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BY W. J. HEASMAN Senior Curator, University Museum of Zoology, Cambridge
Introduction. . Material. . External features. . The axial skeleton and nervous system . The alimentary system . . . The respiratory system . . . The vascular system . . . The excretory and reproductive systems . Discussion Behaviour. . . The line of fusion and specific differences Causes of the abnormality . Conclusions. . . . References . . .
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RECENT advances in experimental embryology have indicated that abnormal embryos may be induced to develop from normal eggs by any of a number of external conditions. While investigation of teratological phenomena may provide valuable clues for understanding normal processes there is a striking difference between the products of experimental embryology and the naturally occurring teratological specimens. The former do not appear to be capable of reaching a post-embryonic condition even under the favourable conditions possible in a laboratory, while examples of the latter category which have been recorded have usually reached this later stage and in a few instances have been sufficiently well adjusted to their environment to become mature. It is possible that this difference between artificially produced and naturally occurring specimens is due to selection from a much larger number of unobserved naturally occurring abnormal forms, but we are nevertheless faced with the problem of how individuals with unusual organisations are readjusted so that they will function under natural conditions. This is but part of the larger problem of adaptation, and before dealing with the problem of how this readjustment is brought about we do well to enquire what modifications are to be found in successful abnormal individuals. Snakes, with their lack of lateral appendages, provide relatively simple teratological examples, especially of double monsters, and as they do not occur frequently it is advisable to place on record whatever information can be obtained to facilitate comparison with any which may be produced under artificial conditions.
W. J. Heasman
Altogether since records have been kept, some seventy cases of doubleheaded snakes have been described, of which only seven have been dissected (Redi, 1684; Vsevolojsky, 1812; Dutrochet, 1829; Dorner, 1873; Borgert, 1896; Cantoni, 1921; Strohl, 1925). As long ago as 1684 Redi made as detailed an examination as has yet been recorded, and the much more recent paper by Strohl gives no more information about his new specimen than that the creature possessed two hearts, one more anterior than the other exhibiting situs inversus.
The present account is of a male double-headed specimen of Coluber (Zamenis) florulentus Schlegel, University Museum of Zoology, Cambridge, No. R 12960. It had been captured at Cairo and preserved in alcohol. The adult of this species reaches a length of just over 1 m. The total length of the preserved specimen was 25 cm., it was therefore probably quite young. Externalfeatures. No significant difference could be detected in the length or breadth of the two heads, but whilst one head undoubtedly followed the axis of the body the other projected at about 250 to the right of this axis and was rotated on the vertebral column until the angle between the ventral surfaces of the two heads was about 120°. That this was not an accident of fixation is indicated by the relationship of the vertebrae. The mouths and oesophagi of both sides appeared to be equally capable of
Ribs At the level of the second neck band, 23 mm. from the tip of each snout, the integument of the two "necks" united to form a common investAnal Scale ment. At this point there was a transverse and somewhat stretched fold of skin. Posterior to this fold the two parts were indicated by asymmetrical marking. A ventral groove extended for the whole length of the animal in the middle line, and that this was undoubtedly more than a phenomenon of fixation was shown by the fact that most of the ventral scales had an irregularity which Tet-ig. showing the ribs as seen scaling took the form of a posteriorly directed projection in a transparent preparation. tod the left of the groove. An enlarged diagram of the anal region shows the slight abnormality of the scaling (text-fig. 1). A Spalteholz transparent preparation showed that the anal scale was attached to two pairs of ribs and that the additional scale was without a rib attachment. At a distance of 2j cm. behind the fold of integument there was a pronounced
The Anatomy of a Double-headed Snake
bend in the body axis. The accompanying photographs make further description of the external features unnecessary (see Plate I, figs. 1 and la).
The axial skeleton and nervous system An X-ray photograph (Plate I, fig. 2) shows that the bend in the body axis occurred at the junction of the right "neck" with that of the main axis. The bend half-way along the right vertebral column took place where doubling was externally apparent and marked the point at which the right "neck" could move independently of the left. Counting from the common vertebrae to this point, it was possible to identify corresponding vertebrae in the two columns; in front correspondence failed as the left column was made up of twenty-four and the right of twenty-five vertebrae. The junction between the two columns extended over four vertebrae.
Text-fig. 2. Transverse sections of vertebrae close to the confluence of the two "necks." Nerve tissue stippled. Skeletal tissue cross hatched.
Serial sections of this region show that on passing backwards the approaching walls of the vertebrae first fused and then disappeared, leaving two separate spinal cords within one vertebra. The sagittal planes of these two cords were then inclined at rather less than 300. The right cord appeared to bend slightly towards the left and the two came into contact, the right cord slightly ventral to the left. Further backwards the central canals of the spinal cords moved towards one another, the left remaining dorsal to the right. Finally the right central canal turned upwards and ran into the left canal (text-fig. 2 a). Fusion of the two spinal cords occurred along a line drawn from the dorsal side running backwards and downwards through the central canals to the ventral side. Ventral to this line two spinal cords could be distinguished, dorsal to it a normal single spinal cord. A pair of abnormal ventral roots arose where the two spinal cords were closely applied to each other but anterior to the confluence of the two central
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canals. Before passing out from the vertebral canal these roots ran into a large mass of nerve tissue which connected the two sides (text-fig. 2 b). From this ganglion the nerves ran backwards and outwards through the vertebra to supply the dorsal musculature.
The alimentary system The jaws, mouths and alimentary canals of both sides were equally well developed and capable of functioning. Measurement of the diameters of the oesophagi in serial sections showed some variation along the length of each, but on the whole the right was slightly smaller than the left. Fusion of the dorsal mesentery 3 mm. behind the transverse fold of skin formed a transverse mesentery which sloped backwards and downwards and which was drawn out to form a partition between the two oesophagi. The alimentary canals did not join until the pylorus, and there were two stomachs. The muscle layer surrounding the endoderm was thick in this region, and the muscle of the two sides formed a single complete tube surrounding the two glandular tubes. When these inner tubes began to fuse the dividing wall first broke down dorsally, leaving for a short distance a septum projecting frond the ventral side. The intestine was normal, and the stage of development was indicated as rather young by the existence of but one intestinal bend whereas in older specimens such bends are numerous. The pancreas was divided, the right part opened into the intestine about 1 mm. behind the left, just at the point of fusion of the two endodermal tubes, not into either the right or left tubes but into a short anteriorly directed diverticulum of the single duodenum which stretched from the ventral to the dorsal side passing to the left of the canals (text-fig. 3). From each of the two gall bladders ran cystic ducts which, becoming bile ducts, passed through the corresponding pancreas and opened into the intestine. The liver was single, but from it there ran two ducts, a normal one to the right pancreas forming a normal bile duct, the other branched at the level of the gall bladders and ran into each cystic duct half-way between the bladder and the pancreas (text-fig. 4). Closely applied to the anterior end of the right pancreas was a single spleen. The length of the left pancreas, which lay slightly anterior to the right, was equal to that of the combined right pancreas and spleen. A groove across the middle of the pancreas suggested that this mass also included both pancreas and spleen, but the serial sections showed that the whole was pancreatic in structure. The single spleen may be regarded as part of the vascular system which was here single. The X-ray photograph indicates the presence of an opaque substance in the alimentary canal which was shown to consist largely of sand grains. Since this lay posterior to the pylorus it was not possible to determine through which mouth it had entered. A two-headed viper which lived in captivity for three days was found by Dorner to have died with the right mouth and stomach full of sand grains. This he attributed to an unsuccessful search for ants' eggs,
The Anatomy of a Double-headed Snake
which are said to form the food of young vipers. Comparison of the condition of the fat body with that of a normal specimen of the same size and species might give some indication of the extent to which the animal had fed.
Text-fig. 3. Transverse section at the confluence of the two alimentary canals. M. circular muscle. P. pancreas. Sp. spleen. St. stomach. In figs. 3, 4, 5 and 6, (R) and (L) indicate that the organs belong to the right and left systems respectively. Text-fig. 4. Ventral view of stomachs and digestive glands. A.A. V. anterior abdominal vein. H.P. V. hepatic portal vein. G.B. gall bladder. 1. intestine. L. V.C. inferior vena cava. L. liver. Other lettering as in fig. 3.
The respiratory system The two respiratory systems were independent and complete in themselves. The walls of the air sacs at the posterior ends of the lungs were very closely applied to each other, but no connection could be found between the two systems.
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The vascular system The disposition and the nature of the vessels was examined by dissection and was verified in serial sections of three regions: (a) the hearts and arches; (b) the junction of the left jugular of the right system with the right jugular of the left system; (c) the transverse fold region. There were two hearts lying in separate pericardia and two circulatory systems (text-fig. 5). In the following account the additional letter R or L before
Text-fig. 5. Ventral view of heart and main vessels. A. auricle. Az. azygos vein. C. carotid artery. J. jugular vein. N. vagus nerve. P.A. pulmonary artery. P. V. pulmonary vein. S. systemic artery. T. thyroid gland. r. right. 1. left. Other lettering as in fig. 4.
The Anatomy of a Double-headed Snake
the name of a vessel indicates that it belonged to the system which served the right and left head respectively. The right system was the more extensive, the heart was normal in size and position, and the most noticeable abnormality was a considerable enlargement of the right auricle which covered the ventricle. This auricle was fed by a normal inferior vena cava, right anterior vena cava and a rather smaller left anterior vena cava. A normal pulmonary vein emptied into the left auricle. The anterior azygos vein was enlarged and drained blood from each pharynx and the dorsal musculature near the transverse fold. The left jugular vein anastomosed with the right jugular of the left system and is discussed with that system. The ventricle, conus and arterial systems were normal. The pulmonary artery was thin walled and the left systemic artery had a diameter about five times as large as that of the right systemic artery. The right carotid was suppressed and supplied the thyroid gland in the usual manner. The left system was incomplete, and the left heart, which was rather more than half as large as the right heart, was placed a little anteriorly. The ventricle was completely exposed and lay in a more than usually transverse direction. The only venous supply from regions posterior to this heart was through the pulmonary vein into a large anterior auricle. There was no trace of a posterior vena cava. The left jugular vein ran dorsally to the heart to open into a posterior swelling ventral to the pulmonary vein. The right jugular vein and the left jugular vein of the other system anastomosed slightly anterior to the thyroid gland. Anterior to the anastomosis the diameter of the jugular of the left system was about four times as long as that of the jugular of the right system, whereas posterior to the anastomosis these proportions were reversed. Thus the main supply appeared to be about sixteen times as great as in the other parts of the vessels and ran from the left head into the right heart. The anterior part of the left jugular of the right system was very closely applied to the carotid artery of the right system. The azygos was quite small and ran into the left jugular vein just anterior to a dilatation of this vessel. Dilatations of all three veins, pulmonary, left and right jugulars, opened into the large anterior auricle dorsal to the ventricle. This single auricle had two separate openings into the ventricle, which was normally divided. The serial sections show that from the ventricle there issued the normal three trunks (text-fig. 6 b); the pulmonary artery and the right systemic arch followed approximately normal courses, but the left systemic trunk was closed off before leaving the heart text-fig. 6 a), so that it was represented by Eve-ry small diverticulum of the ventricle. The right systemic trunk Was elongated anteriorly, and slightly posterior to the level of the thyroid gland it gave off the right systemic artery equal in size to the corresponding vessel of the other system. The anterior branch from this trunk was slightly larger than the systemic artery and, dividing into two, formed equally and well-developed right and left carotids. The development of the right carotid may be connected with the fact that no trace of a thyroid gland could be found on this circulation. The right systemic artery
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from the left heart passed dorsally to the left systemic of the other system to join the right systemic of the right system at almost 900. Slightly posterior to this junction the right and left systemics of the right system united to form the dorsal aorta. Posterior to the heart there was thus one main arterial trunk and a single inferior vena cava returning all the blood to the right heart. The circulation of the blood in such a system must have been abnormal. The azygos vein of the right system returned to the right heart blood supplied by the carotids from the left heart. Any blood which passed through the single systemic of the left system would have been returned to the right heart. In spite of this loss of blood the small ventricle of the left heart was as completely differentiated as that of the right heart. Unfortunately nothing is known of the
Text-fig. 6. Transverse sections through the left heart and blood vessels. A. 30 , anterior to
B. Tr. trachea. Other lettering as in fig. 5.
behaviour of this system from observations on the living animal. Since both ventricles were well developed it is probable that both functioned. There is nothing to indicate that they were synchronized, and since four vagus nerves were traced past the hearts they were probably subject to an independent nervous control. If the ventricles were in systole together but very little blood would have passed from one system into the other. On other and probably more frequent occasions blood would first be pumped by the right heart into the left carotids, and then blood from the left heart would be forced into the dorsal aorta. The volume of blood in the two systems would therefore be subject to fluctuations which would be of considerable significance in the very small left system. The existence of a venous connection between the two systems probably lessened the effect of this variation. This fusion did not appear to be part of the general fusion of the two heads and may have been brought about by a distinct developmental process.
The Anatomy of a Double-headed Snake
The excretory and reproductive systems The excretory and reproductive systems appeared to be normal.
Behaviour Adequate records of the behaviour of such abnormal specimens would be of great value, but although several specimens have been kept in captivity (Wall, Dorner, Silly, Vsevolojsky, Borgert, Fischer) the records are too incomplete to make possible an attempt to correlate behaviour and anatomy except on one point. Most authors have considered only the independence or otherwise of the heads. When each has a complete set of receptor organs, a complete brain and normal musculature it is reasonable to suppose that the heads, being subjected to different stimuli, will behave independently. Strohl concluded that the evidence was conflicting, "les uns ont vu les deux tetes se mouvoir et happer toujours simultanement (Silly, 1841; E. C. Fischer, 1896); d'autres rapportent des observations contraires (Vsevolojsky, 1812; Dorner, 1873; Borgert, 1897; Reuter, 1921)." E. C. Fischer (1896) says clearly that "each head is endued with separate will power." The evidence from Silly is less definite. "Le lendemain matin-l'animal qui conservait encore un reste de vie me fut apporte, mais il mourut au bout de quelques instants." We can only conclude that the account of the similarity in behaviour of the two heads which followed was based on evidence supplied by the labourer who caught the animal. This is the only evidence that two separate heads behave simultaneously. The movements of many posterior parts of Vertebrates are controlled by impulses originating in the head. This is especially clear in animals which progress by means of a wave of muscle contraction which passes along the whole body. Observations of such movements in the single region of double-headed specimens would be interesting. The only recorded observations are those of Borgert, whose viper executed rotatory movements continually either to the
right or left.
The line offusion and specific differences Johnson (1901) reviewed the literature of double-headed snakes and paid special attention to the degree of bifurcation. His conclusions will be considered in the light of the rather more detailed account given in this paper. 1. "That the point of bifurcation of the vertebrae is more posterior than would be supposed from external examination." It is clear that when the anatomy of the whole animal is considered there was a line, not a point, of bifurcation. This line was curved and passed from the dorsal surface at the level of the first neck band, through the vertebrae, the point of union of the right systemic arteries, the pylorus, and flattening out ran to the anus close to the ventral surface. Ventral to this line the animal was double, dorsal to it the animal was single. It is therefore possible to picture a
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plane of separation bounded dorsally by the line of bifurcation. The inner layers of tubular organs, e.g. the alimentary canal, retain traces of bifurcation after the outer layers have assumed the normal condition. On the basis of the rather inadequate published descriptions we may conclude that most of the specimens possessed planes of separation similar to that described above, but Strohl has figured an interesting double-headed snake belonging to the Museum of Natural History at Geneva in which the heads are partially fused, the vertebral columns are distinct, and the posterior third of the body bifurcated. So far as it is possible to form an opinion from the published photographs and description the plane of separation here passes through the dorsal side of the animal and is bounded ventrally by a curve similar in direction but placed more dorsally than that just described. That this simple condition is not universal is indicated by a specimen of Tropidonotus sipedon described by Wyman in 1865, in which the vertebral column was said to be double in the anterior, middle and posterior parts of the body and single in the second and fourth sections. 2. " That the point of bifurcation is most likely to occur in the cephalic half of the snake, between 6 and 13 per cent. of the entire length from the cephalic end." If the first of Johnson's generalizations is modified as suggested above it will be impossible to find a simple ratio for the lengths of the double and single parts of the animal. To define the limits of the plane of separation requires more detailed study of the specimens than has been customary hitherto, but, since this plane of separation probably represents the original extent of bifurcation, comparisons would be valuable. Are the lines of bifurcation parallel to a given line? To what extent may the plane of separation be distorted? 3. "That this abnormality is more abundant in some species than in others." Johnson considered but twenty-five cases. Strohl, who reviewed the whole subject again in 1925, extended this number to sixty-eight, and to his list we can now add: Elaphe vulpina (Baird and Girard), 1 specimen; Amaral, 1927. Lampropeltis getulus getulus (L.), 2 specimens; Amaral, 1927. Natrix sipedonfasciata (L.), 2 specimens; Amaral, 1927.
Bothrops atrox (L.), 3 specimens; Amaral, 1927. Crotalus terrificus (L.), 1 specimen; Amaral, 1927. Pituophis sayi (Syn. Coluber melanoleucas, Daud.), 1 specimen; Pope, 1927. Thamnophis sirtalis infernalis 1 specimen; Fisher, 1928. Coluber (Zamenis) fiorulentis 1 specimen; present paper. Most of these are new species to show the abnormality. Although thirty-three of the eighty belong to the genus Tropidonotus and nineteen of these to the species T. natrix, it would be rash to assume that some genera or species are more susceptible to duplicitas anterias than others. The phenomenon is widely distributed taxonomically and geographically. The groups from which most
The Anatomy of a Double-headed Snake
specimens are known are the most widely distributed in regions from which records are likely to be kept. The habits both of snakes and men will introduce factors to invalidate any conclusions drawn from a simple consideration of the numbers of different species recorded. Confirmation of this generalization must await the results of experimental rearing of different species.
Causes of the abnormality Most of the descriptions of specimens have been followed by a description of an hypothetical early stage or by other hypotheses designed to explain the origin of the abnormalities. Such hypotheses were considered by Strohl, who concluded that it may be possible to classify cases of double-headed snakes on the causes of their origin. Apart from the improbability that they arise from more than one cause, it is better to work on the simpler hypothesis that double monsters arise from a single cause until the evidence makes it necessary to adopt the more complex hypothesis. It is, moreover, highly probable that the three monsters from a brood of 120 Zamenis constrictor (L.) described byMitchill, and which exhibited very different degrees of bifurcation, all owed their abnormality to the same cause. A review of the hypotheses which have been suggested makes clear the fact that in general successive writers have referred the abnormality to earlier and earlier stages of development, e.g. (1) Dorner (1873) assumed the fusion of two embryos. (2) Tornier (1901) suggested regeneration after an embryonic lesion. (3) Spemann (1919) refers to division of the dorsal lip of the blastopore. (4) Dareste (1891) Wilder (1908) and Rabaud (1914) to irregular gastrulation. (5) Graper (1931) relates all chick double monsters to pregastrula streamings of protoplasm. Of these Tornier alone refers to a possible external influence different from that acting in normal circumstances but which might occur under natural conditions. Examination of abnormal but more or less mature animals is unlikely to provide sufficient evidence to show whether such abnormalities are to be assigned to several causes or to one. Comparison of the anatomy of naturally occurring specimens with the anatomy which must be assumed to follow from the various hypotheses may serve to check the false though it cannot verify the correct hypotheses. It is difficult to explain the structure of the specimen herein described on Tornier's hypothesis of wound regeneration. The main axis which should represent the original animal which regenerated is different according to whether the vertebral column or the blood system is considered. The left vertebral column appears to be continuous and to bear a branch on its right side. On the other hand the right heart is connected with a much more complete blood system than is the left. Even if we suppose that the difference in the vertebral columns resulted from a more extensive use of the left head this must have been at least equally well developed when use commenced. In
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Tornier's diagram of a hypothetical two-headed snake more detail is shown in the original than in the regenerated skull. This is not mentioned in the text and it is therefore not clear if a deficiency of the regenerated head was indicated. The X-ray photograph and the Spalteholz preparation of the subject of this paper both indicated a normal structure for the two heads. More recently Graper (19.31) has identified the early stages of duplicitas of all kinds in the chick egg with abnormal protoplasmic streamings preceding the formation of the primitive streak. Such streamings are not all hypothetical, for Graper has recorded some of them with his stereo-cinematograph. He explains duplicitas anterias by the setting in of two streamings eccentrically situated on the germinal disc. Streamings which would normally each be sufficient to form one embryo meet, each turns aside in two halves to form a new anlage with a similar half streaming from the other side (text-fig. 7). Since the positions are eccentric the material from one side will be exhausted before that from the other, leaving the more posterior region single. Although each of the resulting heads is composed of material of different streamings Graper has carried the analysis back so far that we could not expect to detect this in the adult. In one point, however, our specimen does not appear to receive a full explanation on Graper's hypothesis. The two hearts are both formed from material of different streamings, but each one is composed of right and left anlagen in the normal relations. When the material of the shorter side gives out it would be expected that the right systemic arch of one system would join the left systemic arch of the other. In the specimen, however, in spite of the fact that the right systemic from the left heart must pass close to the left systemic from the right heart it goes further and joins the right systemic from this heart (text-fig. 8). Bataillon and several subsequent workers have pointed to the effect of such external conditions as delay in fertilisation and the presence of foreign substances in the environment. Strohl has also pointed out that most of the specimens of double-headed snakes have been caught when a few days old at the end of the breeding season. Of the specimens described since his review information is available for two specimens only. Pope's Pituophis sayi (Schlegel) was collected in the summer of 1902 at Sylvan Grove, Kansas. Fisher's Thaninophis was 2 or 3 days old when found dead in California on 4 September, 1926. The specimen described in this paper was caught near Cairo, Egypt, probably during 1925, but no information could be obtained as to its date of capture. The factors which may be involved in this hypothesis are: (1) Ovarian exhaustion. (2) Prolonged sojourn in the oviduct and (a) toxic effect of secretions, products of metabolism or of destruction of other eggs; (b) delayed fertilization and consequent over-ripeness of ova.
The Anatomy of a Double-headed Snake
(3) Unusual environmental conditions during development. (4) Late hatching may be the effect and not the cause of the abnormality. The first of these factors is not known to have any effect. Both factors included under (2) are known to result occasionally in abnormal development. The cause may be the same in both cases. Abnormal embryos always represent a small proportion of the broods, and some of the specimens belong to viviparous species. It is therefore probable that neither the climate of the nest nor the district is responsible for the abnormalities. The second is therefore the most likely factor to be involved.
Pregastrula streaming which results in duplicitas anterior = Heart rudiment. r' Anterior border of embryo.
Text-fig. 8. Ventral view of hearts and main arteries. A. As required on Griper's hypothesis. B. As in the specimen.
1. The study of the anatomy of a double-headed snake indicates that the boundary of a plane which separates the two heads is well defined but may be distorted. Consideration of this plane may provide some evidence of the manner of origin of the two parts. The example described in this paper does not
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agree with the regeneration hypothesis put forward by Tornier, nor does it support Graper's hypothesis of streaming. 2. Two related developmental phenomena may be distinguished: (a) development of the embryo modified by an unusual stimulus, (b) the readjustment of systems to make a viable entity of this unusually organised matter. The first of these events would result in a specimen single on one side, completely double on the other side of the line of bifurcation. The second process becomes much more important at the end of the embryonic period, especially in the complex systems which pervade the whole animal. Doubling of the alimentary system does not introduce important difficulties, and hence little modification of the simple condition is necessary although in the specimen described a diverticulum of the intestine carried the secretions from the left pancreas into the intestine. In the vascular system the jugular anastomosis which occurred well to the ventral side of the line of bifurcation stood out as an exception to the main scheme of fusion, and since it can be shown to be of considerable functional importance it was probably a secondary development which converted an ineffective into an effective system. The commissure connecting the roots of the last pair of spinal nerves to issue ventral to the line of bifurcation is also considered to have been such a secondary development. Thus the vascular and nervous systems may have followed the normal development in the separate parts until the two sets of systems interfered with each other. They then appear to have been made workable by the development of suitable anastomoses, a process for which no allowance is made in the current hypotheses of experimental embryology. Further investigation of this adjustment may throw some light on the origin of variation, i.e. of individual differences which cannot be attributed to germinal differences and which may be adaptive. 3. The study of the origin and development of two-headed snakes deserves experimental treatment especially by the incubation of overripe or of chemically treated eggs. Fortunately Tropidonotus natrix, at once plentiful and innocuous, appears to be susceptible to the causes of this malformation. In conclusion I wish to express my thanks to Prof. J. Stanley Gardiner, to whom I am indebted forthe specimen and for permission to dissect it thoroughly, to Mr C. Forster Cooper and Mr C. F. A. Pantin for help and advice, to Mr W. Parker, of the British Museum, for the identification of the species, and to Dr Barclay for the X-ray photograph.
EXPLANATION OF PLATE Figs. 1 and la. Dorsal and ventral views of the Coluberflorulenti&. Fig. 2. X-ray photograph of Coluberfloruienti&.
Journal of Anatomy, Vol. LX VII, Part 2
Fig. 2. HEASMAN-THE ANATOMY
The Anatomy of a Double-headed Snake
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