THE H D I T S OF ENHYDRICTIS GALICTOIDES.

241

17. Note on the Habits of Enhydrictis galictoides, with Description of

some Limb-bones of this Mustelid from the Pleistocene of Sardinia.
By DOROTHEA
M. A. BATE,F.Z.S.
[Received February 15,1935 : Read May 7, 1935.1

(Text-figures1 & 2.)

It is to Dr. Forsyth Major that we o m the discovery of the large Mustelid

from the Pleistocene of Sardinia to which he gave the name Enhydrictis
galictoides. He read a preliminary account of the skull of this species at
a meeting of the Zoological Society in 1901, but, unfortunately, no figure of
this specimen was published with the description, and its present whereabouts
are unknown. From his study of this skull Dr. Major was convinced that
he was dealing with an animal specialized for an aquatic mode of life. He
summed up his description (1902, p. 626) by saying :- From the description
and comparisons it results that the Sardinian fossil belongs to an amphibious
member of the Nustelina, coming nearest among recent forms to the South
American Galictis *.
A few years ago drawings which are believed to represent the type-skull of
E. galictoides were found among Dr. Majors papers, together with a figure
of a ramus belonging to this species which is in the Natural History Museum
at Basle. These have recently been published by Dr. Pilgrim (1933), who
redescribed the skull and gave the first description of the mandibular ramus.
He associated this genus in a group including the Recent Grison and Tayra,
thb extinct Pannonictis, Mustelid gen. indet. of Zdansky, and perhaps
Trochictis, but was evidently strongly inclined to consider that Enhydrictis
was not aquatic in its habits, and suggested that a t least this theory had not
been proved.
I n his original description of Pannonictis Dr. Kormos recognized its affinity
with Enhydrictis and accepted the theory of the specialization of the Sardinian
Mustelid for an aquatic mode of lifa. The matter stood thus until last year
(1934), when Dr. Kormos described a new species of Pannonictis and referred
in a footnote (p. 130) to some limb-bones of E. galictoides preserved in the
Bade Museum which he said showed a great resemblance to those of Pannonictis and provided evidence against the theory of specialization for an aquatic
mode of life. The description of the limb-bones of Pannonictis will be awaited
with much interest. Dr. Pilgrim then communicated with the authorities
of the Basle Museum, and through the kindness of Professor H. G. Stehlin
(Director of that institute) and of Dr. Helbing the Sardinianspecimens have been
sent over and their description entrusted t o me, since I was already studying
some limb-bones of a Mustelid from the Pleistocene of Malta. I should like
to record my appreciation and thanks for this opportunity of studying the
Sardinian specimens, and also to Dr. Pilgrim for his interest and encouragement.

Now more generally referred t o as Orison (see Thomas, 1907, p. 162).

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MISS DOROTHEA M. A . BATE : NOTE ON

The collect,ion of Mustelid limb-bones from Sardinia is a small one and

several of the bones are incomplete. The following specimens are included :The proximal end of a left humerus.
The proximal ends of two right femora.
One ( 2 ) complete left tibia.
The proximal end of a right tibia.
A right calcaneum.
A right asbragalus.
Besides the above there are also a caudal and two dorsal vertebrae, a small
left first metacarpal, and a number of phalanges which do not seem of particular interest and perhaps may not all represent E. galictoides. These
specimens were all collected by Dr. Forsyth Major at San Giovanni, near Iglesias,
south-west Sardinia, which is the type-locality of E. galictoides. It may
be of interest to mention that the known range of the species has recently been
extended by the present writer finding a lower carnassial in a Pleistocene
deposit near Cagliari in the south-east of the island. This specimen was
associated with remains of Cuon sardous*, Cermus sp., Prolagus sardus,
Rhclgamys orthodon, and other extinct forms.
The chief points to be considered in the study of this collection of limbbones from Sardinia were :(A) Do these limb-bones represent the same species as the type-skull of
Enhydrictis galictoides ?
(B) What evidence do these limb-bones provide regarding the mode of life
of the species which they represent, more especially, was it a n animal of aquatic
or terrestrial habits ?
(C) What indication do these specimens give of relationship with other
forms, living or extinct 1
(A) The fact that Dr. Major had collected both the skull of E. galictoides
and the limb-bones now under review in a single locality, and yet had not
immediately recognized that they might be associated, made it necessary to
proceed With caution. It should, however, be remembered that miscellaneous
limb-bones in a large collection are apt t o be reserved for study a t a later date,
and since Dr. Major considered the skull to be that of a n aquatic animal no
obvious connection was then suggested by the limb-bones.
When the Sardinian limb-bones were first received it seemed as if they must
be too small to represent an animal of the size indicated by the type-skull.
A careful study and comparison of the collection have, however, modified this
view, and it is now believed that in so far as size is concerned, as well as in
other respects, there is every likelihood that these bones represent E . galictoides.
It was interesting to find that there was little difference between the relative
proportions of the Sardinian skull and limb-bones and those of some other
Mustelids. For instance, a skull of Tayru barbara in the British Museum
collection (3.7.25.3) has a maximum length of 12.1 cm., while that of E. galictoides has a length of 10.7 cm. (Pilgrim, 1933, pl. ii. fig. 1 a). I n a femur of
T. barbara the greatest width of the proximal articular end is 22 mm. ; in two
examples of this bone from Sardinia this measurement is 21.5 mm. and 21 mm.
The correspondence in these measurements suggests that, allowing a reasonable
margin for individual variation in size, there is nothing in this respect t o prevent
the acceptance of t,he Sardinian limb-bones as those of E. galictoides.
(B) Although the collection of limb-bones from Sardinia is small and
incomplete it seems to provide suffcient evidence to show that the skeleton
of E . galictoides was in no way specially modified for a n aquatic mode of life.

See Major, 1901, p. 833.

THE HABITS O F ENHYDRICTIS GIALICTOIDES.

243

The proximal portion of the femur appears to prove this (see text-fig. l),
for it is similar to that of a normal terrestrial species; the shaft is rounded
instead of wide and flat as it is in Lutra, a character seemingly found only in
the Lutrinz. Furthermore, the general shape of the proximal portion of
the Sardinian femur is similar to that in many other terrestrial Mustelids,
whereas in Lutra i t is square and compact, with the head closely attached.
(C) There is little to be said under this heading, for, although these limbbones evidently represent B Mustelid, they do not show a specially close
resemblance t o the corresponding bones of any Recent species which has been
available for comparison. This, after all, is not surprising if Enhydrictis is
a survival from earlier Tertiary times, and since older species are generally
known only from teeth and parts of skulls there has been little opportunity for
comparison.
Description of Specimens.
Metacarpal.-A left first metacarpal having a total length of 14 mm. is
practically identical with the corresponding foot-bone of Grison ajlarnandi,
which is probably a much smaller animal than E . galictoides. Not much can
be gleaned from this single foot-bone, so it seems wisest simply to record it
as possibly that of E . galictoides and await further material.
Vertebra-The Sardinian vertebra are larger but otherwise seem to be
similar to the corresponding bones of G. allamandi a- far as it is possible to
Text.figurc 1.

A. Lutra lutra, proximal portion of right femur, posterio aspect.

B.
,, ,, transverse section of shaft of A.
C. Enhydrictis galictoides, proximal portion of right femur, posterior aspect.
D.
,,
,,
transvarse section of shaft of C.
E.
9,
,,
the same specimen as C, anterior aspect.
All figires approximately natural size.

judge from the skeleton of an immature animal in the British Museum

collection (210 a).
Humerus.-Unfortunately this bone is represented by only the proximal
end of a single example of the left side. This end of the bone is not so characteristic as the distal end, but even this imperfect portion from Sardinia proves
sufficient to show great differences from the corresponding bone of Lutra
lutra. It is important to find t h a t the shaft is straighter, and there is no

244

MISS DOROTHEA M. A. BATE : NOTE ON

expanded surface below the greater tuberosity on the anterior side of the
bone such as obtains in the humerus of Lutra. The lesser tuberosity is strongly
developed and merges into the head of the humerus as in Tayra and Meles,
whereas in Lutra it is separated by a shallow groove. The greater tuberosity
is low and continues anteriorly well beyond the middle line of the shaft as
in Tayra.
Femur (text-fig. 1, C-E).-There
are two examples from S a r W a of the
proximal portion of the femur, each belonging to the right side, and these
are similar except for a slight difference in size. The figures, showing one of
Text-figure 2.

Enhydrictis galictoides Major.

A. Left tibia, anterior aspect.
B. Right astragalus, dorsal aspect.
c. ,,
,,
plantar aspect.
D. Right calcaneum, plantar aspect.
E. 9 ,
dorsal aspect.
A, slightly reduced; the other figures approximately.

x 14.

these specimens and the corresponding bone of Lutra, clearly indicate the contrast between these two bones. The specialization of the femur in Lutra
is shown in the shortening of the complete bone, by the square outline of its
proximal end, and by the wide and flat shaft of the bone. The Sardinian
femur is a comparatively longer bone than that of Lutra and has a rather
slender shaft which is almost circular in section near its centre. The shaft
has a width of 6.2 mm., with an antero-posterior thickness of 6 m.,while
the corresponding measurements in the femur of L. lutra are : width of shaft
9.7 mm. and antero-posterior thickness 7.4 mm.
Tibia (text-fig.2, A).-The tibia shown in text-fig. 2, A, was sent in two pieces,
which were obviously parts of a single specimen, so they have been joined, but

THE HABITS OF ENHYDRICTIS QALICTOIDES.

245

it is possible that the complete bone may have originally been slightly longer ;
its maximum length is 83.7 mm. and the maximum width of the proximal
articular end is 18 mm. This specimen is slender and much straighter than
the corresponding bone of Lutra or Potamotherium. The cnemial crest is
well defined and continues as a sharp edge down the anterior face of the shaft
for two-thirds or more of its length, in a manner rather similar to that seen in
the tibia of Grison furax (Brit. Mus. 210 c). The proximal facet for articulation
with the radius and the grooving of the centre of the shaft on its posterior
aspect are also much as in Grison.
The complete tibia is smaller than might be expected from the size of the
type-skull of E. galictoides, but a second example of the proximal portion is
a stouter bone with a maximum width of the proximal drticular end of 20.5 mm.
Calcaneum (text-fig. 2, D & E).-The total length of the Sardinian bone is
24.3 mm. and its maximum width is 14 mm. ; except that its trochlear process
is more salient it is not unlike the corresponding bone in Grison allumandi.
The distal portion of the bone compared with the proximal portion is longer
in the Sardinian specimen than in Lutra. Further, the calcaneum of Lutra
is more twisted on its long axis and the groove for the tendon achilles is more
deeply excavated than in E. galictoides.
Astragalus (text-fig. 2, B & C).-The astragalus from Sardinia is of the right
side and may be associated with the calcaneum. The total length of the
bone is 17.5 mm. and its greatest proximal width is 12.8 mm. The smooth
surface of the head for articulation with the scaphoid is not so strongly developed
as in Lutra or in Enhydriodon ( 1 ) latipes, in which it is very extensive and is
directly connected with the distal facet for articulation with the calcaneum.
The width of the proximal end of the bone is comparatively less than in Lutra
and the central groove of the trochlea is shallow. The proximal articular
surface for the calcaneum is more concave than in Lutra and the proximal
end of the tibia1 border of the trochlea is not as salient as in Lutra, a character
seen in an exaggerated degree in the astragalus of Potamotherium.

Conclusions.
The study of the Mustelid limb-bones from Sardinia has shown that these
in all probability represent Enhydrictis galictoides Major and that this animal
was not specialized for an aquatic mode of life. These specimens provide
little evidence of close relationship with any one particular group of Mustelid,
and this is probably due to lack of remains of other extinct genera for
comparison.
LISTOF WORKS
REFERRED TO.
KORMOS,
T. 1931. Pannmktis plioc~niecc,n. gen., n. sp., a new Giant Mustelid from
the Late Pliocene of Hungary. Ann. Inst. Reg. Hung. Geol. xxix. pp. 167177, pl. iii.
KORMOS,T. 1934. Neue und wenig bekannte Musteliden aus dem Ungarischen Oberplioziin. Folia Zool. et Hydrobiologica Latvijas Univemit. sistemat. Zool.
Inst. Riga, v. nr. 2, pp. 129-158.
MAJOR, C. I. FORSYTR.
1901. Remarks on Remains of Cyon surdous (Studiati) from
a Cave at Capo Caccia (N.W. Sardinia). Roc. Zool. SOC.London, 1900,
pp. 833-835.
1902. On Enhydrictis gulictoides, B new Fossil Mammal from
MAJOR,C. I. FORSYTH.
Sardinia. Proc. Zool. SOC.London, 1901, pp. 625-628.
PILGRIM,
G. E. 1933. The Genera Trochictis, Enhyd~ictis,and Trocharion, with Remarks
on the Taxonomy of the Mustelida?. Proc. Zool. SOC.London, 1932, pt. iv.
pp. 845-867, pls. i., ii.
THOMAS,
0. 1907. On Neotropical Mammals, etc. Ann. & Mag. Nat. Hist. (7) xx.
pp. 161-168.