THE RED DEER OF THE CAUCASUS. 789

52. The Red Deer of the Caucasus. By EDGAR N. BARCLAY *. (From

the Department of Zoology, British Museum (Natural History).)

[Received May 29. 1934 : Read October 23, 1934.1

(Plates I.-III.-f ; Text-figures 1-5.)

INTRODUCTION. The status of the Red Deer, Eastern Red Deer, or Maral of the Caucasusl

Black Sea, and Caspian provinces has never been clearly understood. Some advance in our knowledge of these deer is to be gained by a study

of the valuable series of skulls and antlers which form a part of the magnificent collection of big game specimens bequeathed to the British Museum by the late St. George Littledale.

The particular specimens which form the basis for these observations were obtained in the Caucasus, and they are of very special interest in that they appear to bridge over the characters which have been advanced as dis- tinctive of the European Red Deer and the various forms of Wapiti of Asia and North America; the presence of Wapiti in the latter country being apparently analogous to the case of the Elk (Alces) and Reindeer (Ranqifer), which presumably spread from Asia to the New World during an epoch when such transmission was possible. The importance of the Caucasian deer therefore liea in the fact that they appear to intergrade with the two great groups of deer which have been referred to Cervus elaphus and Cervus cam- densis. So little is a t present known concerning many of the other forms of Asiatic deer referable to Cervus that some at least of these may eventually prove to be no more than local races of the deer now under consideration.

The chief points of difference between the extreme forms of the Red Deer and Wapiti concern antler pattern, size, colour, and voice ; the aflcinity of the Caucasian deer with their European and Asiatic neighbours may therefore be dealt with in that order.

In connection with antler pattern my special thanks are due to Mr. Martin A. C. Hinton for taking the photographs here reproduced.

ANTLER PATTERN. It may be taken as an accepted fact that no two antlers of any species of

deer are ever exactly alike, and the variation to which they are subject is often so great that in the absence of a series of specimens they might easily be regarded as examples of distinct species. Such variations may cause the partial atrophy of one part of the antler to compensate for the extra development of some other part, and these may be transmitted and fixed by heredity in a relatively short space of time, evidence of which may be found by an examination of the history and antler pattern of various herds of park deer. The production of variations from typical forms, however, does not obliterate the original specific characters, and antIer pattern is sufficiently definite to form a valuable guide for the purposes of classification. The pattern of antlers is always of

* Communicated by M. A. C. HINTON, F.R.S., F.Z.S. t For explanation of tho Plates, see p. 798.

790 MR. EDGAR N . BARCLAY ON

greater importance than their size, since their growth is dependent primarily on the quality of the feeding. In the case of the typical Red Deer and the typical Wapiti the antlers are perhaps the most readily distinguishable characters, and the points of difference are too well known to require more than brief reference.

The typical tines of the fully developed Red Deer stag comprise brow, hez, and trez, together with a varying number of terminal points which con- stitute the crown or surroyals. In the typical Wapiti antler the arrange. ment of the brow, bez, and trez tines is the same as in the Red Deer, but it is distinguished from the latter by the presence of a fourth tine which is usually the largest, and by the remaining terminal points being in the same fore-and- aft plane as the fourth tine. For this reason, and also on account of the fact that with antler deterioration the bez and trez tines may no longer be grown, it is the pattern of the antler above the trez tine which is of the greatest

Text-figure 1.

Right antler of Hungarian stag.

importance. This is well illustrated by the antlers upictei in P1. I. fig. 1, which are those of an old stag killed at Tartarow, Galicia, by the late St. George Littledale in 1906 (B.M. no. 31.6.1.81). In this head the bez and trez tines are absent, the brows are very short, but the beam is still massive and the crowns have remained complex.

The occurrence of a fourth tine, quite separate from and below the points of the crown, must be an exceedingly rare occurrence among the stags of Scotland and England, if, indeed, it occurs at all. Of the hundred and eleven heads from the mainland of Scotland figured in H. F. Wallace’s ‘ British Deer Heads ’ it cannot be said that one definitely displays this characteristic, though in both antlersof nos. 16 and 91, and in the left only of no. 9, a tendency towards this particular development is shown. Of the nine heads figured of English wild stags not one displays it, and in the much more complex and many- pointed heads grown by park stags only nos. 158 and 172 of the seventeen

THE RED DEER OF THE CAUCASUS. 791

pairs figured display fourth tines, and in both these specimens it only occurs on one of the antlers. In no wild-bred British or Scandinavian examples that I have seen have I noted a fully developed fourth tine between the trez tine and the points of the crown, and its absence is probably a definite antler characteristic of the deer of these countries.

An examination of the antlers in the historical collection preserved at Moritzburg, near Dresden, is of interest. Of the sixty heads figured in Dr. A. B. Meyer’s two folio works (‘ Die Hirschgeweih-Sammlung im Konigl. Schlosse zu Moritzburg,’ 1887 and 1883) only four show the development of an independent fourth tine. Many of these heads have from twenty to fifty points, and the rarity of the fourth tine is therefore of particular interest, as its occurrence seems unrelated to the multiplication of the points of the surroyals ; furthermore,

Text-figure 2.

Right antler of Hungarian stag.

in not one of the specimens figured do the points of the crowns appear to be in the same fore-and-aft plane.

NO attempt is here made to make a comprehensive survey of European antlers for the presence of the fourth tine, but that it occurs among the stags of Central and East-Central Europe is certainly the case, and examples are not infrequently seen at the various continental exhibitions of sporting trophies. The three examples figured are taken from photographs ; the first head was exhibited at Vienna in 1910, and the other two at Leipzig in 1930. Text-fig. 1 (Supplement to ‘ Country Life,’ July 30, 1910, p. xlx) is the antler of a Hun- garian stag with a massively developed fourth tine. Text-fig. 2 (‘ Wild und Hund,’ May 30, 1930, p. 547) is also from a Hungarian stag. In this antler the fourth tine is developed midway between the trez and the surroyals, the points of which are roughly in the same fore-and-aft plane. Text-fig. 3 (‘ Wild und Hund,’ May 30, 1930, p. 547) is the antler of a stag from Rumania

792 ME. EDGAR N . RARCLAY ON

which quite definitely inclines to the Wapiti pattern. These examples are not specially selected from all the heads exhibited, but are merely taken from illustrated articles on the exhibitions in question.

The antler pattern of the Caucasian stags appears to be extremely variable, some being indistinguishable from the typical Red Deer pattern, whilst others are markedly of the Wapiti type. Figs. 1 & 2 (Pl. 11.) and fig. 2 (Pl. 111.) are three specimens shot by Littledale in the Western Caucasus, and now in the British Museum (B.M. no. 31.6.1.82-84-85). If these examples be compared with fig. 1 (Pl. III,), which is the antler of a Wapiti shot by Littledale in the Thian Shan (B.M. no. 31.6.1.78), the similarity of pattern is strikingly evidenced. Each of these antlers shows a strongly developed fourth tine with the points of the crown in the same plane. In fig. 1 (Pl. 11.) the lateral compression of the upper portion of the beam and crown is also to be noted, as this is a feature which is very common among the Wapiti of North America.

Text-figure 3.

Xight antler of Rumanian stag.

Of the remaining skulls and antlers in the Littledale bequest no. 31.6.1.87 has already been figured by Lydekker in ' Great and Small Game of Europe, Asia, and America,' 1901, p. 223. It was shot in the Western Caucasus, and is definitely of the Red Deer pattern, with eight points on the right antler and seven on the left. No. 31.6.1.83 has uneven antlers, in that the right has a strongly developed fourth tine intermediate in distance between the trez and the crown, whilst the left has no corresponding fourth tine. No. 31.6.1.86, shot on 22. viii. 1887, is an eleven-point head, and is only remarkable on account of the left bez being represented by a fork or double bez in place of a single tine. No. 87.12.22.4 was presented to the Museum by Littledale in 1887, and was figured by Lydekker in Cat. Ung. Mamm. Brit. Mus. 1915, vol. iv. p. 127. It inclines to the Red Deer pattemthough both fourth tines, which are bifurcated, are somewhat below the crowns.

The antlers of the two mounted specimens of Caucasian stags in the exhibition galleries of the Museum both incline to the Red Deer pattern.

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They are, however, very dissimilar in type, one having heavy beamed, short' tined, and straight antlers with a tendency to palmation in the crowns, whilst the antlers of the other specimen are not so heavy, more curved, longer tined, and with no tendency to palmation.

As to whether the deer of northern Persia, the Caspian area, Turkey, and the Crimea are identical with those of the Caucasus must remain in doubt, as the available material is, in my opinion, quite insufficient to form a basis from which to draw any definite conclusions. Of the few specimens in the Museum single antlers from the Crimea and the Beinbighas Mountains have already been described and figured by Lydekker (Proc. Zool. SOC. 1890, p. 363), and antlers from Circassia have been figured by Sclater (Trans. ZOO]. 8 0 ~ . vol. vii. pl. xxix.), but reference must be made to a single antler from Lake Van, Armenia, presented by Lord Arthur Hay in 1854 (B.M. no.

Text-figure 4.

Ant l~rs of stag from the Thian Shan.

54.4.26.6). As will be seen from the photograph of this antler (Pl. I. fig. 2 ) , the fourth tine is strongly developed and is independent of the crown, which tends to palmate. The brow and bez tines have unfortunately been sawn off, but the development of the whole antler is remarkable, as the following measurements show. The length on outside curve is 484 in., circumference of beam above bez tine 7Q in., circumference of beam midway between trez and fourth tine 8 in., length of trez tine 17 in., length of fourth tine 134 in., greatest width of palmated portion of crown 49 in., points 12.

The antlers of the various forms of Asiatic Wapiti vary considerably, and though some examples are indistinguishable from typical North American specimens, others incline more towards the Red Deer pattern. Fig. 1 (Pl. 111.)) which has been previously referred to, is a fairly typical example of the antler pattern of the Thian Shan and Altai deer, but text-fig. 4 (' Great and Small Game of Europe, Asia, and America,' fig. 14) depicts a pair of antlers collected

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THE RED DEER OF THE CAUCASUS. 795

by Messrs. J. V. Phelps and P. Church in the Thian Shan which depart most markedly from the normal type. Text-fig. 5 is figured from a mounted speci- men in the Museum from Chenkend, Turkestan, the pattern of which is close to that of some heads from the Caucasus.

The typical North American Wapiti antlers need no special reference, but those of the Pacific coast assigned to occidentalis sometimes tend to palma- tion and also to the cupping of the crowns.

Though the material surveyed is certainly limited, I think it is sufficient to show the intergradation of the pattern of Red Deer and Wapiti antlers, and it is apparently in the Caucasus that the intermediate types are most manifest.

Text-figure 5.

Right antler of stag from Chenkend, Turkestan.

CRANIAL MEASUREMENTS. From the foregoing table of measurements the chief points of difference

between the various groups of skulls may be summarized as follows :- I. The skulls of Norwegian and Scottish specimens are much broader in

relation to their length than those of the Caucasus and Galicia, and they are somewhat broader than the Wapiti skulls.

2. The skull from Galicia is very close in all measurements to the Caucasian specimens.

3. The Caucasian and Galician skulls are longer in relation to their breadth than any others in the table.

4. The two skulls from the Thian Shan are of greater relative breadth than those from the Caucasus, and they are also relatively broader than those from North America.

5. The skulls from the Caucasus and Galicia are closer to t.he Wapiti skulls than they are to those from Scandinavia and Scotland.

WEIGHTS. Owing to the variability of the condition factor, weight, as an indication

Qf sue, is a somewhat uncertain guide, and even though stags are shot at the

796 MR. EDGAR N. BARCLAY ON

same season of the year the rut may entail a loss of several stono in the case of a master stag.

The only authenticated weight of a Caucasian stag that 1 have been able to trace is that of an animal shot in 1895 by the Grand Duke Sergius Mikhailo- vitch in the Kouban district, which weighed 53 stone. This is recorded by E. Demidoff, Prince San Donato (‘ Hunting Trips in the Caucasus,’ 1898, p. B), but he does not state whether this weight included the gralloch.

The heaviest stag killed within recent times in Central Europe that I have found recorded is mentioned by H. J. Elwes (Supplement to ‘ Country Life,’ July 30, 1910, p. 6). This animal was killed in Javovina, a forest in the Hohe Tatra, and weighed over 62 stone as it fell, and just over 49 stone clean. E. von Dombrowski has recorded a stag from Munkacs with a clean weight of 44 stone 4 Ib., whilst W. A. Baillie Grohman in ‘ Sport in the Alps ’ states that the average weight of thirty stags shot in one season in the same forest was 29 stone 9 lb. clean, of which the heaviest weighed 44 stone.

From data collected by Dombrowski, Huxley, and Baillie Grohman the following average clean weights of adult stags may be quoted to indicate the decrease of weight in the westward extension of the Red Deer’s range. In order to facilitate comparisons I have converted the weights from kg. to stones : Bukovina, average of fourteen 36.8 stone ; East Russia, average of ten 28.9 stone ; Styria, average of nine 23.5 stone ; N. Bohemia, average of fourteen 21.4 stone ; Bavaria, average of fifty-three 18.5 stone ; Alsace- Lorraine, average of thirty-six 16.2 stone ; Scotland, average of forty-five 15 stone.

I have failed to trace weights of wild examples of Central Asian Wapiti, but W. Winans records it park stag which weighed 837 Ib., or 59 stone 11 lb.

According to H. E. Anthony (‘ Records of North American Big Game,’ 1932, p. 40) the male Wapiti of North America attains an average weight of about 700 lb. (50 stone), “ but specimens of 1000 to 1100 pounds [71 stone 6 lb. to 78 stone 8 Ib.] are known.”

.COAT COLOUR.

There are only two complete skins of the Caucasian Red Deer in the Museum, both of them mounted specimens which have been in the collection for over thirty years. In the following observations therefore the question of individual variation and differences of colour due to sex, age, and seasonal change cannot be dealt with. Of the two specimens available one is an adult stag in early summer coat that has been mounted with the antlers of another individual, and the other is an adult stag that appears to be in winter pelage. It so happens that both these specimens, together with examples of the Thian Shan and American Wapiti, are so placed in the gallery that they may all be viewed together ; and considering that these specimens are representatives of three continents, one cannot fail to be impressed by the general similarity o i coloration and colour-pattern, though the possibility of the specimens having faded some- what must not be lost sight of.

The general colour of the Caucasian stags is a brownish grey, the rump- patch more yellowish than in the more typical races of Cervus elaphw, thighs and underparts darker, median tail-line dark brown.

In the mounted example of the Thian Shan stag presented by Littledale in 1902 the general colour is greyer, head and neck darker than body, rump- patch larger and more yellowish, limbs, flanks, and underparts dark brown, contrasting with the body-colour more markedly than in the Caucasian stags.

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The general colour of the male from the Yellowstone National Park, dated 1884, is more inclined to yellowish brown. The head and neck are darker, rump-patch as in the Thian Shan specimen, but no median line to tail, limbs, flanks, and underparts, dark brown.

According to Lydekker (‘Great and Small Game of Europe, Asia, and America,’ p. 223) the herd of Caucasian deer which was formerly a t Woburn Abbey showed in stags and hinds of three or four years of age distinct yellowish spotting when in the reddish summer garb, whilst when in winter dress the shoulders, thighs, and underparts showed a large amount of black. If this darkening of the underparts is typical, it makes the Caucasian deer still closer in coloration to the Wapiti than the skins in the Museum show. Additional material is greatly needed, but on the evidence of coat colour alone it seems doubtful if there are any differences which can be regarded as of more than racial significance.

VOICE. No comparison between the deer referable to Cervus elaphgs and Cervus

canadensis would be complete without some reference to the question of voice, In the more typical examples of the two forms of deer this is sharply defined, the Red Deer stag uttering a roar and the male Wapiti a whistle or bugle. These facts are of course well known, but little has been recorded concerning the voices of the more intermediate types. Never having heard the roar of the Caucasian stag, I am indebted to Dr. Schwarz for the following information. Dr. Schwarz, who was familiar with the herd of pure bred Caucasian deer which existed for many years a t the Berlin Zoological Gardens, states that the roar of the Caucasian stags was immediately recognizable from the German stags by being much fuller toned and less hoarse.

Whether or not the deer of the Thian Shan and Altai utter a cry identical with that of the American Wapiti I am unable to say, but Radde, Church, and other writers appear to be agreed that it is essentially the same.

With reference to the Wapiti-like deer of Kansu (Cervm kansuensis Pocock), Mr. H. F. Wallace writes (‘ Big Game of Central and Western China,’ 1913, p. 203) :-“ Their roar is quite different from the wonderful ringing bugle of the North American Wapiti, which is one of the most musical sounds emitted by a wild animal. It resembles the sound made by a Red Deer, but it is rather deeper in tone, and in the case of one or two stags I heard there was just a suspicion of ‘ bugling ’ a t the end of the roar.”

This statement is particularly interesting as i t seems to indicate a cry intermediate in tone between the Red Deer and the Wapiti.

At the best this is a di6cult subject to deal with on paper, and further information is much needed.

CONCLUSIONS. The chief points which emerge from this brief survey may be summarized

as follows :- Antler Pattern.-The antler pattern of the European Red Deer stag tends

to assume the fourth-tine characteristic of the Wapiti in the more easterIy latitudes of its range.

The antlers of the Caucasian stags are extremely variable, some being of the typical Red Deer type, others are of the Wapiti pattern, and some combine certain characteristics of both.

This intergradation of patterns appears to be more manifest in the Caucasus than elsewhere.

PROC. ZQOt. $0~.--1934. 52

798 THE RED DEER OF THE CAUUASUS.

Skull Characters.-The skulls of the Caucasian deer are closer both in size and formation to Wapiti skulls than they are to those of the Scandinavian and Scottish deer. The relatively long and narrow skulls of the Caucasian deer may possibly be a specialization due to an inherited tendency to browse rather than to graze. In view of our limited biological knowledge of these deer, however, this explanation is not put forward as more than mere surmise.

Weight.-From the statistics available it would appear that an adult Caucasian stag is about the same weight as an adult American Wapiti, and this weight is approximately twice as great as that of the deer of Western Europe.

Coloration.-From the limited material available it seems doubtful whether the coat colour differences exhibited by the Caucasian deer and the various forms of Wapiti can be regarded as of more than racial significance.

Voice.-The voice of the Caucasian stag, though differing from the stag of Western Europe, is much closer to it than it is to the ( ( whistle ') or " bugle ') of the Wapiti.

From the foregoing facts it would seem that the Caucasian deer definitely intergrade with the Red Deer and the Wapiti, and in most characters they arc closer to the latter than they are to the former.

Since the range of the Red Deer extends continuously from Western Europe to the Caucasus, and since some male examples from the latter country exhibit unmistakably their close afinity to the typical Red Deer, the Caucasian animals can only be regarded as a local race of Cervus eihphus=C. e. maral ; but if this be so, it must follow (on account of their close affinity to the Asiatic Wapiti and the intermediate characters exhibited by C. e. mural) that the various forms of Wapiti-like deer referable to Cervus should also be regarded as local races of Cervus ekphus. Our present knowledge of many of these deer is so imperfect and the available material so scanty that it is difficult to state definitely which forms should be included, but the following Asiatic and American varieties should apparently be regarded as subspecies of Cervus elaphus :-xanthopygus, baicalensis, songaricus, bactrianus, kansuensis, cashmiriensis, canadensis, occidentulis, merriami, and mnnodes.

EXPLANATION O F THE PLATES.

PLATE I. Fig. 1. Skull and antlers of an old stag from Tartarow, Galicia.

2. Right antler of a stag from Lake Van, Armenia. (The brow and bez tines have been sawn off .)

PLATE 11. Fig. 1. Right 'antler of a stag from the Western Caucasus.

2. Left antler of a stag from the Western Caucasus.

PLATE 111. Fig. 1. Left antler of a stag from the Thian Shan.

2. Left antler of a stag from the Western Caucasus.