

— internasale, 24. 

— nasi, 302. 

— pulmo-aorticum, 190. 

— vestibuli, 26. 
Central lymph heart 



190, 256, 260. 
Centrale (of carpus), 42-4, 83-4; of tarsus, 

47- 
Centrum, 18. 

Cera to-branchial elements, 35-6. 
Cerato-hyal, 34. 
Cerebellum, 129, 131. 
Cerebral hemispheres, 128, cavities of, 131; 

plexus of, 245; veins of, 246. 
Cervical plexus, 162. 

— vertebra, 16; homology of, 17. 
Cestode parasites, 10. 
Champy, 285. 

Chevron bones, 20. 

Chin, 54, 66. 

Chioglossa, i. 

Choanae, 22, 264. See also Internal nares. 

Choanal glands, 264. 

Chondrocranium, 22-7. 

Chorda tympani, 49; homology of, 146-8. 

Chorioid plexus of the first ventricle, 245, 
248. 

of the fourth ventricle, 246, 247, 248. 

Choronshitsky, 195, 238-9, 267. 

Chromosomes, number of, 285. 

Ciliary ganglion, 135, 140; possible sympa- 
thetic nature of, 136. 

— muscle, 300. 

— nerves, 135, 140, 301. 
Cisterna linfatica, 255. 
Claus, 19. 



Climbing powers, 12. 

Cloaca, of female, 282; of male, 286-8; 
arteries of, 209, 215-16; external appear- 
ance of, 12; lymphatics of, 252-3, 254; 
veins of, 236, 243. 

Cloacal ganglion (sympathetic), 182. 

— glands, xvii, xix, xxiii, 102, 287-8. 

— papilla, 286, 287. 

— sinus (lymphatic), 252, 253. 
Coccidian parasite, 9. 
Cochlea, anlage of, 296. 
Coeliac ganglion, 182. 
Coeliaco-mesenteric complex, 206. 
CoGGi, 125. 

CoGHiLL, 52-4, 134-7, 142-3* i45> 147-8. 
CoiTER, xiv, XV, xix. 

Coition, 4-6. See also Amplexus, Copulation. 
Colouration, 2, 12-15. 
Columella, xxiv, 32, 294, 295. 

— ethmoidalis, 23. 
Common bile duct, 267. 

— carotid, 197. 

— facial vein, 219, 225. 
Copula, 34, 35, 37, 66, 69. 
Copulation, 4, 5. 
Coracoid, 38, 39, 40, 41. 
Cordylus, xiv. 

Cornua trabeculae, 22, 23. 
Coronoid process, 34, 52, 53. 
Corpus callosum, 131. 

— opticum, 129, 136; artery of, 245. See 
also Optic lobes. 

Cosmocerca commutata, 10. 
Costal arteries, 205. 

Cranial cavity, 23, 29. See also Cavum 
cranii. 

— nerves, xxv, xxx, 132-56. 
Cranium, 24-5. See also Brain-case. 
Crista dorsalis humeri, 41, 79. 

— muscularis, 24. 

— retrosellaris, 25. 

— tibiae, 46, 108, 112. 

— ventralis humeri, 40, 73, 76, 81. 
Cristae acusticae ampullorum, 298. 
Criticism of Perrin, 105-6. 
Crofts, 266. 

Crural plexus, 157, 159, 170. 
Cryptobranchus, 92, 96, 108, in, 188, 231, 

240-3. _ 
Cucullaris muscle, 72. 
Cutaneous glands, xvi, ir, 292-3. 

— respiration, 274. 

CuviER, xix, xx, xxi, 51-3, s^, 65, 67, 73, 75. 
Cystic artery, 207. 

— (bile) duct, 267. 
CzERMAK, xxii, xxx, xxxi, 285. 



INDEX 



Dalecampius, xiv. 

DaUDIN, XX. 

Davy, xix, xxiv. 
— , Dr. J., xxiv. 

— Sir Humphry, xxiv. 

Definitive kidney, 278; of female, 279; of 
male, 283. 

Degenerating larvae, number of, 6. 

Deltoid muscle, 77. 

Dempster, 126. 

Dentary, 33, 34, 53. 

Dentition, 262. 

Development, 3, 4; intra-uterine, 3, 6. 

Diaphragm, xvii, 97, 194, 269, 270. 

Diemyctylus, 42, 45. 

Diencephalon, 128, 130. See also Mid- 
brain. 

Diet, 8; of larva, 7. 

Digits, 40, 42-7; arteries of, 213-18; flexor 
muscles of, 88-9, 117; homology of, 42. 

DioscoRiDES, commentary on, xiii. 

Dipnoi, 227, 232. 

Disease, fungoid, 9. 

Distinction between sexes, 12, 279, 286. 

Distomum crassicolle (in gut), 9; D. cygnoides 
(in bladder), 9. D. endolobum (in gut), 9. 
D. glabrum (in gut), 9. 

Distribution, geographical, 2, 8-9, 13-15. 

Dorsal aorta, xxv, 205 et seq.; origin of, 205; 
branches of, 206. 

— fissure, 132. 

— mesentery, 268. 

— pancreatic duct, 265, 267. 
Dorso-lumbar arteries, 214. 
Drasch, 292. 

Druner, 26, 35-7, 49, 51-69, 71, 72, 123, 
133, 139, 144, 146-56, 159-61, 183, 199, 

201, 221, 222, 225-7, 259, 273. 
Ductless glands, 289-91. 

Ductus Botalli, 63, 185, 186, 192, 195, 197, 

202, 218. 

— Cuvieri, 187, 193, 219, 220, 231-3; left, 
219; right, 191, 219; possible communica- 
tion with lymphatic system, 257. 

— endolymphaticus, 25, 125, 297, 298. 

— naso-lacrimalis, 11, 29, 141, 302, 304. 

— perilymphaticus, 126, 296, 299. 

— thoracicus, 256. 
Du Fay, xvi. 

Ducfes, xxviii, 20, 28, 30, 36-8, 45, 48, 52-3, 
55, 56, 60, 6^, 67, 69, 71-9, 81-3, 85-7, 
90, 99, 100, 102-3, 105? 108-IQ. 

Dujardin, xxxi. 

Dumas, Provost and, xxiv. 

DuM^RiL, XX, xxiv. 

Dunn, 294, 295. 



349 

lymph- 



Duodenum, 265; arteries to, 206-7 

atics of, 255. 
Dura mater, 124, 125, 127, 128, neural layer 

of, 124-8; periosteal layer of, 124, 126-8; 

pigment cells of, 124. 
DiJRIGEN, 15. 
DUTROCHET, xxiv. 

DuvERNOY, xvii, xix. 

Ear, xvii, xxiv, 294-300; intrinsic portions 
of, 297; extrinsic portions of, 297, 299; 
artery to, 246; nerve of, 149. 

— capsule, 25, 26, 29, 30, 138, 150, 295. See 
also Auditory capsule, Otic capsule. 

— ossicles, xxiv. See also Auditory ossicles. 
Ecology, 15. 

Ectoparasites, absence of, 10. 
Ectopterygoid, homology of, 31. 
Edgeworth, 36, 49, 52-8, 60, 62-4, 67-9, 

72, 271, 273. 

Egg(s), xiv, xvi, xviii, xxii, xxiii, xxiv, xxix, 

5, 280. See also Ova. 
Egg membrane, xxii. 
Eimeria salamandrae, 9. 
Eisler, 42, 70, 75, 78-91, 98-100, 107, 

1 10-2 1. 
Ellipsoglossa, 59, 60. 
Embryo, xxiv; early development of, 5; 

hatching of, 6; weight of, 6. See also 

Larva. 
Emery, 43. 
Enchytraeid worms, 7. 
Endorachis, 128. 
Engler, 37, 38, 39, 40. 
Epencephalon, 129. 5"^^ also Cerebellum. 
Epicondylus lateralis humeri, 41, 81-3. 

— medialis humeri, 41, 85-7. 

— lateralis femoris, no, in, 112, 114, 115. 
Epicoracoid, 39. 

Epidermis, xv. See also Skin. 

Epidural space, 128. 

Epigastric veins, 228, 230-1, 232; compared 
with those of Reptiles, 237; median, 237. 

Epiphysis, 129. See also Pineal body. 

Epipubis, 45. See also Ypsiloid cartilage. 

Epithelial bodies, 290. See also Parathyroid. 

Epithelium, of mouth floor, 146, 151, 152; of 
palate, 144, 145; of pharynx, 150, 155; 
olfactory, 134, 141, 145. 

Esterly, 293. 

Ethmoidal region, 22. See also Nasal cap- 
sules. 

Euproctus, I. 

Eurycea, 274. See also Spelerpes. 

Eustachian tube, xxiv; absence of, 262, 264. 

Exaeretus caucasicus, 2. 



350 



INDEX 



Expiration, 276. 

Extensor muscles of fore-arm and hand, 8 1 ; 
of leg and foot, no. 

— nerves of fore-limb, 164-7; of hind-limb, 

171-5- 
External carotid artery, 197-8. See also A. 
carotis externa. 

— gills, XV, xvi, xviii, xxiii; loss of, 7. 

— jugular vein, 219, 220, 225. See also V. 
jugularis externa. 

— mandibular nerve, 143. 

— nares, 11, 23, 28, 29; opening and closing 
of, 303- 

— trochanter, 103. 
Eycleshymer, 129. 

Eye(s), II, 300-1; accommodating powers 
of, 300; disc of, 300; lens of, 300; metamor- 
phosis of, 7, 300; muscles of, 49; refractive 
index of, 300. 

Eyelids, 13, 55, 302; muscles of, 51; nerves 
of, 142. 

Fables, xiii. 

Facial nerve, xxx, 26, 144—9, 150. 

Facialis canal, 26, 144. 

Fascia cephalo-dorsalis, 57, 62, 72, 74. 

— pectoralis, 58, 59. 

— plantaris, 115, 117. See also Plantar 
aponeurosis. 

Fat-body(ies), xv, xvii, xxii, 280, 286; 

arteries of, 208; veins of, 227. 
Fat reserves, 280. 
Fatio, 6. 
Faust, 293. 
Favaro, 250, 254, 255. 
Fecundation, 3, 4, 5. See also Fertilization. 
Feistmantel, 285. 

Femoral nerve, 105, 107, 171, 172, 173. 
Femoro-fibular muscle (Humphry), 105, 

109. 
Femur, 45, 46. 
Fenestra basalis, 22, 23, 145. 

— basicranialis anterior, 25; posterior, 25. 

— dorsalis nasi, 23. 

— infra-conchalis, 23, 141. 

— narina, 23, 304. 

— olfactoria(ae), 23, 24, 134. 

— ovalis, 294. 

— vestibuli, xvii, xxi, xxiv, xxvii, 24, 26, 
294, 295, 299. 

Ferric chloride, use of, 130. 

Fertilization, xiv, xviii, xxix, 4, 5. See also 

Fecundation. 
Fibrae cruciatae, 64. 
Fibula, 46. 
Fibular nerve, lyy-S. 



Fibulare, 46, 47. 

Fifth visceral arch nerve, 153, 155; r. recur- 

rens, 153. 
Fire, reaction to, xiii; resistance to, xvi. 
First hepatic (bile) duct, 267. 
Fischer, xxx, 52-4, 56, 60, 62-7, 132-3, 

135, 156. 
Flemming, 285. 
Flexner, 124, 128. 
Flexor muscles; of the hand, 43, 84-92; of 

the foot, 114-22. 
Foetal membranes, xiv, xv. 
Foetus(es), xiv, xv, xviii, xxii, xxiii. See also 

Larva, Embryo. 
Food, 8; of larvae, 7; method of securing, 61. 
Foot, pronation of, 116, 119. 
Foramen abducens, 138. 

— acusticum(a), 25. 

— apicale, 23. 

— endolymphaticum, 25, 298. 

— for N. sp. I, 17; for N. sp. 2, 17. 

— inferior dental, 146. 

— intervertebral, 17, 19. 

— magnum, 17, 29. 

— metoticum, 150. 

— nutritium, 17. 

— obturatorium, 44. 

— oculomotorium, 24, 135. 

— of Monro, 131. 

— opticum, 24, 134. 

— orbito-nasalis laterale, 23, 141; mediale, 
13, 141. 

— palatinum, 144. 

— perforans carpi, 44. 

— perilymphaticum, 25. Seealso-p^p. 126,299. 

— post-oticum, 26, 150. 

— pro-oticum, 138, 139. 

— rotundum, 126, 299, 300. 

— supracoracoideum, 38, 162, 163. 

— vertebrale, 1 8 . 

Fore-arm, 40, 41-2; arteries of, 194, 212-13; 

muscles of, 81-90; nerves of, 165-9; veins 

of, 233-5. 
Fore-brain, 123, 128. 
Fore-limb(s), 11, 40-4; arteries of, 212-13; 

muscles of, 76-92; nerves of, 163-70; 

sympathetic nerves of, 180; veins of, 233-5. 
Forma typica, 2, 13, 14, 15. 
Fossa cubitalis ventralis, 41. 

— olecranon, 41. 

Fourth (cranial) nerve, xxx. See also N. 
trochlearis. 

— ventricle, roof of, 131; plexus covering, 
246-8. 

— visceral arch nerve, 152, 155; r. post- 
trematicus, 153; r. pre-trematicus, 153. 



INDEX 



351 



37-9' 44. 48, 
81, 85, 93-4, 
-12, 115, 132, 



Foveal depressions, 46. 

Francis, 36. 

Fritsch, 185, 194. 

Frog(s), XV, xvii, xix, xx, xxiv, xxvi, xxviii, 

29, 30, 32, 125, 128, 129, 181, 186, 187, 

191, 240, 296, 301, 302. See also Rana. 
Frontal (bone), 23, 27, 28, 29, 32. 
Fronto-squamosal arch, i. 
Froriep, 17, 147, 160. 
Fungoid disease, 9. 
Funk, xxi, xxv, xxvi, 10, 27, 

52-4, 59, 60, 65-7, 71-9, 

96, 99, 100, 103, 107-8, lie 

194, 229, 283, 287. 

FUNKE, 280. 

FtJRBRINGER, 38, 67-81, 156-7, 161-8, I 70, 
277, 281. 

Gachet, xxvi. 

Gadow, 9. 

Gait, 12. 

Gall-bladder, xv, xvi, 267; artery of, 207. 

Ganglion, acustico-facialis, 139, 144, 149. 

— anterior iliac, 183. 
subclavian, 180. 

— ciliary, 135, 140; possible sympathetic 
nature of, 136. 

— cloacal (sympathetic), 182. 

— coeliac, 182. 

— copulare, 151, 183. 

— Gasserian, 133, 138, 139, 142, 144. 

— glossopharyngeus-vagus, 150, 151, 152, 
156. 

— habenular, 123. 

— laterale VII, 146. 

— otic, 146. 

— palatine, 141, 145, 183. 

— posterior subclavian, 180-1. 

— spinal, 19, 157. 

— subclavian, anterior, 180; posterior, 180-1. 

— trigeminus, 133, 138, 142, 144. See also 
Gasserian ganglion. 

Gape, actual, 262; apparent, 262. 

Gasserian ganglion, 133, 138-9, 142, 144. 

Gastric nerves, 156. 

Gaumenfortsatz, 23. 

Gaupp, id, 21, 29, 33-4, 41, 71, 93, 124-5, 

136, 157-8, 202, 204, 212, 217, 223, 

231, 246-7, 294-5. 
Gegenbaur, 37, 38, 

273. 
Gehirnhiille, 126. 
Gehuchten, 123. 
Gelderen, van, 223, 
Gemmill, 282. 
Generic characters, i. 



40, 42, 43, 263, 272, 



Genio-hyoid muscle, 65. 
Genital ducts, xvi, xxvi. See also Miiller's 
duct, Vasa efFerentia, Wolffian duct. 

— organs, xvi; system, xvii. See also pp. 
277-88. 

Genitalia, xvii, xxii, xxvi, xxvii. See also 

pp. 277-88. 
Geographical distribution, 2,8-9, 13, 14, 15. 
Gesner, xiii, xiv, xvii. 
Gessner, 293; and Craemer, 293. 
Gill(s), xxiii, xxiv, xxvi, 2; external, xv, xvi, 

xviii, xxiii, 7; internal, xvi, xxiv. 
Gill arches, xvi, xxiv. See also Branchial 

arches, Visceral arches. 

— openings, xxiv; slits, xvi, 7. 

Glands, cloacal, xvii, xix, xxiii, 102, 287-8 ; 
ductless, 289-91; Harderian, 301-2; labial, 
262; lacrimal, 301-2; mucus, 292-3; para- 
toid, II, 13, 292; poison ('milk', venom) 
xvi, II, 292; tonsillar, 264. 

Glandula intermaxillaris, 263. See also 
Intermaxillary gland, 24. 

— nasalis externa, 304; interna, 304. 

— palatina, 263. 
Glenoid cavity, 37, 40. 
Glomeruli, 278. 

Glossopharyngeal nerve, xxx, 26, 60, 61, 149, 

150. 
Glossopharyngeus-vagus complex, 133; 

origin of, 129. 

— ganghon, 150, 151, 152, 156. 
Glottis, xix, 64, 65, 271. 
Goette, 43, 231. 

Gonads, 280, 284, 291; arteries of, 208; 

mesenteries of, 268; veins of, 227. See also 

Genitalia, Ovary, Testis. 
Goniale {see Pre-articular), 33, 34. 
GopPERT, 19, 49, 63, 64, 267, 273. 
Graham Kerr, 130. 
Gravenhorst, XX, xxi, xxvi. 
Gray, 18, 277, 278. 
Gregg Wilson, 281, 282. 
Greil, 185, 190, 250, 260. 
Grey matter, 130, 132. 
Grodzinski, 235, 251, 254. 
Gronroos, 5. 
Gubernaculum cordis, 194. 
Gular fold, 11, 58, 59; region, 13. 
Gut, xvi, XX, xxii, xxvi, xxix, 264-6; relief 

pattern of, 264. See also Alimentary canal. 

Habenular commissure, 123, 128-9. 

— ganglion, 123. 
Haberlandt, 186. 
Habits, 8, 9. 
Haemal arch, 20. 



352 

Hall, 282. 

Haller, 123. 

Hand, pronation of, 90. 

Harderian gland, 301-2. 

Haro, xxviii. 

Harrison, 126, 296-300. 

Hasse, 125, 126, 296, 297, 298, 299. 

Head, 11; arteries to, 197-204; lymphatics 
of, 258; muscles of, 49-69; nerves of, 132- 
56; veins from, 220, 221-7. 

Heart, course of blood through, 192; descrip- 
tion of, 186-91 ; external features of, 186; 
historical, xv, xvi, xviii, xix, xxi, xxii, xxiv, 
XXV, xxvi, xxvii, xxix, 185; internal details 
of, 187; nerves of, 155-6; 186; physiology 
of, 185-6; 191, 192-3; wax-model of, 186. 

Heidenhain, 182, 183, 286, 287. 

Helminth parasites, 9, 10. 

Heloderma, 244. 

Henle, xxviii, 63, 64, 274. 

Hepatic artery, dorsal, 207; ventral, 207. 

— duct(s), xvi, 267. 

— portal system, 220, 235-9. 

— veins, 228-9. 
Hepato-cystic (bile) duct, 267. 
Hermann, xvii. 
Hermaphrodite, xvi, 285. 
Herrick, 303. 

Hertwig, 262. 

Hibernation, 8. 

Hind-limb, 45-7; arteries of, 214-18; com- 
pared with fore-limb, 171, 173, 174, 177; 
muscles of, 98-122; nerves of, 170-9; 
sympathetic nerve to, 183; veins of, 240-3. 

Hinsberg, 303. 

Hochstetter, 194, 206, 210, 228-31, 235, 
238, 239. 

Hoffmann, xv. 

— C. K., ID, 17, 30, 32, 37, 42, 45, 52-5, 
57, 60, 65, 68-91, 94, 96, 99-112, 114- 
21, 133, 135, 137-43, 146-8, 150, 156-7, 
159, 165-6, 171-3, 176-9, 184, 194, 198. 

Hohlvene Fortsatz, 228. See also Post-caval 

lobe. 
Horizontal (semicircular) canal, 24, 297; 

ampulla of, 149. 
HoYER, 252; and Udziela, 250, 252, 254-7, 

260. 
Humerus, 38, 40-2; condyles of, 41, 81-3, 

85-7. 
Humphrey, 284-5. 
Humphry, 48, 57, 65, 67, 69, 71-88, 90-1, 

96, 99-105, 107-12, 114, 1 16-21, 165-6, 

171-2, 176-8. 
Hunter, John, xviii, xxiv. 
Hunterian museum, xviii. 



INDEX 



HUSCHKE, xxiv. 

Hynobius leechi, 295. 

Hyo-branchial apparatus, 34; functional 

mechanism of, 6r; muscles of, xxvi, 57 et 

seq. 

— skeleton, xxi, xxvi, 34, 35. 

Hyoid, arch, xix, 34-5; cleft, 147-8; muscles 

of, xix, xxvi. 
Hyoidean sinus (of Selachii), 227. 

— (lymphatic) sinus, 259. 
Hyomandibular cleft, 147-8. 

— nerve, 145. See also N. facialis, truncus 
hyomandibularis. 

Hypobranchial elements, 35-7. 

— nerve, 161. 

Hypochordal commissure, 25. 
Hypo-glossal nerve, 66, 67, 71, 132, 157, 159, 

160, 161; anastomosis with vagus, 154, 
160; homology of, 161; relation to sub- 
clavian vein, 233. 

Hypo-hyal, 34-7. 

Hypophysis cerebri, 123, 125-7, 130. See 
also Pituitary body. 

Hyrtl, 17, 206-7, 228. 

Ichthy aphis, 123. 
Ilia(um), 20, 44, 45, 
Iliac artery, 214-18. 

— (sympathetic) ganglia, 183. 

— (lymphatic) sinus, 253. 
Imperato, xiii, xiv, xv. 

Impregnation (of the female), 4. See also 

Fertilization. 
Incisura coracoidea, 38. 
Inferior dental foramen, 146. 

— jugular sinus (of Selachian), 227. 

— median plexus (of diencephalon), 245, 248 

— palpebral nerve, 142. 
Infundibulum, 123, 126-7, 130, 131. 
Inguinal (lymphatic) sinus, 253. 
Inspiration, 276. 

Inter-auricular septum, 185, 188. See also 

Septum atriorum. 
Intercostal arteries, 205. 
Interdural space, 124-8. 
Intermaxillaries, 27. See also Pre-maxilla. 
Intermaxillary gland, 24, 263. 
Intermedium, 46, 47. See also p. 43. 
Internal carotid artery, 30, 198-201. 

— jugular vein, 219, 220, 221, 222-5. 

— mandibular nerve, 143-4, 146. 

— nares (or narial opening), xx, 22, 31, 145, 
264, 304. See also Choanae. 

Internasal septum, 24. 
Intervertebral foramen, 17, 19. 

— cartilage, 17. 



INDEX 



353 



Intestine, 265; arteries to, 207-8; lymphatics 
of, 255; relief pattern of, 265; veins from, 
238. See also Alimentary canal, Gut. 

Iris, 300, 301. 

Ischial spine, 44. 

Ischium, 44, 45. 

Investing bones, 22; of lower jaw, 33-4; of 
skull, 27-32. 

Jacobaeus, xiii, XV. 

Jacobshagen, 8, 265. 

Jacobson, xxi. 

Jacobson's anastomosis, 145. 

— gland (organ), 134, 302-4. 

— veins, 196, 239, 244. 
Jaquet, 179. 

Jaw(s), 21; articulation of, 31, 33; lower, 32- 

4; muscles of, 49; upper, 27-8. 
Johnson, 56, 300-2. 
Joseph, 228. 
jourdain, 194. 
Jugal ligaments, 27-8. 

— tendon, 21. 

Jugular (lymphatic) sinus, 259. 

— vein, external, 219, 220, 223, 224, 225, 
227; internal, 219, 220, 221, 224. 

Kallius, 36, 263. 

KaMMERER, 5, 12. 

Kaufman, 6, 7. 

Kehrer, 42. 

Keith, 270. 

Kesteven, 30, 31. 

Kidneys, xvii, 268, 278; of female, 279; of 

male, 283; arteries of, 208; definitive, 278; 

development of, 278; sexual, 278; veins of, 

227, 239. 
Kingsbury, 264; and Reed, 71, 294. 
Kingsley, 133, 142. 
Klaatsch, 268. 
Kleeman, xvii. 
Klingelhoffer, 300. 
Knoblauch, 13. 
Konigstein, 195. 
Kuhn, 296, 297, 298, 299. 

KUPFFER, 5. 

Kurdistan, 2. 

KUREPINA, 303. 

Labial glands, 262. 

Labyrinth, 24, 25; membranous, 296-8. 

Lacerta salamandra, 3 . 

Lacrimal gland, 301-2. 

Lagena, 150, 297, 298; sensory area of, 299. 

Lamina terminalis, 131. 

Langer, 185. 

Langerhans, 185. 

4038 . 



Lapage, 29. 

Larva(ae), xiv, xv, xvi, xxi, xxii, xxiv, xxviii, 
xxix, xxxi, 3, 7, 22, 30, 33-6; action of cilia 
in respiration of, xxviii; cranial nerves of, 
133; I5°> 160; dead, 6; deposition of, 6; 
food of, 7; growth of, 7; lymphatics of, 
250, 254, 255, 257; metamorphosis of, 7; 
monstrous, 7; number of, 6. 

Laryngeal artery, 218. 

— cartilages, 36, 271; homology of, 272. 

— muscles, 49, 62-5. 
Laryngo-tracheal chamber, 271. 

Larynx, xxviii, 36, 271; artery of, 218; 

homology of, 272; muscles of, 49, 62-5; 

nerves of, 154-5. 
Lateral glands (of pharynx), 264. 

— line organs, xxxi. 

— lymph hearts, 249, 253, 255, 256, 259-60. 

— palatine nerve, 145. 

— plexus (of fore-brain), 131. 

(of sinus Endolymphaticus), 247, 249. 

— (lymphatic) plexus, 251. 

— vein, 230, 231, 232; connexion with 
lymphatic system, 232, 253, 255; relation 
to V. iliaca transversa, 244. 

Lateralis component, 151; system, 139. 

Latreille, xvi, xviii, xx. 

Laurenti, xvii, 3, 13. 

La Vallette St. George, 285. 

Leeuwen, van, 5. 

Leg, 46. See also Hind-limb. 

Legends, xiii. 

Length of life, 8. 

Lens (of eye), 300. 

Leuckart, xiv. 

Leydig, xxi, xxiii, xxix, xxxi, 277, 278, 282- 

3, 288, 292. 
Lieno-gastric artery, 206. 

— vein, 238. 
Life, length of, 8. 
Life-cycle, xxiii. 
Life-history, 3, 4-9. 
Ligaments, cotyloid, 45. 
--jugal, 27-8. 

Ligamentum coronarium hepatis, 269, 281. 

— denticulatum, 127. 

— dorso-laryngeum, 64, 65. 

— gastro-lienale, 255, 267. See also p. 268. 

— hepato-cavo-pulmonale, 269, 272. 

— hepato-entericum, 267, 268-9. 

— hyo-quadratum, 34, 57, 58, 61. 

— piae ventrale, 127. 

— quadrato-maxillaris externum, 27; in- 
ternum, 27. 

— suspensorium hepate, 269. 

— tubae, 269, 281. 



354 

Limb(s), xviii, xxii, xxiv, xxviii 



INDEX 



i; lymph- 
atics of, 252; movement of, 12; muscles 
of, xxii, 11; nerves of, 157. See also, Fore- 
limb, Hind-limb. 

Limb-girdles, xxii. See also Pectoral girdle, 
Pelvic girdle. 

Linea alba, 40, 45, 55, 95. 

Lingual vein, 219, 226. 

— nerve, 151. 
Linnaean species, 3. 
Linnaeus, 3. 

LiNSTOW, VON, 9, 10. 

Lips, 262; glands of, 2625 sores on, 9. 

List, 19. 

Liver, xvi, 266, 291; arteries to, 206-7; fat 

content of, 280; mesenteries of, 268-9; 

post-caval lobe of, 228, 238, 266; veins of, 

228. 
Lizard(s), xiv, xvi, xvii, xx, xxii, xxvi. 
Locality, 8, 9; influence on breeding habits, 

4. See also Geographical distribution. 
LoNNBERG, 3, 274. 
Low, 99, 100, 102-10. 
Lower jaw, 27, 32-4; nerves of, 143, 145. 
LUBOSCH, 49, 52, 53. 
Lung(s), xvi, xvii, xix, xxvi, xxviii, 45, 272, 

arteries to, 218; mesenteries of, 269, 272; 

nerves of, 155-6; veins of, 220, 235. 
Luther, 49, 52-5. 
Lymph heart(s), xxvi, xxvii, xxix, xxx, 249; 

relation to blood-vessels, 259. 
Lymph heart, central, 190, 250, 256, 260. 

lateral, 249, 253, 255, 256, 259-60. 

Lymphatic cistern, xxvii, 255, 256. 

— system (or lymphatics), xxv, xxvii, xxix, 
xxx, 249-59; caudal, 250; of the bladder, 
xxix, 253; of the head, 258; of the gut, 
xxix, 254; of the larva, 250, 254-5; of the 
limbs, 252; of the palate, 258; of the pec- 
toral region, 251, 256; of the pelvis, 251, 
252-3; of the pharynx, 258; of the tail, 
254; of the tongue, 258; of the trunk, 251, 
254, 256; subcutaneous net-work, 251. 

Lymphoid glands (of mouth and pharynx), 
264. 

MacBride, 13. 

McMuRRiCH, 70, 84-5, 87-91, 99, loi, 114- 

21, 157, 168. 
Macula acustica neglecta, 150, 299. 

recessus utriculi, 299. 

sacculi, 299. 

Maculae acusticae, 298; histology of, 299. 
Malbranc, xxxi. 
Malpighi, xvii, xix. 
Malpighian bodies, xv, 278, 283. 



Man, de, 99, 100, 102-5, i°7-9- 

Mandible, 33. See also Lower jaw. 

Mandibular arch, 32-4. 

— (lymphatic) sinus, 258. 

Mann, 301. 

Manno, 217-18. 

Marsupial bone, xx, 45. See also Ypsiloid 

cartilage. 
Martin Saint-Ange, xxvii. 
Mathes, 268. 
Mating, 4, 5, 6. 
Mattiolus, xiii. 

Maturity, age at which attained, 8. 
Maupertuis, xvi. 
Maurer, xxxi, 92, 94-7, 289, 290. 
Maxilla, 21, 27-32, 141. 
Mayerhofer, 19. 
Meckel, xxi, 69, 72-9, 81-2, 85, 88, 92, 99, 

100, 102-4, 107-8, 115. 
Meckel's cartilage, 33, 34, 53. 
Median epigastric vein, 230, 232, 237. 



[32, 138, 144, 



Medulla oblongata, 129, 130, 

150, 156; arteries of, 246. 
Mehlsheimer, 8. 
Membrana intercartilaginea, 35. 

— nictitans, 55, 302. 

— sterno-coracoidea, 40. 
Membranes, foetal, xiv, xv. 
Membranous labyrinth, 296-8. 
Meninges, 124-8. 

Meninx primitiva, 124, 128. 

— secundaria, 124, 128. 
Mental foramina, 144. 

— nerves, 144. 
Mento-mandibulare, 33. 
Mento-Meckelian, 33. 
Mercurochrome, use of, 9. 
Mercury, use of, 249, 254, 256. 
Mesencephalon, 129, 130, 135. See also Cor- 
pus opticum. Optic lobes. 

Mesenteries, 268-9. 
Mesial cerebral artery, 245. 
Mesocardial ligament, 194. 
Mesorchia(um), 268. 
Mesovaria(um), 268, 280 

256. 
Metacarpals, 40^ 

91- 

Metamorphosis, xvi, xxiv, xxvi, xxviii, xxix, 
xxxi, 7; factors influencing, 7, 8; of vis- 
ceral muscles, 48, 54, 57, 58. 

Metatarsals, 46, 47; muscles of, 117, 1 19-21. 

Meves, 284. 

Meyer, xxx, 249, 255, 256. 

Middle visceral (sympathetic) nerve, 182. 

Mihalkovics, 302. 



lymphatics in, 
43, 44; muscles of. 



INDEX 



355 



MlVART, 52-4, 57, 60, 62, 65-7, 69, 71-9, 
81-3, 85-8, 90, 96, 99-105, 107-10, 112, 
115. 

MiYAWAKr, 150. 
Molge 'vulgaris, 260. 
Monilia batrachea, 9. 
Monro, xiii; foramen of, 131. 
Monro secundus, xvii. 
Monsters, 7. 
MOOKERJEE, 17, 160. 

Mouth, 262; epithelium of, 263; lymphoid 
glands of, 264. 

— floor, lymphatics of, 258-9; oscillation of, 
274, 275. 

Mucus glands (of skin), 292, 293. 

MuLLER, Johannes, xxvi, xxvii, 249. 

Miiller's (or MuUerian) duct(s), xvii, xx, xxxi, 
280-2; arteries to, 208-10; development 
of, 281-2; external opening of, 281; in- 
ternal opening of, 269, 280; lymphatics 
of, 256; of male, 283-4; vascularization of, 
281; veins from, 229, 240, 281. 

Murray, 19. 

Muscles, arm, 75-81. 

— body-wall, xxii, 94-8; arteries of, 210-11. 

— eye, 49-51, 55-6; nerves of, 135. 

— fifth nerve, 51-6. 

— fore-limb, 75-92. 

— general, xxv, xxviii, 48, 121. 

— head, xxii, 48-69; supplied by spinal 
nerves, 65-9. 

— hind-limb, 98-122. 

— historical, xxii, xxv, xxviii, 48-9, 69-71, 
92, 98-9. 

— laryngeal, 49, 62-5. 

— ninth and tenth nerve, 60-5. 

— pelvic, 99-107. 

— seventh nerve, 56-60. 

— shoulder, 71-5. 

— trunk and tail, 92-8; arteries to, 214; 
lymphatics of, 251, 254, 256; veins of, 
229. 

Musculus abdominis rectus, 94. 

— abductor digiti V, 120. 

— abductor et extensor digiti primi, 83, 
174, 213. 

— anconaeus, 70, 79, 211, 233. 

coracoideus, 79, 80, 166. 

humeralis, lateralis, 80, 165, 

medialis, 80, 165-6. 
scapularis medialis, 79, 166. 

— basiradialis, 67. 

— caput longum musculorum contrahen- 
tium, of fore-limb, 88, 169; of hind-limb, 
117, 118, 178. 

— caudali-femoralis, 102, 103, 176, 216. 



ii3> 



lee-. 



Musculus caudali-pubo-lschio-tibialis, roo, 
loi, 102, 103, 176, 216. 

— caudo-pedalis, 10 1. 

— cephalo-dorso-subpharyngeus, 62, 64, 
153, 154; relation to aortae, 63. 

— constrictor laryngis, 64, 65, 154, 155, 272, 
273. 

naris, 304. 

Musculi contrahentes digitorum, of fore- 
limb, 89, 169; of hind-limb, 118. 

Musculus coraco-brachialis brevis, 71, 78, 
168. 

longus, 71, 78, 168. 

— coraco-radialis proprius, 76, 77, 163. 

— cucullaris, 62, 71, 72, 154, 212, 221, 233. 
minor, 72, 162. 

— deltoideus, 77. 

— depressor mandibulae, 56, 62, 149. 

— dilatator laryngis, 63, 65, 154, 272, 273, 
274. 

naris, 304; accessorius, 304. 

— dorsalisscapulae, 70, 73, 77, 164, 211, 233. 
trunci, 37, 92-3, 158. 

— dorso-humeralis, 70, 73, 164, 211, 233. 

— epischio-hyoideus, 94. 

— extensor antibrachii et carpi radialis, 41, 
81, 82, 165, 167; ulnaris, 41, 81, 82, 167. 

— extensor cruris et tarsi fibularis, 114, 173, 
174. 

— extensor cruris fibularis, no, tibialis, no, 
112, 172, i74._ 

digiti brevis superficialis II, III, IV, 

and V, 113. 

— extensor digitorum communis, of fore- 
limb, 41, 81, 166; of hind-limb, 110, in, 
174- 

— extensor ilio-tibialis, 46, 107, 108, 172, 
173, 214, 215, 243. 

lateralis digiti quarti, 84. 

tarsi tibialis, 111, 174. 

Musculi extensores digitorum breves pro- 
fundi, 113. 

breves digitorum, of fore-limb, 84, 165, 

166; of hind-limb, 112, 174. 

Musculus femoro-fibularis, 108, 109, 176, 
216. 

— Uexor accessorius lateralis, of fore-limb, 
85, 87, 169, 234; of hind-limb, 116, 119, 
178,217. 

medialis, of fore-limb, 85, 87, 90, 

169, 234; of hind-limb, 116, 118, 119, 178, 

217- 

antibrachii et carpi radialis, 85, 122, 

antibrachii ulnaris, 85, 87, 122, 169, 



356 



Musculus flexor carpi ulnaris, 86, 12 

cruris, caput ventxale, 99. 

primordialis communis, of fore-limb; 

84, 85, 87, 169, 212, 234; of hind-limb; 

114, 119, 122, 177, 178. 
Musculi flexores breves profundi, 118, 119. 
superficiales, of fore-limb, 85, 88 

169; of hind-limb, 116. 
digitorum minimi, of fore-limb, 91 

169; of hind-limb, 120. 
Musculus genio-glossus, 55,66, i6r, 198, 225 

— genio-hyoideus, 35, 36, 55, 59, 65, 66, 68 
161. 

lateralis, 59, 60, 66. 

tertius, 66. 

— glutaeus maximus, 107. 

— gracilis, 99, 100. 

— hebosteoypsiloideus, 69. 

— humero-antibrachialis, 71, 81, 168, 212. 

— hyoglossus, 61, 67, 161. 

— hyoideo-ypsiloideus, 94. 

— ilio-caudalis, 98. 

— ilio-extensorius, 107, 108. 

— ilio-femoralis, 106, 109, 173, 214. 

— ilio-fibularis, 108, 109, 173. 

— ilio-tibialis, 107, 108. 

— interhyoideus, 54, 55, 58-60, 
197, 224, 225. 

posterior, 58, 149, 224, 225. 

— intermandibularis, 54, 58, 59, 14 

anterior, 54; posterior, 54, 55 

Musculi intermetacarpales, 84, 91. 

— intermetatarsales, 121. 

Musculus interossesus antibrachii, 90, 169. 
cruris, 119, 178, 217. 

— interphalangeus digiti III, 91. 
Musculi interphalangei, 121. 

— interspinales, 18, 93. 

Musculus intertransversarius, 93, 158. 

capitis inferior, 94, 160; superior, 93. 

160. 

— ischio-caudalis, 102, 176. 

— ischio-femoralis, 104, 175. 

— ischio-flexorius, 100-1, 175, 
217. _ 

Musculi laryngei, 65. 
Musculus laryngeus ventralis, 65 

— latissimus dorsi, 73. 

— levator arcuum V, 273. 

bulbi, 49, 55, 137, 142, 223, 224; pars 

principalis, 55, 56; pars sagittalis, 55, 56; 
pars transversalis, 56. 

mandibulae, 27, 49, 51, 143, 200, 223. 

anterior, 52, 143, 223; superficial 

portion, 52; deep portion, 52. 

externus, 53, 143. 



INDEX 
69 



[44, 149, 



57, 58- 



176, 



273. 



Musculus levator mandibulae posterior, 53, 

143. 

scapulae, 71, 154, 295. 

Musculi levatores arcuum branchiarum, 62, 

63, 64. 
Musculus longissimus capitis, 93. 

— mylohyoideus, 54, 57. 

— obliquus externus, 96. 

profundus, 97 ; superficialis, 76, 

96, 158. 

inferior, 51, 55. 

internus, 97, 158, 163, 194. 

superior, 50, 136, 137. 

— occipitalis, 93. 

— opercularis, xxiv, 24, 37, 71, 160, 221, 
233; function of, 295-6. 

— pectineus, 105. 

— pectoralis, 40, 41, 58, 71, 75, 76, 95, 
211. 

— pectori-scapularis, 68, 69, 161, 162. 

— plantaris superficialis medialis, 10 1. 

— procoraco-humeralis, 58, 70, 71, 74, 75, 
77, 163, 164, 212. 

— pronator manus, 87, 88. 
pedis, 118. 

profundus, of fore-limb, 87, 88, 89-90, 

169; of hind-limb, 118-19, 178, 217. 

— pubi-femoralis, 105, 106, 175, 216. 

— pubo-extensorius, 106. 

— pubo-ischio-femoralis externus, 99, 103, 
106, 171, 175, 215, 216. 

internus, 105, 106, 107, 1 10, 171, 172, 

215, 216. 

— pubo-ischio-tibialis, 99, loi, 102, 103, 
175, 215, 216, 217. 

— pubo-tibialis, 100, 105, 108, 175, 216. 

— pyramidalis, 94, 96. 

— pyriformis, 103. 

— rectus abdominis, 45, 94, 163, 164, 194; 
relation to diaphragm, 270. 

profundus, 68, 95, 158; super- 
ficialis, 40, 68, 76, 94, 96, 158. 

anterior, 50, 201. 

capitatis posterior, 93, 160. 

cervicis, 61, 67, 161, 162, 197. 

profundus, 35, 67, 68; super- 
ficialis, 35, 40, 68, 95. 

femoris, 99, 107. 

inferior, 50, 55, 135, 201. 

posterior, 50, 138, 200. 

superior, 49, 132, 135, 201. 

— retractor bulbi, 31, 50, 51, 135, 136, 138; 
200, 301, 302. 

— sartorius, 100. 

— serratus magnus, 74. 

— scapulare-cutaneus, 72. 



INDEX 
60, 6 



357 



Musculus subarcualis rectus I, 35 
152, 153, 155, 198. 

— subhyoideus, 35, 59, 60, 61, 149, 198; 
homology of, 59, 6G. 

— subscapularis, 41, 78. 

— subvertebralis, 97. 

— supinator pedis, in, 113. 

— supracoracoideus, 41, 58, 71, 75, 76, 163. 

— thoraci-scapularis, 37, 71, 74, 162, 163, 
164, 232. 

— tibialis anticus, 112. 

— transversalis, 97 ; relation to diaphragm, 
270. 

— transversus ventralis, 273; IV, 63, 154; 
V, 273. 

— trapezius, 72. 

— ypsiloideus anterior, 95. 

posterior, 94, 96. 

Myelencephalon, 129. See also Medulla 

oblongata. 
Mylohyoid muscle, 57. 
Myosepta, 11, 92. 

Nakano, 186. 

Nares, external, 11, 23, 28, 29; opening and 
closing of, 303. 

— internal (or posterior), xx, 22, 31, 145, 
264, 304. See also Choanae. 

Nasal (bone), 23, 27, 28, 29, 141. 

— capsules, 22-4, 27, 28, 29, 141, 145, 302; 
glands of, 304; veins of, 222. See also Olfac- 
tory capsules. 

— gland, 304; organ, 302. 
Naso-lacrimal duct, 11, 29, 141, 302, 304. 
Navratil, von, 303. 

Neck, 11; muscles of, 93-4; nerve plexus of, 

162; vertebra of, 16, 17. 
Necturus, 20, 92, 167, 228, 295. 
Nematode parasites, 9. 
Nematoxys commutatus (in gut), 9, 10. 
Nephrostomes, 278. 
Nerves, body-wall, 157-9, 170. 

— cardiac, 155, 156. 

— ciliary, 140. 

— cranial, 133, 134-56. 

— eye-muscle, 135-8. 

— gastric, 156. 

— general, 132-4. 

— mental, 144. 

— of fore-limb, 163-70. 

— of hind-limb, 170-9. 

— pharyngeal, 150-4. 

— pulmonary, 155-6. 

— spinal, 156-79. 

— spino-occipital, 156. 

— sympathetic, 179-84. 



Nerves, visceral, 133, 148, 150-4. 
Nervous system, xvii, xxix, xxx, 123-84. 
Nervus abducens, xxx, 50, 51, 130, 138. 

— accessorius X (Willisii), 72, 15 1-2, 154. 

— acusticus, 25, 149; endings of, 298-9; r. 
anterior, 149, 299; r. medianus, 149, 150; r. 
posterior, 150,299. S'^^'a/^o Auditory nerve. 

— axillaris, 164. 

Nervi brachiales longi superiores, 164. 
Nervus brachialis, 167; r. profundus, 167, 
168;r. superficialis, 78, 81, 86, 167, 168. 

— brachialis longus inferior, 167. 

— cardiacus anterior, 155; posterior, 156. 
Nervi coraco-brachiales, 78, 168. 
Nervus cruralis, 171 ; anterior, 171. 

Nervi cutanei brachii superiores laterales, 164. 

— cutanei cervicales, 162. 

occipitales anterioreset posteriores, 151. 

Nervus cutaneus abdominis lateralis, 158; 

medialis, 158. 

antibrachii dorsalis, 165. 

brachii dorsalis, 166. 

inferior lateralis, 168; medialis, 168. 

superior lateralis, 164; medialis, 

167; r. axillaris, 167. 

ventralis, 170. 

dorsi lateralis, 158; medialis, 158. 

femoralis posterior, 176. 

ventralis, 175, 177, 178. 

femoris internus, 172. 

medius, 171. 

surae lateralis, 173; medialis, 176. 

Nervi digitales dorsales, 172, 174, 175; ven- 

trales, 170, 175, 178. 

— dorsales pedis, no. 

Nervus dorsalis humeralis, 74, 164. 

manus intermedius, 83, 84, 165. 

radialis, 165. 

ulnaris, 82, 84, 166, 167. 

pedis fibularis, 114, 174; r. acces- 
sorius, 174, 175. 

intermedius, in, 112, 114, 173—4. 

tibialis, 112, 172. 

scapulae, 73, 77, 163. 

— extensorius, 108, 109, no, 173. 

caudalis, 79, 82, 83, 166 ; relation to N. 

extensorius cranialis, 167. 
cranialis, 80, 82, 165, 166; relation to 

N. extensorius caudalis, 167. 

— facialis, xxx, 26, 34, 144 et seq., 150. 
r. alveolaris, 34, 144, 146; homology 

of, 146-8. 

r. buccalis, 139, 142. 

r. jugularis, 148. 

r. mandibularis externus, 139, 148; 

internus, 147. 



INDEX 

ophthalmicus superficialis; 
144, 145, 148 



142, 



116, 



i49> 



Nervus facialis 

139- 

r. palatinus, 14 

r. pre-trematicus, 148. 

truncus hyomandibularis, 145. 

— femoralis, 105, 107, 171, 172, 173. 

— iibularis, 120, 177; r. medianus, 115 
118, 177, 178. 

— gastricus, 156. 
dorsalis, 156; ventralis, 156. 

— glossopharyngeus, xxx, 26, 60, 61, 
150. 

r. communicans ad VII, 148, 149, 150. 

r. post-trematicus, 151, rr. cutanei 

jugulares, 151; rr. pharyngei, 151; r. 

lingualis, 151. 
r. pre-trematicus, 150; r. pharyngeus, 

150. 

— hypobranchialis, 161. 

— hypoglossus, 66, 67, 71, 132, 157, 159, 
160, 161; anastomosis with vagus, 154, 
160; homology of, 161; relation to sub- 
clavian vein, 233. 

— ileo-hypogastricus, 96, 171. 

— interosseus, of fore-limb, 89, 90, 91, 92, 
169; of hind-limb, 119, 175, 177, 178. 

— intestino-accessorius X, 63, 64, 153. 

— ischiadicus, 102, 104, 109, 176-7. See also 
N. sciaticus. 

ventralis, 100, 102, 104, 106, 173, 175. 

— jugularis VII, 57, 59, 60, 148. 

— laryngeus dorsalis, 155; ventralis, 154. 

— lingualis, 151. 

— mandibularis, 143. 

externus, 143; internus, 143-4, ^A^- 

— maxillaris, 142; homology of, 143. 

— obturatorius, 44, 104, 105, 171. 

— oculomotorius, 50, 51, 132, 133, 135, 139, 

— olfactorius, 23, 129, 134. 

— ophthalmicus profundus, 26, 28, 133, 137, 
139, 145; homology of, 143. 

— opticus, 24, 130, 134. 

— palatinus, 133, 141, 142, 144, 145, 148; 
lateral branch of, 145. 

— patheticus, see N. trochlearis. 

— pectoralis, 76, 167, 170. 

— perinealis, 176. 

— peroneus, no, 114, 173, 177. 

— post-choanalis, 145. 

— post-palatinus, 145. 

— pudendus, 102, 103, 176, 179. 

— pulmonalis, 155. 

anterior, 156. 

lateralis, 155. 

— recurrens intestinalis X, 65, 152, 154. 

— saphenus, 112, 172. 



Nervus sciaticus, 102, 104, 109, 176-7 ; anoma- 
lous condition, 177. 5«'^(2/i-o N. ischiadicus. 

— spinalis i, 156, 158, 159 et seq.; anasto- 
mosis with vagus, 154, 160; anastomosis 
with N. spinalis 2, 161, 162; ganglion of, 
160; morphology of, 159-60. See also 
N. hypoglossus. 

— spinalis 2, 159, 160, 161; anastomosis 
with N. spinalis i, 162; anastomosis with 
N. spinalis 3, 162. 

— spinalis 3, 162, 163, 165; anastomosis 
with N. spinalis 2, 162. 

— spinalis 4, 163, 164, 167. 

— spinalis 5, 170. 

— spinalis 10, 157; ganglion of, 157; r. dor- 
salis, 157; r. ventralis, 158. 

— spinalis 14, 96, 171. 

— spinalis 15, 96, 170, 171. 

— spinalis 16, 171, 173. 

— spinalis 17, 173, 175-6. 

— spinalis 18, 176, 178. 

— spinalis 19, 179. 

Nervi spino-occipitales, 156. 
Nervus suboccipitalis, 159-60. 

— subscapularis, 79, 163. 

— supracoracoideus, 38, 77, 162, 163, 171. 
Nervi thoracici superiores, 3, 163. 

Nervus thoracicus inferior 2 anterior, 16 r; 

posterior, 162. 
superior 2, 72, 161, 162. 

— tibialis, 176. 

— trigeminus, xxx, 34, 126, 138 et seq. 
rr. frontales et palpebrales superiores 

anteriores, 140. 

r. mandibularis, 143. 

r. maxillaris, 133, 142. 

r. opthalmicus profundus, 26, 28, 133, 

137, 139, 145; homology of, 143. 
r. temporalis superior, 139. 

— trochlearis, xxx, 29, 51, 129, 135, 136. 

— ulnaris, 85, 86, 87, 89, 91, 92, 168, 169, 
170; r. medianus, 88, 89, 169. 

— vagus, xxx, 132, 150, 151; anastomosis 
with N. hypoglossus, 154, 160. 

r. communicans ad VII, 148, 149. 

rr. cutanei jugulares, 152, 153. 

fifth visceral arch nerve, 153. 

fourth visceral arch nerve, 152; r. re- 
currens, 152. 

r. laryngeus ventralis, 154. 

rr. musculares, 152, 153, 154, 155. 

rr. pharyngei, 152, 153, 154. 

rr. recurrens, 152, 153, 154. 

r. recurrens intestinalis, 152, 154; dis- 
section of, 155. 

r. intestino-accessorius, 63, 64, 153. 



INDEX 



359 



Nesterov, 2. 

Neural arch, i6, i8; of atlas vertebra, i6j 
foramina in, 17. 

— canal, 18, 19, 132. 

— spine, 16, 18. 
Neuromasts, xxxi. 

Newt(s), xiii et seq. I. See also Triton. 

Nicolas, 285. 

Nictitating membrane, 55, 302. 

NiERENSTEIN, 292, 293. 

NlSHI, 92. 

Noble, i, 10, 24, 108, 191, 192, 281, 290. 

Nodus chorioideus, 128, 131, 245, 246, 248. 
See also Paraphysis. 

Nomenclature, priority of, 3. 

NoRRis, 145. 

Nose, 302-4. See also Nasal capsules. Olfac- 
tory organ. 

Nostril, xxi. See also Nares. 

Obturator foramen, 44. 

— muscle, 104. 

— nerve, 44, 104, 105, 171. 
Occipital condyles, 26; facets for, 17. 

— region, 21, 25; nerves of, 156; veins of, 
221. 

— ring, 22, 26. 

— segment, 26. 
Occipito-petrosal, 26. 
Occurrence (of Salamanders), 8. 
Oculomotor foramen, 24, 135. 

— nerve, 50, 51, 132, 133, 135, 139. 
O'DoNOGHUE, 200, 217, 237, 239, 244. 
Odontoid process (of Atlas vertebra), 17. 
Oesophagus, 264; arteries of, 218. 
Okajima, 150. 

Olecranon fossa, 41. 

— process, 42, 79. 

Olfactory capsules, 22-4, 27, 28, 29, 141, 
145, 302; glands of, 304. See also Nasal 
capsules. 

— chamber, 302. 

— lobes, 128, 131. 

— nerve, 23, 129, 134. 

— organ, 302. 

— passages, 302. 
O'Neill, 124-8, 296, 299. 
Operculare, 33. See also Splenial. 
Operculum, xvii, xxi, xxiv, xxvi, xxvii, 24, 

26, 32, 71, 294, 295, 299. 
Optic fenestra (or foramen), 24, 28, 134. 

— nerve, 24, 130, 134. 

— peduncle, 134, 301. 

Orbit, 23, 24, 28, 31, 32; arteries of, 199, 
200-1, 203-4; glands of, 301-2; lymphatics 
of, 258; rim of, 21; veins of, 222, 225. 



Orbital sinus (of Selachian), 227. 

Orbito-sphenoid, 24, 25, 29, 32. 

Orbito-temporal region, 24. 

Oro-nasal groove, 303. 

Os transversum, 31. 

Os triangulare, 20, 34, 35, 37, 38, 65, 66. 

OSAWA, 10, 48, 52-4, 56, 60, 62-3, 65, 67, 
69, 71-3, 76-91, 99, 100, 102-5, i07-i4> 
1 16-21, 164-6, 171-2, 176, 188, 200, 202, 
206-18, 222, 230-3, 240, 242-5. 

Osborne, 131. 

Ostia(um) tubae, 269, 280. See also under 
Oviduct. 

Ostium venosum sinus, 187. 

Otic capsules, 22, 24-6. See also Auditory 
capsules. 

— ganglion, 146. 
Otolith(s), xvii, xxv, 298, 300. 
Ova, xiv, XV, 280. See also Eggs. 
Ovary(ies), xv, xvi, xxii, xxiii, 280; arteries 

of, 208; mesenteries of, 268; veins of, 227. 

Oviduct(s), XV, xvi, xxii, xxiv, xxviii, xxix, 4, 
280-2; arteries of, 208-10; development 
of, 28 1-2; external opening of, 281; internal 
opening of, xvi, xxii, 269, 280; lymphatics 
of, 256; vascularization of, 281; veins of, 
229, 239-40, 281. See also Miiller's duct. 

Ovisac, 280. 

Owen, xiii, 48, 51-4, 57, 67, 71-6, 8r, 94, 
99, 100, 107. 

Oxysoma bre^icaudatum (in gut), 9, 10. 

Oyarzun, 123. 

Pachytriton, i. 

Pairing, description of, 5. 

Palate, arteries to, 203-4; glands of, 263; 

lymphatics of, 258; nerves of, 144-5; veins 

of, 221, 223-4. 
Palatine (bone), 31. 

— foramen, 144. 

— ganglion, 141, 145, 183. 

— glands, 263. 

— nerve, 133, 142, 144, 145, 148. 
Palato-nasal artery, 203. 
Palato-quadrate, 22, 25, 26-7. 

Pancreas, xvi, 266, 267, 291; arteries to, 
206-7; ducts of, 265, 267; veins of, 238. 

Pancreatic ducts, 265, 267. 

Panizza, xxvi, xxvii, xxviii, xxix, xxx, 249, 
252^ 254-7. 

Parachordal plate, 25-6. 

Paraphysis, 125, 127, 128, 129, 131. See also 
Nodus chorioideus. 

Parasites, 9, 10. 

Parasphenoid, 26, 27, 30, 32. 

Parathyroids, 290. 



360 

Paratoid glands, 



INDEX 



I, i3> 



292; arteries to, 



Paratre, 4, 5. 

Par^, xiii. 

Parepigastric (lymphatic) trunk, 253. 

Parietal(s), 27, 29, 32. 

Parker, 17, 20, 21, 24, 28, 29, 31, 33, 34, 37, 

38, 39, 40. 
Pars basilaris (of labyrinth), 296, 297, 298; 

relation to perilymph, 300; sensory area of, 

299. 

— initialis (of labyrinth), 296. 

— neglecta (of labyrinth), 296, 297, 298 ; 
relation to perilymph, 300. 

— posterior utriculi, 297. 

— superior utriculi, 297. 
Pathetic nerve, see N. trochlearis. 
Pectoral girdle, xxviii, 37-40 ; attachment of, 

75. 5"^^ a/so Shoulder-girdle. 

— nerve, 76, 167, 170. 

Pelvic girdle, xix, 44-5; muscles arising 
from, loo-i, 103-10. 

— gland, xxiii, 287. 

— (lymphatic) sinus, 253. 

— veins, xxvii, 220, 244. 

Pelvis, lymphatics of, 251, 252, 253; muscles 
of, 99-109; skeleton of, 44-5. 

Penis(es), xvi, 286, 287, 288. 

Peri-anal (lymphatic) sinus, 253, 254. 

Pericardial cavity, 193. 

Pericardio-peritoneal canal, 193. 

Pericardium, 40, 68, 95, 97, 193-4; attach- 
ment of ventricle to, 193; relation to dia- 
phragm, 270; relation to mesenteries, 
269-70. 

Perilymph, 297, 299 ; function of, 300. 

Perilymphatic tissue, 297. 

Peri-orbital network, 222, 223. 

— (lymphatic) sinus, 258. 
Periotic, 30. 
Peritoneum, 269. 
Perrault, xiii, xvi. 

Perrin, 37-40, 42,44, 68, 70-5, 77-91, 98- 
114, 1 16-21; criticism of, 105-6. 

Perso-Turkey, 2. 

Pes anserinus, 149. 

Peter, 17, 159. 

Pfitzner, 292. 

Phalanges, of hand, 40, 44; of foot, 46, 47. 

Pharyngeal plexus, nerve, 151, 152; venous, 
224, 230. 

Pharynx, 38, 264; arteries of, 218; epithelium 
of, 263; lymphatics of, 258; lymphoid 
glands of, 264; nerves of, 15 1-2; veins 
of, 226, 230. 

Phylogeny of thigh muscles, 104. 



Pia mater, 124, 128. See also Primary vas- 
cular coat. 

Pineal body (or organ), xxxi, 129, 131, 291. 

Pituitary body, 123, 125-7, 130, 131, 291; 
artery of, 245; veins of, 248. 

Plantar aponeurosis, 10 1. 

Planum conchale, 23. 

— internasale, 23, 25. 

Plessen, VON and Rabinowicz, 52-4, 132-3, 

136, 138-9, 142-3, 146, 156. 
Plethodon, 19, 293. 
Plethodontidae, i. 
Pleural ligament, 269. 

— mesenteries, 272. 
Pleurodeles, i. 

Pleuro-peritoneal membrane, 269. 
Plexus iliacus (sympathetic), 182. 
Plica gastro-lienalis, 268. 

— hyomandibularis, 147. 

— recto-lienalis, 268. 
Puny, xiv. 

PoHL, xxi. 

Poison, xiv, xvi, 293; chemical nature of, 
293. See also Venom. 

— glands, II, 92, 292. See also Venom 
glands. 

Poisonous secretion, xiv, xvi, 293. 

PoUex, 42. 

Popliteal region, 10 1, 109. 

Portal vein, 220, 237. 

Post-branchial body, 290. 

Post-cardinal vein, xvii, 196, 220, 227, 229- 

30, 234; relation to anterior oviducal 

arteries, 210. 
Post-caval lobe (of liver), 228, 238, 266. 

— vein, 187, 196, 219, 220, 227-9; abnor- 
malities of, 228-9. 

Post-choanal nerve, 145. 
Posterior (semicircular) canal, 297; ampulla 
of, 150. 

— groin, 103, 242. 

— oviducal veins, 239, 240. 

— subclavian ganglion, 180. 
Post-orbital process, 29. 

— sinus (of Selachian), 227. 
Post-palatine nerve, 145. 
PoucHET, xxxi. 

Pre-articular, 33, 34, 146. See also Goniale. 

Pre-frontal, 23, 27, 28, 29. 

Preglottideal glands, 264. 

Pre-hallux, 47; cartilage of, iii, 112, 113. 

Pre-maxilla(ae), 27, 28, 29, 30. 

Pre-poUex, 43, 91. 

Pre-sacral vertebra, 18. 

Pre-sphenoid, 24. 

Pre-vomer, xx, 22, 27, 28, 30, 31, 32, 145. 



INDEX 



361 



Prevost, and Dumas, xxiv. 

Prey, seizing of, 12, 61. 

Primary cranium (or skull), 22-7, 31. 

— sub-dural space, 125, 127. 

— tongue, 263. 

— vascular coat, 124, 125, 127, 128. See 
also Pia mater. 

Primordial (or primary) skeleton, 22, et seq. 

Pro-atlas, 17. 

Process, olecranon, 42, 79. 

— pre-pubic, 44. 
Processi articulares, 18. 

Processus ascendens (of palato-quadrate), 26, 
142, 199. 

— basalis (of palato-quadrate), 26, 27, 31, 
145, 199. 

— coronoideus (of lower jaw), 34, 52, 53. 

— oticus (of palato-quadrate), 26, 30, 142, 
145, 199. 

— prenasalis inferior lateralis, 23. 

— pterygoideus, 27. 

— spinosus, 18. 
Pro-coracoid, 38, 39. 
Procter, Dr. Joan, 9. 

Profundus nerve, terminal branches of, 
140-1. See also N. ophthalmicus profun- 
dus. 

Pronephros, xxix; possible vestige of, 278-9. 

Prosencephalon, 128, 130, 134. See also 
Cerebral hemispheres. 

Prostate (s), xix, 288. 

Protozoan parasites, 9. 

Pterygoid, 27, 28, 30, 31, 32. 

— cartilage, 27, 31. 

Pubis(es), xxviii, 44, 45. See also Pelvic 

girdle. 
Pubo-ischium, 44, 45. See also Pelvic girdle. 
Pudendal nerve, 102, 103, 176, 179. 

— (lymphatic) trunk, 253. 

Pulmonary arch, 195, 197; branches of, 218. 

— arteries, xxv, 218. 

— nerve (s), 155-6. 

— respiration, 275-6. 

— system, capacity of, 192. 

— vein(s), xxv, 187, 188, 235; relation to 
sinus venosus, 235. 

Pulvinar, 129. 
Pylorus, xxvii, 265. 

Quadrate(s), 21, 27, 29, 30, 31, 34, 58. 
Quadrato-jugal, 27, 30, 32. 
Quadrato-mandibular joint, 30. 



Rabinowicz, von Plessen and, 52-4, 

136, 138-9, 142-3, 146, 156. 
Rabl, 43, 269, 270, 282. 



32-3. 



Radial condyle, 41. 

Radiale, 41-4. 

Radials (of copula), 35, 36, 37. 

Radius, 40, 41, 42, 43. 

Ramus communicans IX + X ad VII, 56, 

57- 
Rana, 21, 32, 231. See also Frog. 
Rath, 285. 

Rathke, xix, xxi, xxii, xxiii, 283. 
Rawitz, 285. 
Receptaculum seminis, 4, 5, 282; discovery 

of, xxi, xxiii, 277, 282; sperms wintering 

in, 5. 
Recessus dexter (of auricle), 188. 

— dorsalis (of auricle), 188. 

— intermedins (of auricle), 188. 

— maxillaris, 302. 

— partis basilaris, 300. 
neglecta, 299-300. 

— praeopticus (of brain), blood plexus of, 

245; 

— sinister (of auricle), 188. 

— utriculi, 149, 297-9. 
Rectal gland, 266. 

Rectum, 266; arteries of, 207, 215; lymph- 
atics of, 254, 255; veins of, 236, 238. 
Reed, 294; Kingsbury and, 71, 294. 
Regeneration, xviii, xxiv, xxvi. 
Reichert, xxviii. 
Renal arteries, 208. 

— organs, xvii. See also Kidneys. 

— (sympathetic) plexus, 182. 

— portal system, 220, 239—40. 

vein, xxi. See also Jacobson's vein. 

Renal (lymphatic) sinus, 254, 256. 
Reproductive habits, xx, xxvi, 4-7. 
Reptile(s), xviii, xix, xxvii, 69. See also 

Lizard. 
Respiration, xix, xxv, xxvii; bucco-pharyn- 

geal, 275-6; cutaneous, 274; of larva, 

xxviii; organs of, 271-2; physiology of, 

274-6; pulmonary, 275-6. 
Respiratory movements, 274, 275, 276. 
Retzius, 285, 295, 296, 297, 298, 299. 
Rex, 124-5, 195' 245-9- 
Ribbing, 42, 70, 81-2, 84-91, 99, 1 10-12, 

114-22, 157, 165-6, 168-9, 172. 
Ribs, xvii, xxi, 18, 19, 20; articulation of, 

18-19- 

ROESEL, xiii, xvii. 
Rome, 15. 
romeiser, 228. 
Rose, 185, 188. 
Rostrum prenasale, 24. 
RoTHiG, 123. 
Rubaschin, 123. 



362 INDEX 

RUDINGER, 69-79, 81-3, 85-6, 88-9, 91. 

RUGE, 54, 57, 58. 

RuscoNi, xxi, XXV, xxviii, xxx, 4, 5, 10, 36, 

48, 52-4, 56-8, 60, 65, 67, 69, 132, 194, 

210, 229, 238, 243, 249, 255. 
Rusconi's vein, 194, 238. 
Rylkoff, 70-81. 
Rymer Jones, 48, 81-2, 85-7, 94, 100, 

102-4, 107, 109-10, 114-15. 



Sabatier, 37, 39, 44, 45. 

Sacculus, 149, 297, 298, 299; relation to 

perilymph, 299, 300. 
Saccus copularis, 258, 259. 

— endolymphaticus, 125, 126, 128, 297, 
298; function of, 300; plexus of, 247, 
249. _ 

— perilymphaticus, 125, 126, 299. 
Sacrum, 19. 

Saint-Julien, xviii. 
Salamander's wool, xiii. 
Salamandra (genus), xvii, i. 

— (species), 3. 

— aquatica, xiii. 

— atra, xiv, xvii, xviii, xxiii, xxiv, xxviii, 
xxx, 2, 17, 36, 39, 160, 230, 254, 264. 

— caucasica, 2 . 

— maculosa, xxiii, xxiv, xxviii, xxx, xxxi, 2, 
21; nomenclature of, 2-3. 

— semeno'vi, 2. 
Salamandridae, i. 
Salamandrina, i, 274. 
Salamandroidea, i. 
Samandar, 3, 293. 

Samandaridin, 293; freezing-point of, 293. 

Samandarin sulphate, 293. 

Sardemann, 301. 

Scapula, 19, 37, 38; muscles of, 71-5. 

Scarpa, xviii. 

Schmidt, Goddard, and van de Hoeven, 

48, 52-4, 6^, 68, 71, 73, 75-6, 78, 81-2, 

85-6, 110-12, 114-15, 120. 
Schneider, xiv, 284. 
ScHOLB, 195, 199, 244, 245, 247. 
Schreiber, 7, 8, 9, 15. 
Schreibers, xxiii, xxiv, 4. 

SCHREITMiJLLER, 15. 

SCHUBERG, 292. 

SCHULTZE, C. A. S., xxi. 

SCHULZE, F. E., xxxi. 

Schwalbe, 2, 4, 135, 136, 281. 

schweitzerbarth, 15. 

Sciatic artery, 214, 215, 216, 217, 218. 

— nerve, 102, 104, 109, 176-7. 

— vein, 240, 241, 242. 
Scott, 9. 



Seba, xiii. 

Second hepatic duct, 267. 

Secondary tongue, 263. 

Segmental duct, xxix. 

Segmentation, xxix, 5. 

Seifert, 263. 

Selachian, sinuses of, 227. 

Semen, xxiii. See also Sperms. 

Semicircular canals, xxi, xxiv, 24, 25, 296, 
297. 

Semi-lunar valves, xxv. See also p. 189. 

Sense capsules, 22. See also Auditory cap- 
sule, Nasal capsule. 

— organs, xxv, 294—304. 
Seoane, 14. 
Septo-maxillary, 29. 

Septum atriorum, xxv, 187, 188. See also 
Interauricular septum. 

— accessorium, 190. 

— aortico-pulmonale, 190. 

— auricular, xxv, 185, 187, 188. See also 
Interauricular septum. Septum atriorum. 

— bulbi, 189; function of, 193; origin of, 
193; significance in respiration, 191— 3. 

— carotico-aorticum, 190. 

— caroticum, 190. 

— internasale, 24. 

— principale, 193. 

— pulmonale, 190. 

— semicirculare anterior, 25, 296; laterale, 
25; posterior, 25. 

— transversum, 270. 
Sewertzoff, 43. 

Sexes, distinction between, 12, 279, 286. 
Sexual kidney, 278; of female, 279; of male, 
283. 

— maturity, age of, 8. 
Seydel, 23, 302, 303, 304. 

Shoulder, lymphatics of, 257; muscles of, 71; 
skeleton of, 37. 

— girdle, 37. 

SiEBOLD, VON, xiv, xxi, xxiii, xxvi, xxxi, 4, 
37, 52-4, 56-60, 65-6, 68-9, 282, 285. 

SlEGLBAUR, 42, 70, 81, 83-4, 90, 99, 105, 
107-8, IIO-14, 157, 165-7, 169, 171-4, 
.178. 

Sinus anastomoticus (lymphatic), 257. 

— lateralis nasi, 302, 303. 

— lymphaticus, 259. 

— mandibularis, 258. 
profundus, 258, 259. 

— maxillaris, 303. 

— sagittalis, 246. 

— superior utriculi, 297. 

— thyroideus, 257. 

— venosus, 185, 187, 192, 219, 228. 



INDEX 



363 



-475 



Sinuses, of Selachian, 227. 

Sire don, 147. 

Siren, 145. 

Skeleton, xix, xx, xxii, xxviii, xxix, 

appendicular, 37-47; axial, 16-37. 
Skin, 292; as respiratory organ, 274; glands 

of, xvi, 11,292-3. 
SkulJ, XX, xxviii, 20-37; development of, 21; 

metamorphosis of, xxix; segmentation of, 

156. 
Smith, 59, 66. 
Snout, II, 13. 
Spanish peninsula, 13. 
Spatium interdurale endolymphaticum, 125. 

hypophyseos, 126. 

paraphyseos, 127. 

perilymphaticum, 126. 

— meningeale, 299. 

— sacculare, 298, 299. 

Spelerpes, xxix, 19, 59, 60, 66, 194, 274. 

Spengel, 277, 278, 284. 

Sperm(s), xxiii, xxviii, xxx, xxxi, 285; in 

receptaculum seminis, xxiii, 4. See also 

Spermatozoa. 
Spermatic arteries, 208. 
Spermatogenesis, xxx, 285. 
Spermatophore(s), 4, 5, 288. 
Spermatozoa(on), xxiii, xxiv, xxx, xxxi. 

also Sperm. 
Sphenethmoid, 24. 
Sphenodon, 200, 217, 244. 
Spinal cord, artery of, 246, anatomy of, 

membranes of, 127. 

— ganglia, 18, 157, 158, 161. 

— nerve(s), 19, 156-79; a typical, 157-8; 
connexions with sympathetic, 158, 161, 
179-82; ganglia of, 157, 158, 161. 

Spino-occipital muscles, 24. 

— nerves, 156, 180. 

Spiral valve, xxv, 189. See also Septum 

bulbi. 
Spleen, xv, 266, 267, 291; artery to, 206; 

mesentery of, 268; veins of, 238. 
Splenial (bone), 33, 34. 
Squamosal, 24, 27, 29-30, 32. 
Stadtmuller, 7, 21, 22, 25-31, 33-4, 144, 

150. 
Stannius, 37, 38, 54, 56, 60, 6^, 73, 75-9, 

82-7, 99-104, 107-9. 
Stapedial artery (of human embryo), 200. 
Steiner, 42, 43. 
Stellio, xiv. 
Steno, XV. 

Sternal plate, 39, 95. See also Sternum. 
Sterno-hyoid muscle(s), xix; xxviii, 67-8. 
Sternum, xxviii, 39-40 68, 95. 



See 



1325 



Sterzi, 124, 126, 128, 296. 

Stohr, 17. 

Stomach, xv, xvi, 264, 265; arteries to, 206; 
blood-supply, double nature of, 239; 
lymphatics of, 255; mesenteries of, 268-9; 
nerves of, 156; veins of, 238. 

Strasser, 42, 43. 

Strong, 133, 147. 

Strongylus auricularis (in gut), 9. 

Stuve, 281. 

Subclavian artery, 210-13; relation to post- 
cardinal veins, 230. 

— ganglion, anterior, 180; posterior, 180-1. 

— vein, 219, 220, 230, 233, 235. 
Sub-dural space, 128. 

Sub-hepatic mesentery, 267. See also Ventral 

mesentery. 
Sublingual glands, 264. 
Sub-maxillary (lymphatic) plexus, 251. 
Sub-pectoral (lymphatic) plexus, 251. 
Sub-pelvic (lymphatic) plexus, 251. 
Subscapular (lymphatic) sinus, 256. 
Subvertebral (lymphatic) sinus, 254, 255, 

256. 
SUCHARD, 186, 191, 272. 
Sulcus coronarius, 187. 

— lateralis, 94, 96. 

— longitudinalis dorsalis, 132. 

ventralis, 127, 132. 

SuMi, 123, 130. 

Superior median plexus (of diencephalon), 
245, 248. 

— palpebral nerve, 142. 
Supinator muscle, 82. 
Supra-pericardial body, 290. 
Supra-renal nodules, xv. See also Adrenal 

bodies. 
Supra-scapular, 24, 37 ; relation to auditory 

organ, 295. 
Suspensorium, 25, 26, 27, 32, 33. 
Suspensor-stapedial ligament, 295. 
SwAMMERDAM, XV, xvii, xxii, xxix. 
Sympathetic nerves, size of, 184. 

— ganglia, xxxi, 179, 180, 181; connexion 
with adrenal tissue, 179, 181. 

— nervous system, xxv, xxx, 135, 136, 146, 
151, 179—84; anterior subclavian ganglion 
of, 180; anterior visceral nerve, 181; branch 
entering antrum, 183; caudal portion, 180, 
183; cephalic portion, 180; 183-4; cervical 
and abdominal portions, 180-3; connexion 
with facial nerve, 183; connexion with 
vagus, 179, 183; cranial portion, 183; his- 
torical, 179; middle visceral nerve, 182; 
posterior subclavian ganglion, 180. 

Systema Naturae, 3. 



364 INDEX 

Systematic arch, 195, 196; branches of, 

201-5; cutaneous branch of, 201. 
Systolic rhythm, 185-6, 192. 

SZAMOYLENKO, 6o. 



Tadpole, xix. See also Larva. 

Tail, xxiv, 11; artery of, 20; length of, 12; 
lymphatics of, 250, 254; muscles of, 98, 
102; nerves of, 179; vein of, 20, 220; ver- 
tebrae of, 16, 20. 

Tarapini, 36. 

Tarsus, 42, 46, 47. 

Tectum internasale, 24. 

— synoticum, 25, 29. 

Teeth, xvi, 27, 28, 33, 262-3 ; nerves to, 144; 
pre-vomerine, i, 31, 62, 262. 

Telencephalon, 128. S^^ a/yo Cerebral hemi- 
spheres. 

Testis(es), xvi, xx, xxii, xxiii, xxvi, 284; 
arteries to, 208; formation of multiple 
lobes, 285; mesenteries of, 268; seasonal 
development of, 284-5; ^^'"s o^' ^^7- 

Thalamencephalon, 127, 128, 130, 134; 
cavity of, 131. 

Theis, 293. 

Thigh, 45, 46; muscles of, 99-110; phylo- 
geny of, 104-5. 

Third (aortic) arch, 197. 

Thoracic duct, xxvii. See also Subvertebral 
(lymphatic) sinus, p. 255. 

Thymus gland, 290 ; artery to, 201 ; veins of, 
225. 

Thyreoid gland, see Thyroid gland. 

Thyroid gland, 65, 289; artery of, 198; 
lymphatics of, 257; vein of, 219, 226. 

Tibia, 46, 47. 

Tibial crest (or spine), 46, 108, 112. 

— nerve, 176. 
Tibiale, 46, 47, in, 112. 
Toad(s), xxiv, xxvi. 
Todd, 48. 

Tongue, xvi, xxvi, i, 34, 66, 262-3; artery 
of, 198; development of, 263; extrusion of, 
61; glands of, 263; lymphatics of, 258; 
muscles of, 66-7; nerves of, 151; primary, 
263; secondary, 263; vein of, 266. 

Tonsils, 264. 

Tooth, 262-3. See also Teeth. 

TowNSON, xix, XX, xxi, 45. 

Trabecula(ae), 23, 25, 138. 

Tracheal cartilages, 271. 

Transverse processes, xxi, 18, 20. 

Trapezius muscle, 72. 

Trematode parasites, 9. 

Tretjakoff, 258. 

Tricoilia, xviii. 



Trigeminal ganglion, 133; connexion with 
sympathetic, 183. See also Gasserian 
ganglion. 

— nerve, xxx, 34, 126, 138 et seq. 
Trigeminus root, 129, 130. 

Triton, xiii et seq., 34, 39, 42, 108, 123, 125, 
138, 148, 159, 160, 169, 182, 194, 247, 
254, 260, 273, 274, 277, 286, 301, 303; 
development of ribs, 18; of kidney, 277-8. 

Triturus, i. See Triton. 

Trochanter, 46; external, 103. 

Trochlear groove, 41, 46. 

— nerve, xxx, 29, 51, 129, 135, 136; relation 
to N. trigeminus, 137. 

Trunci lymphatici longitudinales laterales, 

254, 255, 257, 260. 

parabdominales, 251, 254. 

parepigastrici, 254, 257. 

subvertebrales, 254, 255. 

Truncus arteriosus, xxv, 63, 66, 185, 186, 

189, 190, 193, 260; dexter, 187; sinister, 

187; structure of walls, 190. 

— iliacus, 253. 

— impar, 190. 

— intestino-accessorius X, 153. 

— lymphaticus circumflexus scapulae, 257. 
longitudinalis paravertebralis, 256. 

— peri-pharyngeus, 256, 257, 259, 260. 
Trunk, 11; lymphatics of, 251, 254, 256; 

muscles of, xxii, 92; veins of, 220, 227, 229, 

232, 239; vertebrae, 16, 18. 
Tylototriton, i. 
Tympanal areas, 300. 
Tympanic cavity, absence of, xxiv, 294.