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THE ANATOMY OF A DOUBLE-HEADED SNAKE

BY W. J. HEASMAN
Senior Curator, University Museum of Zoology, Cambridge
CONTENTS
PAGE
Introduction. . 331
Material. . 332
External features. . 332
The axial skeleton and nervous system . 333
The alimentary system . . . 334
The respiratory system . . . 335
The vascular system . . . 336
The excretory and reproductive systems . -339
Discussion
Behaviour. . . 339
The line of fusion and specific differences 339
Causes of the abnormality . 341
Conclusions. . . . 343
References . . . 345
INTRODUCTION
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 natu-
rally 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.
332 W. J. Heasman
Altogether since records have been kept, some seventy cases of double-
headed 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.

MATERIAL
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 pre-
served 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 relation-
ship of the vertebrae. The mouths and oesophagi
of both sides appeared to be equally capable of
functioning.
At the level of the second neck band, 23 mm. Ribs
from the tip of each snout, the integument of the
two "necks" united to form a common invest-
ment. At this point there was a transverse and Anal Scale
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 phenome-
non of fixation was shown by the fact that most
of the ventral scales had an irregularity which Tet-ig. 1.owingthraiewoftsen
scaling showing the ribs as seen
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 333
bend in the body axis. The accompanying photographs make further descrip-
tion 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.

K.

. */ , .~~~~~~/ \

A~V

7, /

-A 13
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 approach-
ing 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
334 W. J. Heasman
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 sur-
rounding 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 335
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.

Fig. 3.

Fig. 4.
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.
336 W. J. Heasman
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 337
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 pos-
terior 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 pul-
monary 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 develop-
ment 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
338 W. J. Heasmcan
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

Y'r

5. A~~~~~~~~I, f)

A B

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 sub-
ject 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 339
The excretory and reproductive systems
The excretory and reproductive systems appeared to be normal.

DISCUSSION
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
340 W. J. Heasman
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 con-
clude 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 341
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 ab-
normality 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 gastrula-
tion.
(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
342 W. J. Heasman
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 sub-
stances 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 343
(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.

[Text-fig. 7. Pregastrula streaming which results in duplicitas anterior (Graper)


= Heart rudiment. r' Anterior border of embryo.

A B
Text-fig. 8. Ventral view of hearts and main arteries. A. As required on
Griper's hypothesis. B. As in the specimen.

CONCLUSIONS
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
Anatomy LXVII 23
344 W. J. Heasman
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 pro-
cess 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 chemi-
cally treated eggs. Fortunately Tropidonotus natrix, at once plentiful and in-
nocuous, 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 Plate I

Fig. 1. Fig. Ia.

Fig. 2.

HEASMAN-THE ANATOMY OF A DOUBLE-HEADED SNAKE


The Anatomy of a Double-headed Snake 345
REFERENCES
For a full bibliography to 1925 see J. Strohl.
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