26. The Leopards of Africa

THE LEOPARDS OF AFRICA. 543

26. The Leopards of Africa. By R. I. POCOOK, F.R.S., F.Z.S., Unofficial Assistant in the Mammal Dept. of the Natural History Museum.

[Received January 16, 1932 : Read April 19, 1932.)

(Plates I.-IV.* ; Text-figures 1-9.)

CONTENTS.

The Leopard of Morocco and Algeria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

The Leopard of Abyssinia and Erythrea . . . . . . . . . . . . . . . . . . . . . . . . .

The Leopards of Kenya, Tanganyika, and Zanzibar . . . . . . . . . . . . . . The Leopards of the Upper Nile and the Rift Valley ... ~. .........

The Leopard of Sinai ....................................................

The Leopards of Somaliland ...........................................

The Leopards of Tropical West Africa.. ............................... The Leopards of Northern Rhodesia .................................... The Leopards af South Africa .... .................................

Page 544 545 548 552 558 564 573 580 58 1

Summary .................................................................... 588 Explsnation of the Plates ............................................... 591

INTRODUCTION. This paper is an attempt to bring together the scattered bibliography

relating to the large number of geographical races of African leopards that have been described, and to settle, SO far as the material a t my disposal permits, the names, range, and status of these races and the resemblances and differences between them. For this purpose, and following the plan adopted in my paper on the Leopards of Asia, published in 1930t, the subject-matter has been divided into a series of sections, each dealing with the leopards of a particular geographical area, as indicated in the table of contents. A brief summary of the results is given at the end.

The cranial and dental measurements entered in the tables, the dimensions of the skull being in English inches, of the teeth in millimetres, are as follows :-

The total length is from the tip of the occipital crest to the edge of the premaxilla above the incisors. The condylo-basal (cond. bas.) length is from the same point in front to the margin of the occipital condyles. This is a better dimension than the total length, because it is independent of the development of a muscular ridge. The zygomatic width (zygom.) is across the zygomatic arches at their widest point. The mastoid width is across the mastoid processes just behind the auditory orifices. The waist is across the postorbital constriction at its narrowest pcint. The inter-orbital (int. arb.) width is the narrowest point of the forehead. The maxillary (max.) width is taken across the maxillse just above the socket of the canine teeth. The upper and lower carnassial teeth (upper and lower carnal.) are measured along the outer edge of the crowns. The dimension of the upper canine is its anteroposterior length close to the scckct.

For explanation of the Plates, see p. 591. t Jonrn. Bombay Nat. Hist. SOC. vol. rrxiv.

644 MR. R. I. POCOCR ON

THE LEOPARD OF MOROCCO AND ALGERIA. No detailed comparison has apparently ever been made between thc leopards

inhabiting this isolated corner of Africa and t,hose from other parts of the Continent or of Asia ; nor between the leopards occurring in different environ- ments in Barbwy. Analogy suggests the possibility, a t all events, of the discovery of differences between those, for example, found in the mountain- forests of the Atlas and the lower-lying districts or Morocco of Algeria bordering on the Sahara. The only example I have seen which came from Morocco is widely different from the type of the Algerian race ; but neither in the Rritish Museum, nor elsewhere, so far as I am aware, is there sufficient material to settle these questions.

PANTHERA PARDUS PANTHERA Schreb. La F,ntMre Buffon and Daubenton, Hist. Nat. ix. pp. 152 & 160, pl. xi.

(d), pl. xii. (?), 1761. Felis panthera Schreber, Die Saug. iii. pp. 384 & 586, pl. 99, 1777 (based

on the described by Buffon). Pelis pardus G. Cuvier, Mbnag. du Museum, pl. 1801 ; F. Cuvier and Geoffr.

St.-Hilaire, Hist. Nat. Mamm. iv. pt. 65, 1832 ; Loche, Expl. Sci. de 1’Algbrie : Mamm. p. 36, 1867.

Felis pardus barbarus Blainville, OstQgr. ii. 0, Felis, p. 186, Atlas, 11. pl. viii., 1839-1864 (preoccupied name).

Loeality of type.-Algeria. Distribution.-Morocco, Algeria, and Tunis. The specimens referred to in the above-given synonymy came from Algeria.

The descriptions indicate considerable variation in colour. In Buffon’s examples it was described a8 more or less deep tawny (fauve plus ou moins fd) on the back and sides and white below. The Cuviers, Geoffroy St.-Hilaire, and Loche, on the contrary, described the specimens they saw as bright tawny (fauve Claire), straw-yellow washed with grey (jaune paille l&&rement isabelle), or greyish tawny (fauve isabelle). But most of the descriptions and figures indicate the pattern as consisting of very large rosettes, with entire or broken rims, and separated by wide interspaces. But in the old 3 figured by F. Cuvier and Geoffry St.-Hilaire the rosettes are mostly unusually disintegrated. In the female, the type figured by Buffon, the rosettes are “ jaguarine I ’ in size, many of them containing smaller black spots.

Buffon is a little vague about the size, the head and body being, he said, from 5 to 6 ft., and the tail over 2 ft. Assuming that this measurement includes both sexes, the male would be over 8 ft., and the female over 7 ft., in total length, both very large leopards. Loche, for an unsexed specimen, gives the head and body as 4 ft. 4 in. and the tail as 2 ft. 10 in., making a total of 7 ft. 2 in. ; but Cuvier and Geofioy St.-Hilaire give the measurements of an old male as : head and body 4 ft. 3 in. ; tail 2 ft. 6 in. ; total 6 ft. 9 in., the head and body being very nearly the same as in Loche’s specimen, with the tail shorter.

In the British Museum there is a subadult 6, represented by the skin and skull (no. 69.3.5.12), which was formerly exhibited in the Zoological Gardens. It came from Morocco, and was probably the original of the wood- cut of ‘‘ The Morocco Leopard ” in James Greenwood’s ‘ Wild Sports of the World,’ p. 234 (Ward, Lock & CO., 1861). so far as I am aware this is the only figure of the Morocco leopard in eaisbnce ; and since no accounts have


1 Total Locality and sex.

length.

- -1- Morocco, CJ imm. .../ 8.8-

Algeria, $? ad. ......I 8.2

ever apparently been published of an example from that country I subjoin the following description of the one above referred to.

Coat longish and noticeably thick and close, with plenty of underwool. Ground-colour tawny or dusky brownish yellow, without the brightness of the typical savannah leopards of tropical Africa, the interspaces being darkened by the infuscation of the tips of the hairs. The back rather darker, richer tinted, and less grey than the flanks. Fore and hind legs washed with buff to the paws ; the back of the fore-paw below the carpal pad and of the hind leg below the hock unusually pale, greyish white. Upper side of the tail unusually grey, the tawny hue vanishing a short distance below the root. The lower surface of the body and tail and tho insides of the legs clean white. The rosettes mostly small and rather widely separated ; the spots on the paws few, small, and scattered.

In ih general hue this skin doe3 not differ much from the skin of the Senegal leopard referred to below (p. 574) ; but the coat is much thicker and closer in texture.

The skull of an adult Algerian female of this race, which had lived in the Jardin des Plantes, was figured by Blainville. It is fully adult and of large size, larger than the female skulls of all the known African races except the one, described as chui, from the savannah district of the north-eastern Belgian Congo (see below, p. 566), with the skulls of which it agrees tolerably closely in general dimensions; and it shows no signs of the modifications which sometimes affect the skulls of menagerie-kept carnivores. It also agrees very closely in measurements with the female skull of tulliana from Osmanieh in Asia Minor referred to below (p. 547).

The skull of the Morocco specimen in the British Museum is that of an immature male with the basioccipital suture and the other cranial sutures open. It is a normal skull in its shape and proportions, and shows no evidence of having been affected by its life in the Zoological Gardens.

The dimensions of these two skulls are as follows :-

Cond*/ Zygom l:& width.

8.0'5.6 7.4+ 1 5.3

~ _ _ _

In English inches. I

I 1 I _ _ _ I

I- -_I __ -'

______ -

In millim. I

___

THE LEOPARD OF SINAI. m e n Hemprich and Ehrenberg described Felis pardus nimr, referred to

a t greater length below, they mentioned the leopard of Sinai, regarding it as belonging to the same race as the typical example of nimr, a very small leopard from Arabia Felix. They did not, however, secure a specimen from Sinai.

The material I have seen from that country docs not agree with any African or Asiatic leopards known to me, and seems to suggest an undescribed race. I am indebted to Col. C. S. Jarvis for the donation to the Museum of the skull of an adult female and the skin of a young specimen, still with milk-dentition ; and to Mr. R. Hayne for the loan of the skin of an adult of doubtful sex, but probably a female.

546 MR. B. I. POCOCK ON

PANTHERA PARDUS JARVISI POC. (Pl. I. ; text-figs. 1 & 2, a, b . ) Pantbra pardwjarvisi Pocock, Abstr. Proc. Zool. SOC. 1932, no. 347, p. 33.

The adult skin, apparently in winter pelage, has the coat full and very soft, with underwool, the hair of the back measuring just over 4 an inch and on the belly 14 inches long. The general tint, perhaps a little faded, pale creamy-buff on the back, creamy-grey on the flanks, and creamy-white below, the contrast between belly and back being comparatively dight. The rosettcs are well defined, of average size and spacing, with their centres slightly

Text-figure 1.

Skull of adult female (type) of the Sinai Leopard (Paiithera parduojmcisi). x 4.

darkened ; on the belly and inner side of the fore legs the spots are brown. The skin measures : head and body 4 ft. 4 in., tail 2 ft. 6 in., total 6 ft. 10 in. The tail is of normal length as comparcd wit'h the head and body, and the dimensions of the skin, even allowing for some stretching, indicate an adult leopard of average size.

In the skin of the young specimen (B.M. no. 29.12.23.1) the coat is long, over 1 inch on the back, thick, and somewhat shaggy, thus breaking up thc edges of the black spots and obscuring the pattern. The back and upper side of the tail are faintly washed with greyish buff, which becomes pallid high UP on the flanks, thighs, and neck, and passes into white below. The spots


Locality and sex.

-__-___- Sinai, 9 old . . . ... ._.

Sinai, 0 ._. ._. ... ... ...

Dsmanieh, 9 ad. ...

are everywhere black. The colour of this unfaded skin is very like that of the Dwarf Leopard (nunupardus) of Italian Somaliland, but is rather paler. The woolliness of the coat is probably a juvenile character such as is exhibited by many leopard cubs.

From its shape I take it to be that of an oldish female. It is without trace of cranial sutures, but has the teeth unworn and is about the normal length of typical female leopard skulls, but is decidedly robust, with the interorbital area relatively unusually wide, and the forehead depressed between the postorbital processes, which are elevated, making the skull high a t this point ; for a female skull the sagittal crest is well developed, but the occipital crest is rather exceptionally narrow. The mesopterygoid fossa is wide, with arcuate lateral margins, the auditory bullae are small, 0attish anteriorly, and separated by a wide basioccipital space.

In its pallid hue this leopard closely resembles one or two races widely separated from it geographically, but inhabiting similar arid districts with comparatively small annual rainfall. One of these is the so-called Persian leopard ( P . p . saxicolor), the origin of the error that the Snow leopard or Ounce occurs in Persia ; the others are the leopards of parts of southern Arabia, of Erythraea, and of Somaliland. The Sinai leopard is, however, decidedly larger than those southern races, and thicker in the coat, at least, than the Somaliland leopards. It is apparently more nearly akin to the northern Persian form, which intergrades with the h i a Minor race ( P . p . tullianu). But the coat of the Persian leopard, when in good condition, is noticeably thicker, longer, and more woolly, and when in poor condition, presumably in summer, is harsher. I have, unfortunately, no skull of typical saxiwh to compare with that of the Sinai leopard ; but an adult female skull from Osmanieh in Asia Minor, assigned to tullknu, differs in several particulars from the Sinai skull. It is certainly considerably younger and probably hardly fully developed ; but it is noticeably larger in all its dimensions as the following table, containing the measurements of the Osmanieh skull, of the type of jarvisi and the teeth in the skin belonging to Mr. Hayne, shows.

The skull sent by Col. Jarvis is unsexed.

~ ~

~ o t a l “c::. Zygom lendh’llength. width.

1- -- 7.6 I 7- 5

- I - -

8.2 7 5 5.2

I In English inches.

Max. I Upper width.1 carnal.

1.9+ 22 ,

---- Upper oanine

13

1 In millim.

Int. orb.

width.

1.6 + -

-

1.6 +

The occipital crest of the Osmanieh skull is over a of an inch wider than in the Sinai skull, and the auditory bullae are considerably more inflated anteriorly. But owing to the known variability of leopard skulls, no definite conclusions can be drawn from the evidence of the two described above. The differences mentioned are, however, worth recording for future reference in case additional skulls of these leopards are collected.

pB00, ZOOL. S0C.-1932. 36

548 MR. R. I. POCOCK ON

TEE LEOPARDS OF ABYSSINIA AND ERYTHRIEA. The first authors to give a technical name to an Abyssinian leopard were

Hemprich and Ehrenberg, who, in 1833, described a small pale specimen from that country as Felis pardus nimr. With it they associated under the same name a similar leopard from Arabia Felix ; and since this specimen formed the subject of the plate, and in the main of the description of nimr, Arabia Felix has been regarded as the locality of the type of nimr by writers of the present century. Now, for the most part, the possibility of the occurrences of nimr on both sides of the Red Sea has been ignored mainly, apparently, on geographical grounds. But since there are some features connected with the description and figure of typical nimr from Arabia Felix suggesting that i t may be racially distinguishable from the Abyssinian leopard associated with it, i t seems that the South Arabian leopard must be set aside as an unknown form until specimens come to hand for examination.

In 1863 a large dark leopard from the Highlands of Abyssinia was described by Brehm, who adopted for i t the inadmissible name Felis poliopardus.

At the beginning of the present century Camerano described some leopards from Erythrsea (Boll. Mus. Zool. Torino, xxi. no. 543, p. 5, 1906, and in Abruzzi’s I1 Ruwenzori,’ i. Zool. p. 108, 1909). The ground-colour, he said, varies from clear yellow brown to reddish-yellow brown, the rosettes being very variable in size, spacing, and the extent to which they are broken up. In 1909 he published a photograph of a skin, with tolerably large rosettes (‘ I1 Ruwenzori,’ pl. iv. fig. 1). He also gave some measurements of four male skulls and reproduced LL photograph of one of them (tom. cit. pl. ii. fig. 3). This figure has all the characters, so far as can be judged, of a normal female leopard skull, differing considerably by its smaller size, shape, and deficient muscular development from the skull of the typical male of the Ruwenzori leopard, which he figured alongside it for comparison, a skull which agrees in all its characters with that of the male of the ordinary Kenya and Tanganyika leopard usually cited as suahelim. The dimensions Camerano gave of four adult male Erythraean skulls all point to the same conclusion. In their size, shape, and the size of their teeth they resemble, more or less, female skulls of ordinary African leopards.

Gnnberg a1.m briefly referred to some leopards from Gheleb in Erythrea (Sv. vet. Akad. Handl. xlviii. no. 5, pp. 76-80, 1912). He had four skins which he described as less rufous than the brightest skin of British Eaat African leopards, but not so pale as the palest. Unfortunately he published very few particulars about the skulls, quoting only the condylo-basal length of an adult male as 7.7 inches and of two females as 7 and 6.7 inches respectively. The points to be noticed about these measurements are that the difference in size between the male and the largest female skull is comparatively small, indicating failure on the part of the male skull to acquire the marked secondary sexual features characteristic of the skulls of ordinary male leopards; also that this male skull is probably about the same total length as the largest male skull measured by Camerano. It may be a little bigger, but it is a small male leopard skull. On the other hand, the two female skulls should prove to be about average in size, if the occipital crest is normally developed.

In 1923 De Beaux described an adult male, the type, and a female from Keren in the Bogos district as antinorii, and definitely included in this race the three smallest specimens recorded by Camerano, leaving the largest as doubtfully referable to it. Lirnnberg’s specimens he mentioned as indicating the existence of a small leopard in the country.


The colour of the type, a mounted specimen, was described as somewhat bright or light (chiaro), and the rosettes as small, numerous, interrupted, rather thin rings, with the centres perceptibly a little darker than the ground- tint. The specimen, as mounted, measures approximately: head and body 3 ft. 6 in., tail 2 ft. 3) in., total 5 ft. 9) in., a very small adult male leopard.

The skull is ‘‘ small and globose,” and since i t has a wide “ waist ” it clearly resembles the skull of a normal female in defective muscular development, like the skull figured by Camerano. Its total length is almost the exact average of the four male skulls measured by Camerano, but the carnassial teeth are larger. The adult female skull is only a little smaller than that of the male, as in the examples recorded by Liinnberg.

This leopard is only known to me from its bibliography, but since i t seems to be a distinguishable form, although closely related, as De Beaux detected, to the dwarf Somaliland leopard, I adopt De Beaux’s name for it.

PANTFIERA PARDUS ANTINORII De Beaux. Felis pardus wimr Hemprich and Ehrenberg, Symb. Phys. pt. 2, 1833 (in

Felie pardus a n t i w i i De Beaux, Atti SOC. Ital. Sci. Nat. pp. 276 & 278,

Locality of type.-Keren in Bogos, Erithraea. Distritnhm-Erithraea and Abyssinia. A small leopard, but varying in size, pattern, and colour, the colour ranging

from clear yellow brown to reddish yellow brown, the typical form from Bogos being apparently the palest and the smallest, the total length being only about 6 ft. Skulls of males only exceed to a small extent those of the females in size, are comparatively poorly developed muscularly, and in dimensions and shape resemble tolerably closely, apparently, the skulls of ordinary female leopards.

The measurements of the skulls given in the table (p. 652) a~ from data supplied by De Beaux, Camerano, and Lonnberg. It may be noticed that the teeth of the male from Bogos are considerably larger than those of the three smallest, recorded by Camerano, which De Beaux refers without hesitation to the same race, getting over the difficulty by assuming a difference in the method of measurement. The point is of some interest because the teeth of Camerano’s specimens, if correctly measured, agree very closely with those of the dwarf Somaliland leopard (nanupardus). Yet De Beaux diagnosed a d i w i i as differing from nampardus in the larger size of its teeth.

The largest of Camerano’s male skulls and the one from Gheleb mentioned by Liinnberg no doubt belonged to leopards intermediate between typical antiwii and the large leopard (adusta) of the highlands of southern and western Abyssinia, and those of tbe lowlands to the south of that country towards Kenya Colony.

part ; Abyssinian specimen).

1923.

PANTHERA PARDUS ADUSTA POC. Felis ’pardus Schwarze varietiit-Hedgln, Peterman’s Mittheilungen, p. 14,

1861.

(not poliopardus Fitz., 1855) ; SB. Kais. Akad.-Wien, 54, 1. p. 555, 1866.

Leopardus poliopardus Brehm, Reise nach Habesch, pp. 106-108, 1863 Schwarzer Leopard, Fitzinger & Heuglin,

Panthara nimr niger Fitzinger, SB. Kais. Akad. Wien, 58, 1. p. 466, 1868. 36*


Felis pardus suahelicu Hollister, Bull. US. Nat. Mus. 99, p. 171, 1918 (in part ; specimen from Adisababa) ; De Beaux, Atti SOC. Ital. Sci. Net. xlii. p. 276, 1923 (specimen from Shoa).

Panthera pardus adustu Pocock, Ann. & Mag. Nat. Hist. (9) xx. p. 212,1927.

Locality of type of adusta.-Highlands of Abyssinia. Distribution.-According to Heuglin, Highlands of Southern Abyssinia. Notes on the synonymy.-Brehm suggested that this large dark Abyssinian

leopard possibly belonged to the form Fitzinger had described and figured as poliopardus in 1855 (SB. Kais. Akad. Wien, xvii. p. 295) ; but in 1868 Fitzinger declared the type of poliopardus to be probably a menagerie-bred hybrid between a jaguar (Panthera o m ) and an ordinary black leopard. The name niger, which Fitzinger in 1868 gave to this Abyssinian leopard, was inadmissible owing to its having been previously applied by several authors to the black variety of the jaguar. Hollister, who had a single black skin from Adisababa, and De Beaux, who had a skull from Shoa, considered the animal to be the same as the ordinary leopard of Kenya and Tanganyika. Hence the name adusla, which I proposed in 1927, seems to be the correct title.

The type of adusta from the “ Highlands of Abyssinia ” (Ernest Poland ; B.M. nd. 28.11.18.1) is dusky, olivaceous-tawny on the flanks, cheeks, and legs, fading on the middle of the belly to buffish white and to greyish white on the hind belly and inside of the thighs. The summit of the head is blackish, and from the shoulder backwards there is an irregular black band, from 3 to 4 inches wide, which narrows on the loins and does not reach the root of the tail. Externally to this band the rosettes are distinct, large, moderately widely spaced, and have very dark, sometimes wholly black, centres ; on the thighs they are solid, and the hind leg below the hock is sooty black,

The other specimens in the Museum came from Wallega, western Abyssinia (E. Gerrard, B.M. no. 31.1.4.1.2). One of them (31.1.4.1) is much the same dusky olivaceous hue as the type, but the ground-colour is slightly more buffy, and the belly is a cleaner white. There is, moreover, no dorsal black band, the rosettes along the spine being well defined. On the flanks the rosettes are as large as in the type of adusla, but their centres are not so black.

The back is jet black, but the flanks are very deep brown, so that the rosettes, which seem to be smaller, are traceable. The whole skin is sparsely speckled with quite white hairs. It is apparently that of a female, at all events the upper canine tooth in the jaws attached to the skin measures only 12 mm. in antero-posterior basal length.

In addition to the type, a series of five skins in Messrs. Poland’s store, and traded from the “ Highlands of Abyssinia,” cover the variations in colour exhibited by the type, and the two examples from Wallega described above, three of them being “ black ” and two spotted, one of the latter having a black band down the back, the other being merely blackish on this area so that the spots are traceable, and in this example the rosettes generally are closer together, more jagged in outline and inclined to fuse, the interspaces being sufficiently narrow to be almost describable as a network. The rosettes in this race vary considerably in size, being in many specimens larger than in typical suahelica, in others about the same size, as recorded by Hollister in the ‘‘ black ” example from Adisababa he assigned to suaheliea.

The writings of Brehm and Heuglin attest the prevalence of this dark leopard in the Highlands of Abyssinia ; and in the sixties of the last century

The British Museum has three skins of this leopard.

The second specimen from Wallega (31.1.4.2) is a “ black leopard.”

THE LEOPARDS OF AFRICA. 55 1

Total.

ft. in. 7 l l t 8 2

8 ot 7 7 6 Ilk 6 6 6 6

---

Major Leveson (‘ Sport in Many Lands,’ p. 491) recorded the shooting of a “ black ” leopard in the passes above Ailet in that country. Writing, moreover, comparatively recently, of the leopards of Somaliland and Abyssinia, Col. H. G. C. Swayne said : “ The best of all, in which the spots are so large and so closely planted as to give the centre of the back almost a black appearance, are obtained from the Highlands of Abyssinia. . . These very handsome dark leopards skins are used in thousands by the Abyssinian soldiers as cloaks.” His further statement that “ there is a very large animal inhabiting the hills which would be described by Indian sportsmen as a panther ” is borne out by the following measurements of the skins I have seen, all of which are native- prepared, and apparently but little, if at all, stretched ;-

1

Locality and sex.

Wallega, d . . . . . . . . . . . . . . . . .

‘I AbyRsinia,” . . . . . . . . . . . .

“ Abyminia,” d . . . . . . . .

“ Abyssinia,” 2 . . . . . . . . . . . . . .

“ Abyssinia,” ? ............... I ‘ Abyssinia,” $! . . . . . . . . . . . .

Wallega, 0 . . . . . . . . . . . . . . .

Head and body.

ft. in.

6 0

~ - -

5 n

4 9k 4 7

4 2t 3 9

3 8P

Tail.

ft. in. 2 l l h 3 2 3 3 3 0 2 9 2 9

2 9t

~-

A male skin from Mareb, recorded by De Beaux, measured : head and body 5 ft. 31 in., tail 2 ft. 8 in., total 7 f t . 11: in. It was apparently longer in the head and body and shorter in the tail than the largest males in the table. Possibly the difference is due t o difference of opinion as regards the origin of the tail.

De Beaux published the dimensions of a skull of an adult d from Shoa, which he identified as swlheliea * ; and there is in the British Museum the skull of a 8, not fully grown, with the basioccipital suture open, from Gabela (C. A. Willis, B.M. no. 19.11.13.1).

The measurements of these two skulls agree closely with those of the leopard of Kenya and Tanganyika. Probably the two races intergrade in the lowlands to the south of Abyssinia ; but the typical forms of the two are easily distinguishable by colour. It probably also intergrades with the smaller paler leopards of Erythraea and Somaliland.

The following table of available measurements of skulls of antinorii and adustu, taking the condylo-basal length of the skull as the standard, shows that even the young-adult example from Gabela is noticeably larger than the largest example recorded by Lonnberg from Erythraea, and still larger than the type of antinorii. It shows also that the “ waist ” of the type of the latter is much wider than the interorbital width and actually wider than in the considerably larger skull of adusta from Shoa, in which the waist is narrower ~ h a n the interorbital. There is not a great difference between the type of antinarii and the examples of adusta in the size of the carnassials, although the canine is smaller ; but the average of all the teeth of antinorii is less.

* Also when writing of the race he named aritznoiir he recorded the length of the aocket of l h i s waR 17 mm., a8 in the

Of skulls of this race little is known.

the upper canine of the large example from Mareb, above referred to. Shoa specimen.

652 bfR. R. I. POCOCK ON

Waist width.

__ Locslity and sex.

, Total ilength :::: M ~ X .

width. width

--I-- antinorii. Gheleb, ad. . . . . . Erythrsa, d ad. . . Erythrrea, d ad.. . Erythrsa, d ad.. . Erytiinea, d ad.. . Bogos (type), d a d Gheleb, 2 ad. . . . . . Gheleb, 0 ad. . . . . . Bogos, 0 ad. ..... ----

aduatn. Shoa, d ad .........

Gabela, d yg. ad. Mareb, 8 nd. .....

W a b g a , 0 ad. ..

14 12 11+ 13

12

17 17 15 12

In English inches.

- - -

17 -

16 18+

15.5

_ _ - - --

19 -

19 -

Cond. ' Mas-

-I---- - 73 1 5, - 1 5 -

4'6 - -

- , 4.7 7 1 4.8 7 1 - 6.7 I -- 6.3 4.6 + - -- ' 6.2

_ _

In millim. ! I

- 23 22 20 21 24 - - 21.3 - 25

25 -

- __

THE LEOPARDS OF SOMALILAND.

There is evidence that the leopards of Somaliland are referable to three distinct races, as pointed out by Dr. Drake-Brockman in 1910. Thanks to him and to other donors of specimens to the British Museum, I have been able to examine several skins and skulls of both types.

The first to be described is the dwarf leopard of Italian Somaliland.

PANTHERA PARDUS NANOPARDUS Thos. Felis pardus nonapurdus Thomas, Ann. & Mag. Net. Hist. (7) liv. p. 94,1904;

Drake-Brockman, Mamm. of Somaliland, p. 17, 1910 ; De Beaux, Atti Soc . Ital. Sci. Nat. xlii. p. 276, 1923 ; id. Atti Soc. Ligust. iii. pt. 2, p. 59, 1924.

Locality of t y p e . 4 0 miles west of Gorahai. Distribution.-Italian Somaliland. A small, short, and smooth-coated leopard with the colour of the dorsal

surface a pale dull olivaceous or greyish-buff, gradually fading on the flanks and passing into white on the belly and limbs, without any of the bright tawny- yellow of typical leopards. The rosettes are smallish, well defined, moderately close set, and with darkened centres showing up especially on the flanks. The tail is of normal length, about two-thirds the length of the head and body.

There are only three skins of this leopard in the British Museum from Italian Somaliland, namely, an adult female (type) collected by Major Dunn, 40 miles west of Gorahai (no. 4.9.5.34), a youngish female from Garabwein, near Obbia (no. 12.12.18.17), collected by Dr. Drake-Brockman, and a still younger, unsexed specimen frem Garrero, in Dolbahanta, collected by H. E. Johnstone Stewart (no. 4.7.30.1). These skins are all very similar in colour, pattern, and length of coat.

There are no available flesh measurements of adult skins. The stripped skins* of the adult 9 and of an immature from Gorahai were measured

The male skin wae kept by the collector and is not available for examination, and the female

(Text-fig. 2.)

skin was subsequently made up in the stereotyped method for preservation in store.

"HE LEOPARDS OF AFRICA. 563

by Thomas as follows :-8, head and body 3 ft. 10 in,, tail 2 ft. 2 in., total 6 ft. ; 9, head and body 3 ft. 7 in., tail 1 ft. 11 in., total 5 ft. 6 in. It is of course possible, perhaps probable, that these xkins were somewhat stretched in the stripping. The only specimen with flesh measurements is the young female from Garabwein. The head and body were 2 ft. 11 in., the tail 2 ft., total 4 f t . 11 in. The skull, however, shows this animal to be immature. She would certainly have surpassed 5 ft. when adult. Probably ths adult male, when known, will prove to be abclut 6 ft. in total length.

It is fully adult, with scarcely a trace of cranial sutures, and is small, low, and

Text-figure 2.

The only available adult skull of nunopardus is that of tho typical 9.

Skull of adult female (type) of the dwarf Leopard of Somaliland (Panthera pardiis Itanopardus). X &.

elegantly Rhaped, decidedly long for its width, the waist snd interorbital areas being narrow as compared witb those of ordinary femalu leopard skulls. The occipital crest is well developed, but there is no sagittal crest, the temporal ridges forming a lyriform pattern 3 of an inch wide on the parietals. The teeth are small. A comparison of the figure of this skull (above) with that of the Sinai leopard (p. 546) shows the marked differences between them. The skulls are approximately of the same age.

The male skull from the same locality (no. 4.9.5.34) is immature, with the median frontal, the interparietals, and the basioccipital sutures still open. It is a little shorter than the female skull, but is a little wider everywhere and much wider in the waist. Its muscular development is poor, and the temporal ridges on the parietal are 1 inch apart. The teeth are larger then


in the female. From the structure of this skull I suspect that the adult male skull of nanopardus will be found to be like that of antirorii in its defective muscular development.

The skull of a young female from Garabwein, near Obbia (It. E. Drake- Brockman, B.M. no. 12.12.28.27) is about the same age as the male skull and very like it, although smaller. The temporal crests are over 1 inch apart on the parietals. The bulls are much less inflated than in the specimens from Gorahai.

An immature unsexed skull from Garrero, Dolbahanta Coy. (Johnstone Stewart, B.M. no. 4.7.30.1), is very like the other immature skulls described, but has a wider mesopterygoid fossa, and small bulls like the skull from Garabwein. These three immature skulls have the permanent teeth fully erupted. In this respect they differ markedly from a skull similar in length and width from the Masai Reserve, Kenya, which is most manifestly that of a cub with milk dentition and the palatal sucking-ridges quite strong.

Typical nawpardus appears to differ from antinorii in being a Little smaller, paler, and greyer in colour and in having smaller teeth.

In the foregoing account of nanopardus the examples assigned by De Beaux to that race have not been considered because, judging from his description, they are not typical, but in some respects transitional between i t and the larger race from British Somaliland. He had three specimens ; an adult pair came from Mogadiscio on the coast of Italian Somaliland, midway between Obbia and the Juba River. The female was described as ochraceous tawny on the back, the sides of the body and external surface of the limbs being light buff and the underside pure white. The male, which De Beaux thought might be artificially discoloured, though t h e evidence for this is not satisfactory, was more intensely coloured, amber brown. Even the female of these examples seems to be darker and brighter in tint than typical nanopardus, approaching in that particular the leopards of Kenya Colony. This is also true of a young male he identified as nanopardus from Gelib in Jubaland. He described the colour of the back and of the centres of the rosettes as ochraceous tawny, the head being ochraceous buff, the flanks and outside of the limbs light buff, and the underside pure white. The flesh measurements of the adult male from Moga- discio were : head and body 3 ft. 6 in., tail 2 ft. 4 in., total 5 ft. 10 in., indicating a leopard about the same size as the male from Gorahai. But the example from Gelib, although not full grown as indicated by its skull, was larger in the head and body, which measured 3 ft. 11 in., the tail being 1 ft. 11 in., making a total of 5 ft. 10 in. This leopard when full grown would have surpassed a total of 6 ft., probably considerably.

The skulls of the pair from Mogadiscio also bear out the conclusion derived from the skins, that these leopards are not typical nampardus. They are stated to be ‘‘ adultissima ” and are noticeably larger than the skulls of naw- p r d u s , the female skull being bigger in all its dimensions than that of the type of nanopardus, of apparently about the same age. But both the skulls from Mogadiscio have small teeth, smaller than those of the skulls of typical nampardus. De Beaux also referred to the skull of a young male from Gelib in Jubaland, which he identified as namprdus. This skull also has the small teeth of the Magadiscio examples. In its total length i t is very slightly shorter than the young male skull of nanoprdus from Gorahai ; but apart from identity in the maxillary width i t is otherwise larger in every particular ; and since the approximate equality in length between its condylobasal and total lengths suggests that i t is a younger skull than the skull from Gorahai, it is impossible to doubt that it is potentially a larger skull, thus agreeing with the Mogadiscio specimens.


Upper Sheikh, d yg. ad. .........

Upper Sheikh, Q ad.

Guban, 9 ad.

. . . . . . . . . . .

.....................

-

It seems probable that the small size of the teeth in these examples from M gadiscio and Gelib influenced De Beaux in distinguishing the Erythraean dwarf leopard a n t i w i i from the Italian Somaliland nunopardus. A t all events he briefly diagnosed a n t i w i i as distinguishable from nanopardus by its longer teeth. The difference is not so marked in the case of typical nanopardus.

The suggestion made above that the immature male skull of nunopardus, from Gorahai, would have exhibited poorly developed secondary sexual characters, if i t had reached maturity, is borne out by the male skull from Mogadiscio, in which the ‘‘ waist,” as shown in the subjoined table (p. 557), is considerably wider than the interorbital area, a feature characteristic of most female leopard skulls.

f t . in. It. in. ft. in. 4 0 2 7 6 7 1

5 9 I 3 31 2 51

3 54 2 4 t 5 10 ,

PANTHERA PARDUS BROCKMANI, subsp. nov. Felis purdus Drake-Brockman, Mamm. of Somaliland, pp. 14-16, 1910.

Locality of type.-Upper Sheikh in the Golis Range. Distribution.-British Somaliland, apparently as far south as Jubaland. Larger, and on the average more brightly coloured, than nunopardus, which

i t links with the southern Kenya leopard. Skulls like those of the latter, but smaller, showing well-marked secondary sexual characters, the skull of the male muscularly well developed, that of the female robust and broad, as compared with the skull of nunopardus.

By Drake-Brockman this leopard was quite correctly, in my opinion, distinguished from nanopardus by its superior size.

The following are the flesh measurements of three specimens recorded by him.

These dimensions indicate a smallish leopard. Judging from its skllll, which is nearly full sized, the male would at most have exceeded slightly 7 ft. in total length when completely adult.

Drake-Brockman described the colour of this British Somaliland leopard as tawny yellow, varying according to altitude, those from the Golis Range, referred to as “ hill ” specimens, being darker than those from Guban, “ lowland,” specimens. Nevertheless, the adult unfaded female from Upper Sheikh, presented to the British Museum, is practically indistinguishable in colour and pattern from the examples of nanopardus I have seen, being describable aa a pale greyish-tawny leopard with small close-set-rosettes. The skin of the male from the same locality, which is in his possession made up as a rug, is, however, slightly more brightly tinted, and is no doubt somewhat faded. He has also kindly lent me for examination two skins from Ogo, 4000 to 5OOO ft. alt. One of them is very much the same tint as the unfaded fern48 from Upper Sheikh, the other is decidedly brighter, and clearly apwoaches the southern Kenya leopard in that respect. I have also seen the skin of an


adult female shot at the junction of the Tug and Shebeyli Rivers. This skin is a dull greyish tawny, but its donor, Mr. F. Lort Phillips, tells me that it is no doubt faded since it was exposed to the light for many years made up as a rug.

In Rowland Ward’s ‘ Records ’ are entries of leopards from Somaliland ranging from 6 ft. t o about 74 ft. in total length. But since the precise localities, ages, and sexes of the specimens are unknown, i t is impossible to identify them racially.

The skull of the male from Upper Sheikh, with the interparietal bone still unfused, I regard as that of a subadult (the basioccipital is shot away). It exhibits well-marked secondary sexual characters due to muscular development. The waist is slightly wider than the interorbital area ; but the waist would probably have become the narrower of the two, if the animal had lived a few more years, and the zygomatic arches would have expanded concomitantly. It resembles a small skull of the better-known leopard of Kenya and Tanganyika, and of the large dark Abyssinian form (adusta). In the size of its teeth it closely resembles the typical form of antinorii from Bogos ; but although not full sized it is decidedly larger in every detail except the “ waist,” which is noticeably narrower.

The skull of the fully adult female from Upper Sheikh appears to be about the same age as the typical female skull of nunopardus. It is a little longer and, except across the mastoids, is wider in every particular, exhibiting the generally robust build, with the broad waist, characteristic of ordinary female leopards. The temporal ridges vanish dorsally over the parietals, the sagittal crest being undeveloped, except posteriorly over the interparietal, where i t forms an erect crest. The canine tooth is equal to that of the male skulls of narwpardus in anteroposterior basal length, and noticeably larger than in the female of that race.

The skull of the female from the junction of the Tug and Shebeyli Rivers (Lort Phillips) is also fully adult and about the same age, size, and shape as the last ; but the temporal ridges are lyrate, and there is no interparietal sagittal crest. In this it resembles the tspe of nunupardus. The carnassial teeth are about the same size as those of male and female nunopardus, but the canine, as in the Upper Sheikh female, exceeds that of nunopardus.

A third adult female skull from Beila Majeasa in Jubaland (Dr. Bevan, B.M. no. 31.1.28.8) probably belongs to this race, but it is a little larger than the last two, and is very nearly as large as the smallest female example of the Tanganyika race recorded from Kigoma in the table below. It is like the female skull from the Tug-Shebeyli River in the development of the sagittal crest and has the same sued canine tooth. The measurements of these three skulls reveal no differences from those of female skulls of antinorii.

In the table of cranial measurements of Somaliland leopards (p. 557), the examples from Mogadiscio and Gclib assigned by De Beaux to nunopurdus have been separated from the typical specimens of that race to indicate clearly their superiority in size and the intergradation they exhibit between nunopardus and brwkmani, except in the waist-width of the males.

It may be added that Hollister assigned the Somaliland leopards known to him from skins and skulls to the Kenya race which he called suahelica. There is, indeed, no reason to doubt that brockmani intergrades with this larger southern form. A skull from Somaliland in Ward’s ‘Records,’ entered as measuring in total length and width 9 by 5.7 inches, belongs probably to the Kenya form. This also applies no doubt to a very large leopard from Hargeiea

'PHE LBOPARDS OF AFRICA. 557

Locality and sex.

----- 7m1lopatrlur.

Gorahai, d imm. . Dolbahanta, d yg. Gorahai (type),

Garabwein, Q imm. 0 ad.

which from its stripped skin was estimated by Drake -Brochn as not less than 8 ft. in total length.

In reply to my inquiry for further particulars about this leopard from Hargeisa, Dr. Drake-Brockman informed me in a letter, dated Nov. 30th, 1931, after I had finished the account above printed of the Somaliland races, that the leopard in question was not shot by himself, but that he saw it measured ; and he added that he secured a far finer one close to the coast near Bulhar. It was not measured in the flesh, but when the skin was stripped and pegged out, good care being taken not to stretch it, i t surpassed 9 ft. ; and its skull was of enormous size, almost as large as that of a tiger. The skull was, unfortunately, mounted in the skin and is not available for measurement.

These large leopards, represented by the Hargeisa and Bulhar specimens, frequent the low hills and undulating country from the Feanar Highlands to the coast near Bulhar and to the west and north of it. Dr. Drake-Brockman believes them to be the biggest leopards in existence. They are probably, he thinks, related to the Abyssinian leopards, being heavily built, although much lighter in colour, cases of melanism being frequent in the Abyssinian animal. They are the only leopards which occasionally kill camels in British Somaliland. He has never known the other leopards of the country, nunopardus and the Sheikh variety, which are distinctly smaller and lighter, kill camels, only sheep and goats.

Finally, he states that he has for years known the following types of leopards in British Somaliland :-(1) nunopardus ; (2) the intermediate Sheikh variety ; (3) the " Hargeisa " variety. This epitome by a systematic zoologist, acquainted with the animals in the field, completely confirms my conclusions expressed above.

The following table records the dimensions of the skulls of nanopardwr and brockmani which I have seen, and also the published measurements of the examples referred to nunopardm by De Beaux.

_- Tota

lengtl -

6.5 6 - 6.7

6.1

Mogadiscio, d ad.. Gelib, (r yg. . . . . . . Momdiscio, 0 ad..

- ~ _ _ - bmckmatti. Upper Sheikh, d

I I

YS. ad. Uppr Sheikh, Q

ad. Tug-Shebeyli, 0 ad. Majeam, 9 ad. . . .

I

8.3 6.4 7.3 -

8.3

71

7.0 7.3

In English inches. 1 In millim. I

Cond. bas.

ength

6.1 5'6 + 6.0

5'7

7 4 6.3 6.4 +

-

-

7.6

6'5

65 6.7

Zygom. width.

4.1 3 7 4'0

3.7

5.4 i 4.6 + 5 -

5.1

4-5 + 4.4 4%

I 2*6+ I 1.7 - 1.1 + 2.5 ~ 1.6 1 1 + 2.8t 1.3 l - l +

3.2+ ' 1.7+ 1.4+ 28t 11.7i 12 3.0 1 1.6+ , 1.2

1*5+ 1*4+

Max. Upper Upper Lower

~-

1.5 ls+i 10 j 16- -c_---

2.0 ' 23 1 14 1 16

I 1.7 ~ 20 12 I 16

1.8 21 I 12 17

1.9 I 22 12 , 16' I

658 MR. R. 1. POCOCK OXd

THE LEOPARDS OF KENYA, TANQANYIKA, AND ZANZIBAR.

PANTHERA PARDUS FUSCA Meyer. Felis fusca Meyer, Zool. Annalen, i. p. 394, 1794. Panthera pardus fusca Pocock, Journ. Bomb. Nat. Hist. SOC. xxxiv. pp. 307-

316, 1930. Felis (Leopardus) nimr Matschie, Saug. Deutsch. Ost. Afr. p. 69, 1895

(not nimr Hempr. & Ehrenb.). Felis pardus suahelicus Neumann, Zool. Jahrb. Syst. xiii. p. 551, 1900 ;

Heller, Smiths. Misc. Coll. xli. no. 9, pp. 6-7, 1913 ; Hollister, Bull. U.S. Nat. Mus. xcix. p. 171, 1918.

Panthera pardus suahelica Allen, Bull. Amer. Mus. Nat. Hist. xlvii. p. 252, pls. 42, 43, 1924.

Felis pardus fortis Heller, Smiths. Misc. Coll. xli. no. 19, p. 5, 1913 ; Hollister, Bull. U S . Nat. Mus. xcix. p. 175, 1918; Allen, Bull. Amer. Mus. Nat. Hist. xlvii. p. 257, 1924.

Type-locality off usca, Bengal ; of suahlica, selected by Allen, Lake Manyara, northern Ugogo, Tanganyika Territory ; of fortis, Loita Plains, Kenya.

Distribution.-India and Ceylon ; Kenya Colony and Tanganylka Territory, and thence southwards at least into Nyasaland.

In my paper on Asiatic panthers, quoted above, I pointed out that the statements of Blanford, de Winton, Lydekker, and others regarding the difference in pattern between African and Indian leopards are unsupported by the facts ; and since I cannot find any differences either in size, colour, pattern, cranial, or dental characters between ths leopards of India and Ceylon, for which the oldest name appears to be fusca, and those of Kenya and Tanganyika, known for the last quarter of a century as suahelica, I am compelled to cite them under the same subspecific name, since I do not admit subspecific titles in a purely geographical sense.

Since Neumann’s description of his so-called suuhelica is valueless, and the specimens described by Heller and Hollister were from Kenya or districts to the north of that country, it may be useful to give an account of some skins in the British Museum, most of them from Tanganyika Territory, where Neumann’s examples were procured.

These Tanganyika Territory specimens were collected by C. H. B. Grant in the Kigoma District, to the south-east of Lake Tanganyika.

(1) 9. From Lubira, Ukaranga, June 14th (B.M. no. 27.2.11.13). General colour the yellowish tawny, characteristic of most leopards, passing into white below and on the inside of the limbs. Rosettes smallish, about average distance apart, rather narrow-rimmed, nearly complete or broken, with darker centres showing up on paler flanks, the yellowish-tawny intermediate areas with scarcely a trace of infuscation a t the tips of the hairs. For0 and hind paws washed with yellowish, their spots spaced and of average size ; rosettes becoming solid spots on the thighs. Coat not short and sleek, longish and full.

(2) dd. Kasaba, Ukaranga, Sept. 15th (B.M. no. 27.2.11.11-12). Practi- cally identical with the preceding in colour, pattern, and coat.

(3) 6. Kanala Pamba, Uvinza, June 30th (B.M. no. 27.2.11.9). Almost exactly like the preceding, just a shade less brightly coloured, and with the coat a little fuller.

(5 ) 9. Kamelapenta, Uvinza, June 30th (B.M. no. 27.2.11.10). Decidedly less brightly tinted than the others, the rosettes a little smaller and cloeer together. Coat about the same.

(Text-figs. 3 & 4.)

Distal end of underside of tail white.


~ - - ~ - ~ ---- f t . in. ft . in.

........................ I

...........................

Naivasha, 4 2; 2 11;

Mevu, d 4 2; 2 9:

Kabalolet Hill, d 4 2 3 o i

Jnga Farm, Q . . I 3 4 2 54

...............

..................

(6). Kenya Colony, without precise locality ; undated and unsexed ( h d y MiwMillan, B.M. no. 25.6.15.1). Less brightly coloured than most of the Kigura skins, about the same tint as the 9 from Kamelapenta, but with the rosettes more broken and with their centres not noticeably darker than the ground-tint, except low down on the flanks. Coat shorter and smoother. In the breaking-up of the rosettes on the back and their undarkened centres, this skin seems to approach fortis ; but the general colour is evidently much paler.

The above described specimens bear out Hollister’s statement that the skins of ‘‘ suuhelica ” from mony localities in Kenya Colony which he examined were a “ very uniform lot in coloration and size.” The same, however, cannot be said of a very fine series of twelve skins procured by Mr. G . W. Foster *, mostly at 9300 ft. alt., on Mt. Elgon in Uganda, which are of special interest on account of their colour-variations t.

In general brightness of tint nearly half of these are very similar to the skins above assigned to fuscu, but they are not quite so uniform in hue, or in the size and spacing of the rosettes. One of them, an immature male, is paler than the rest, less bright on the back and greyer on the flanks. Its coat, too, is remarkably long, and the large and close-set rosettes are less well- defined on account of the length of the hair.

Most of the rest, one procured a t 3800 ft., are noticeably richer and darker in the general tint of the ground-colour, which is deep orange-ochraceous, reddish bpown by comparison with the paler suahelicu-type. The tint indeed is suggestive of Heller’s description of the type of fortis as “ cinnamon-brown.’’ They are the handsomest and richest hued African leopard skins I have seen.

A third type of colour is exhibited by a single skin which is almost olivaceous tawny in ground-colour, much more like the dusky forest leopard of West Africa (leopardus) and one of the skins of iturensis, below referred to, than it is like the other Mount Elgon skins above described. But it is a little more bully than other olivaceous tawny skins mentioned.

Since none of the many skins of this race in the British Museum was measured in the flesh, I subjoin the dimensions of a few recorded by Hollister from Kenya.

---- ft . in.

7 0

5 9;

I Total. I Headand Tail.

; body. Locality and sex.

The British Museum in greatly indebted to Mr. 0. W. Foster for thie series, procured at the request of Capt. Pitman, the Game Warden of Uganda, who was especially informed of the Museum’s need of the skins of big carnivora from tbat part of East Africa.

A skin he had apparently resembled the Gondukuro leopard described as clnti by Heller, und he identified i t r C that race, although with some hesitation (Acta. Univ. Lund. xxi. no. 3, p. 17,192.5). The Mt . Elgon leopards are in a measure a transitional type. I assign them, provisionally, to the same race as the Kenya leopards because of the eize of their skulls, which are smaller on the average than those of chui,

t Granvik reported leopards a8 common in the dense forests of M t . Elgon.

P

1 D

NO 830306 'I ?I '8AI. 099


The adult d skulls from Kenya Colony measured by Heller and Hollister, and assigned to smhelicu, were under 94 in., the average of the d skulls, according to Allen, being gin. Now one of Heller’s primary characters for distinguishing fortis from suuhelicu was the larger sue of the only known skull, from the Loita Plains, Kenya Colony, which is close upon 10s in. in total length *. But some adult 6 skulls in the British Museum from Kenya and adjoining districts of East Africa are larger than those that Hollister and Heller measured, exhibit considerable individual variation, and put the average length of the skulls higher than Allen’s calculation. For example, two from Kenya are 9-9 and 9.7 in. respectively, a series from Mt. Elgon ranges from 8.6 to 9.6 in., and from Tanganyika Territory from 8.8 to 9-8 in. Hence the largest Kenya skull is only about half an inch shorter in total length than the skull of fwtis, and much less than that in condylo-basal length, and the largest from Tan-

Text-figure 4.

n . Frontal region of ~ l i ~ l l of female Leopard (Panthera pardtts fitsca=fiitohelien) from

L. The same of a normal female from Kigoma. Nyeri with abnoimally narrow “ waist.”

X t .

ganyika Territory, ticketed Kigoma, is almost as long in total length. Moreover, i t is noticeable that this skull (no. 24.8.11.1) is 1 inch longer than a smaller fully adult skull (no. 24.8.11.2) from the same locality. The difference indeed between these two is considerably greater than the difference between the skull of fortis and the largest skulls from Kenya, Mt. Elgon, and Tanganyika Territory.

These data justify the view that in its cranial characters fortis merely symbolises a somewhat exceptionally large individual of this East African leopard, with an unusually robust canine tooth. My table of measurements of the skulls of normal adult d leopards from India and Ceylon, published on p. 314 of the paper quoted, shows that they range in total length from 8.9 to 10 inches, the average of nine skulls being almost 94 inches. In other words, they agree as closely as possible with the skulls of the East African leopard commonly assigned to suuhelicu.

The male skulls vary considerably in other characters than length, more * Heller wrongly claimed this to be the largest leopard skull on record at the time he wrote.

But in 1903 I published a figure and description of a male skull from Cette Cam8 in the Gaboon exceeding the skull of fortis by nearly one inch, and referred to two otber~ from the Name locality slightly longer.

Som

e Sk

ull-m

easu

rem

ents

of L

eopa

rds (

P. p

. fus

ca) f

rom

Ken

ya, T

anga

nyik

a, a

nd N

yasa

land

. _

__

__

~ -.

._

__

__

__

_

__

__

-

-~

-

Wai

st

wid

th.

-.--

Loc

alit

y an

d se

x.

Int.

orb.

w

idth

. --

I Total

length.

-

1 '6

1.7

1.7

1% -

1 '6

16+

1.7

+ 1.7

1.7 -

1.5

1.6

1'8

1'8

1'5

+ 1.6

1-7

1'7

1.7

1.7

1.7

1.8

1.5

1.8

1.6-

-___--

I--

1

Loi

ta P

lain

s (fovtin), d

ad.

....

..

Ken

ya,

d ad

. ...

......

......

......

. ,,

d ad

. ...

......

......

......

...

,, d

gg

......

......

......

......

M

t. E

lgon

, 6 ad

......

......

......

....

,, ,,

d ad

....

....

....

....

....

..

,, ,,

6 ol

d ...

......

......

......

,,

,, 6 ol

d ...

......

......

......

,)

,) d

yg ).

_. ....

....

....

....

..

Kig

oma,

T.T

., ad

. ...

......

......

((

da

d ...

....

....

....

. U

krrv

anys

. T.T

., d

ad.

....

....

....

D

ar-e

s-Sa

laam

, T.T

., 8

ad.

......

Si

ngid

da, T

.T.,

CJ ad.

......

.....

Mila

nji,

Nya

sal.,

ad

. ...

......

...

Fort

Man

ning

, Nya

sal.,

d ad.

Ken

ya,

$? a

d.

......

......

......

......

N

yeri

, Kenya,

9 ad

...

......

......

I It

. Elg

on,

0 ad

. ,,

,, 9

suba

d.

......

... ..

....,

,, ,,

9 su

bad.

...

......

......

' K

igom

a, T.T., Q ad.

......

......

... )

,, 9ad .

....

....

....

... I

...

......

......

......

...

,, 2

nd

....

....

....

....

I ...

......

... I

....

....

...

Uvi

nza,

T.T

.. 9

suba

d.

Mila

nji,

Nya

ssl.,

9 4;

,, 2

nd

....

....

....

....

I ...

......

... I

....

....

...

Uvi

nza,

T.T

.. 9

suba

d.

Mila

nji,

Nya

ssl.,

9 4;

lk4

9.9

9.7

8.1

9.6

9'2

8.8

8.6

7.9

9.8

8.8 -

94

90

9.4

9.1

7.8

7.9

7.8

7.9

8.0

7.3

7.6

7.5

+ 7.

7 -

-

lond

. bas

. length.

9.4 +

9.1

8.8

7.4

86

8.6

8.0

7.9

7.4 -

8.8

7.9

8.6

8-0

86

84

7.1

7.3

74

7.3

7.3

+ 69

6.8

67 +

7.1

---

-

-

I ! In

Eng

linh

inch

es.

Zyg

om.

wid

th.

--

63 -

6.7

6.2

4.8

6-

60

5.8

5.7

5.1

60

5'3

6.1

5'8 -

57 +

5'8

4.8

5-1

5 4.9

5.1

+ 5'1

48

4.9 -

4.4

4.9

-

Mas

toid

w

idth

.

4t

3.9

3.8

32 +

4.1

-

4'0 3.6

3.5

3.8

+ 35 -

3.5

3.7

+ 3.

7-

3.8

3.6

3.0

3.3

32

3.2

3.4

3.2

+ 3.2 -

3.2

3.0

3.3 -

-

I-- ._

__

__

_

1-7 +

1.8

2.8

1.2

1.6

1.5

1.6

1.6

1.3

1.5

1.5

1.5

1.7

1.7-

1'7

1.5

1.4

1.3

1.3

1.3

1'4 I .3

1.3

1.2

1 '5

1.3-

Max

. 1

Upp

er

wid

th.

carn

al.

--I-_ 2'6-

2.4

2,5

1 2.0

23

22

2.2

2.1

2.3

2.1

2.2

2.3

2.3

2.8

2.2

1.9 +

2.0

1'9

1.9

2.0

,

1'9

1.8-

~

2.1

2.2

'

2'1

1 1'9+

,

27 -

27

27

26

26

26

25

25

23

24

23

26

26

24

27

24

22

24

25

23

24

25

23 -

22

23

24

In m

illim

.

Upp

er

cani

ne.

18 +

17

16

15

16

16

17

15 +

15

15

17

16

16

14

14

16

12

15-

14

14

15

13

13

13

12

14

---

__

-

-

Low

er

carn

al.

19 +

20

20

19

20

19

19

18

'

18

19

i 19+

18

19

18

15

18

19

16

18 -

18

17

16

' 1

--

18

I 16

j

18

i

THE LEOPARDS OF AFRIOA. 663

particularly in the size and shape of the auditory bullao. On p. 660, text- fig. 3, c-f, are given sketches of the base of the skull of the larger of the two male skulls from Kasulu, near Kigoma, and of the example from Sengidda to illustrate the range in the variation of the bull%. All intermediates between them occur in Tanganylka Territory, the smaller specimen from Kasula having considerably less expandpd bullae than the larger, although not so d u c e d as in the specimen from Sengidda. The series entirely bears out Hollister’s statement that variation in the size of the bulle is not a character of systematic importance, as Heller supposed.

The variation in the size of the female skulls is less than in those of the males, the greatest amounting to just under 2 of an inch in total length, as illustrated by the two juxtaposed examples from Kigoma. Otherwise they ars tolerably similar, apart from the specimen from Nyeri, presented by Capt. Murray. This skull is unsexed ; but has been identified as a female by the sum total of its dimensions. Although only just adult, it has a remarkably narrow, constricted waist, only as wide as its interorbital space instead of being considerably wider, as .is usual in female skulls. The waist, indeed, is as narrow as in the much smaller skull of the adult female of the Somaliland dwarf leopard, nanopardus. The postorbital processes are correspondingly prominent, the two features combined imparting to this skull a singularly 6 aspect.

Note (May 11, 1932).-During the passage of this paper through the presa, the Museum received two female skins, with skulls, collected by Mr. A. T. Hopwood, of the Geological Department, a t Olbalbal, 4500 ft., some 40 miles N.W. of Lake Mangara, the district selected by Allen as the locality of typical suahelicu described by Neumann. This gives them a. particular interest. The skins, moreover, vary in tint in a way previously unrecorded and modifying in a measure Hollister’s observation and my own regarding the general uniformity in colour and pattern of the leopards of Tanganylka and Kenya. One of them exactly matches Grant’s series from Kigoma in brightness of hue. The other is a little duskier. It is, in fact, only slightly brighter than the Senegal example of P . p . leopardus described below (p. 574). A young male skin is nearly intermediate between the two others. The skulls are perfectly normal in shape and dimensions.

PANTHERA PARDUS ADERSI POC. (Pl. 11.) Panthera pardus adersi Pocock, Abstr. Proc. Zool. SOC. 1932, no. 347, p. 33. Locality of type.-Near Chuaka, Zanzibar ( W . M . Aders, B.M. no. 19.9.30.2). General colour yellowish tawny on the back, only a little paler than in the

mainland form ; but all the rosettes practically disintegrated into smallish rather closely packed spots, small rosettes being merely indicated by darker yc.llowish-tawny patches between some of the spots ; no solid rosettes on the thighs ; the paws scarcely washed with tawny-yellow, their spots reduced to closely packed speckling. Dressed skin : head and body 4 ft. 3 in. ; tail 2 ft. 3 in.

A second skin, also from Zanzibar ( J . H . J‘aughan, no. 29.4.l.l), is like the first in its pattern and dimensions, but is paler, less well coloured, and possibly faded. The tail in both is obviously shorter than in the mainland form. These two skins are distinguishable at a glance from other described races of leopards by being closely spotted all over, with the rosettes obscurely indicated. The skin that comes nearest to them and foreshadows the dis- integration of the rosettes is the one above referred to from Kenya Colony presented by Lady MacMillan.

~ O O . ZOOL. S00.-1932. 37


In English inches. _ _

In millim.


PANTHERA PABDUS PARDUS Linn. Felis pardus Linn. Syst. Nat. ed. 10, p. 41, 1758. Felis pardus pardus Cabrera, Bol. SOC. Esp. Hist. Nat. x. pp. 422427,

1910; id. op. cit. xviii. pp. 472482, 1918; id . op. cit. xxviii. p. 79, 1928 (Panthera) ; Thomas, Proc. Zool. SOC. 1911, p. 135 ; Hollister, Bull. US. Nat. Mus. xcix. p. 170, 1918.

Typical locality.-Egypt (Cabrera & Thomas) ; restricted to the Egyptian Sudan by Hollister.

There has been great difference of opinion regarding the allocation of the name pardus Linn., in a subspecific sense. Cabrera in 1910 decided that Egypt was the locality of typical pardus; and this view was confirmed, apparently independently, by Thomas, in 1911, who showed that the first author quoted by Linnaeus, with citation of a definite locality, was Alpinus, who saw specimens in captivity in Cairo and Alexandria. It is, of course, not certain whence these specimens came ; but more likely than not they were captured locally or at some point higher up the Nile. Adopting this latter supposition, and following the verdict of Cabrera and Thomas as to the restriction of the subspecific name pardus, Hollister in 1918 definitely assigned that name to some leopards from Khartoum and El Dueim in the Nile Valley. Allen, however, in 1924 set aside the opinions of Cabrera, Thomas, and Hollister on this point and, appealing to Buffon and Daubenton as the first revisers of Linnaeus, decided that Algeria was the original country of pardus, although there is nothing in Linnaeus’s citations to support that view. One of Allen’s reasons for rejecting “ Egypt ” as the locality of pardus was the statement made in 1740 by Mascrier that there are no leopards in that country. That is not true. Leopards, as I learn from Major S. S. Flower, late Director of the Zoological Gardens, Cairo, are still found in the western deserts of Egypt between Siva and Dabaa, Dabaa being on the coast only some 100 miles west of Alexandria. Hence the animals seen by Alpinus may have been locally captured, as stated above. Acting on this information received from Major Flower, I last year cited the deserts in question as the typical locality of P . pardus pardus; but at the time I had overlookcd Hollister’s previous decision that the specimens Alpinus saw were brought down from the Upper Nile, which is not unlikely. The priority of his selection of the locality renders mine null and void.

Since there are no skins or skulls of leopards frcim the Egyptian Sudan in the British Museum, I can add nothing to the little that is known of this race ; but, according to Hollister, who examined two skins, one from Khartoum, the other from El Dueim, preserved in the United States National Museum, i t differ3 from the Kenya Colony leopard he assigned to suahelica and from the leopard of Gondokoro and the Lado Enclave, cited below as chui, in being “ much more ochraceous buff -coloured.” Allen, unfortunately, did not refer to these two skins when he described the great individual variation in the colour and pattern of chui. It is quite possible that the two forms, provisionally kept distinct, will prove to be identical when more Sudanese examples comc to hand.

Since writing the last paragraph I have received the following notes on Sudanese leopards from Dr. J. Bryant, of the Sudan Medical Service, to whom I applied for information on the subject :-L‘ There are two very distinct types in this country. The leopard of the Dinder River, the Northern Sudan; and the Blue Nile is usually a small, very dark beast with spots set close together. I saw the bodies of four in one locality in Sennar lmt year [1930].

(Not Panthera pardus pardus Allen, 1924.)

37*

566 MR. R . I. POCOCK ON

Three of them were small and very dark, the fourth was a big boldly marked tawny beast. I had previously marked him down on account of his large pugs, which I at first mistook for a lioness’s. The leopards I have seen here [at Rumbek in the Bahr el Ghazal Province, over 500 miles S.W. of Sennar and 200 miles N.W. of Gondokoro] have all been boldly marked, large and tawny red or bright gold.”

The four specimens referred to above from Sennar are of particular interest. From their distribution they must belong to the same race as the skins f r an Khartoum and El Dueim which Hollister identified m P. p r d u s pardus. Yet of the four only one, the “boldly marked tawny beast,” approached the “ ochraceous-buff ” specimens he described. The facts above recorded show that typical pardus from Sennar may be dark and small spotted or paler and richer in tint and boldly spotted, exactly as in the leopards from the Upper Welle identified by Allen as chui and referred to below. On the other hand, the leopards from the Bahr el Ghazal Province, described by Dc. Bryant, apparently agree very closely with Hollister’s Sennar skins of pardus. But from the comparative nearness of Rumbek to Gondokoro, i t seems probable that they are the same as chui, the type of which was procured at Gondokoro. Dr. Bryant’s observations, for which I am very much indebted to him, bear out my suggestion that chui will prove to be indistinguishable from pardus as restricted by Hollister.

PANTHERA PARDUS CHUI Heller. Felis p r d u s chui Heller, Smiths. Misc. Coll. lxi. no. 19, p. 6,1913 ; Hollister,

Bull. U.S. Nat. Mus. xcix. pt. 1, pp. 170-173, 1918 ; De Beaux, AttiSoc. Ital. Sci. Nat. xlii. p. 276, 1923.

Panthera pardus chui Allen, Bull. Amer. Mus. Nat. Hist. lvii. pp. 254-256, pls. 45-54, 1924.

Locality of type and co-type.-Gondokoro and the Lado Enclave. Distribution.-From the typical locality westwards into the savannah-

district of the Belgian Congo to the north of the Ituri rain-forest area. There are no leopards in the British Museum from the geographical area

of this race. It was based upon the skin and skull of an adult from Gondokoro and an adult skull from the Lado Enclave by IIeller, who distinguished i t from the Kenya Colony leopards, identified as smhelica and ruwenzorii, by colour and pattern alone ; the spots were said to be fewer, large, and widely separated, and the ground-colour dominant with the hind legs and the underside of the end of the tail whiter.

Hollister subsequently assigned to i t an adult female from the Nzoia River on the Guasin Gishu plateau, thus extending its range into Kenya ; and Allen in 1924 referred to i t a fine series of specimens collected fmm 100-150 miles to the west of Gondokoro in the veldt country of the north-eastern Belgian Congo drained by the Welle River and its tributaries. In the same year De Beaux published some measurements of some skulls he identified as chui frcm Bussu.

Allen’s series shows the colour to be much more variable than Heller and Hollister supposed. The ground-colour is usually cinnamon-buff, more rarely pale bufly white, the belly being clear white or faintly tinted with yellowish. The pattern similarly varies, the extremes, as illustrated by his plates, being found in two examples from Faradje. I n one of these (pl. 51) the rosettes are large, comparatively few in number, and “ jaguarine,” many of them enclssing one or two small black spots on the dark central ground-colour. Even

(Text-fig. 5.)


_---_____I__

Gondokoro, d (type) . . . . . . . . . . . . . . . . . .

Faradje, d ad. (largest) . . . . . . . . . . . . . . . .

,, d ad. (smallest) . . . . . . . . . . . . . . . . .

,, d ad (average of 7 8) ......... Nzoia R. (Guaain Gishu), 0 aubd. Faracije, 9 ad. .............................. Garamba, Q ad. ..........................

the spots on the nape are arranged in very definite rosettes. In this skin, too, the interspaces are narrow, forming an irregular meshwork. In the other specimen the rosettes appear to be about half the size, much more numerous without forming a reticulated pattern, those on the nape not being arranged in rosettes. Allen does not endorae Heller's opinion that this race is distinguishable from the Kenya Colony leopard by colour or pattern ; but keeps it distinct on account of an average superiority in the size of the skull.

In the following table are reproduced the flesh measurements of specimens recorded by Hollister and Allen.

-

Locality and mx.

-I--.%- --- Gondokoro, d ad. 1 9 7+

Lado, d ad. . . . . . . . I 9.9 Faradje, d ad. . . . . . 11.3

(type).

Bussu, d ad. ....... . ' 100 + ,, d ad. . . . . . . 9.6 ,, d ad. . . . . . . 9.2

Upper Welle, d ad. 11.0 Nzois R., 9 subad. 8.5+ Bussu, 2 ad. ........ 1 8.6

,, Q ad. . . . . . . . . 8.4

Faradje, 2 ad. . . . . . . 8.1 ,. Q ad. . . . . . . . . 7.9 +

Garamha, ad. . . . . 7.8

Head and body.

9 + 8'9 + 9.8 8'5 8.5 9.7 7.8- 7.9+ 7%+ 7'2 + 74 7.0 +

3.9 ~

4.0 -

--- ft. in. 4 1) 4 8

3 93

4 06 3 8? 3 7)

4 2

- 1.7- 1.6 1-6

Tail.

f t . in. ---

3 9) .-

-

2 9% 2 9% 2 76 2 6)

2.3+ 23+ 2% 2.4 2.2 2.1- 2.0 2.O+

2-0 2.0- 1%+

2.0-

Total.

f t . in. 6 116

--

-

. -

6 11) 6 10)

6 3% 6 2

26+1 I7+ 25 17 26 20+ 24 1 17- 24+ 18 28 1 18 23 + , 14- 25 1 13-

23+' 13 24 I 13+ 23 ~ 13-

24 ! 1 2 t

The table shows that the average size of the males is slightly less than that of the three males from Kenya recorded by Hollister, and very much less than the average I have computed by including the Kenya males in Ward's

Also that the differences in average size between the males and females, amounting to only about six inches in total length, is considerably less than in the case of the Kenya form. Finally, whererts the males of chui do not surpass those of Kenya, the females are noticeably larger.

The following table of skull measurements is based upon data supplied by Hollister, Allen, and De Beaux, with the addition of a single specimen in the British Museum from the Upper Welle (Sir Alfred Sharpe).

Records.'

4.3 1% 33+1 ~-

Localitv and sex. I

1*8+ 1'4

In English inches.

Zygom. width.

6.0

6.1 + 5'9 - 6 7 6.0 + 5'8 6 4 5-2 -

5.3 -

5.0 +

--

-

- 5.0 -

I I

In millim. I Max. Upper Upper Lower

width. carnal. canine. carnal.

2*3+ 24 l6+ 19- ----__.--

t 204 19

18 19 20 18- 17 17 17 17 .t

I 17 ,

568 MR. R. I. POCOCR ON

When this table is compared with that of the Kenya and Tanganyika leopards, it will be seen, in the case of the males, that the variation in total length, amounting to two inches in skulls from Faradje, is very much greater and that the average length is about one inch longer. The average length of the female skulls is half an inch longer. I suspect that the skull from the Nzoia River, on the Guasin Gishu plateau, referred by Hollister to chui, should be regarded as an unusually large skull of the typical Kenya leopard ; geographically this leopard should be identical with the series from Mount Elgon ; but the female skulls from that mountain, like those of the males, fall into the same dimensional category as typical Kenya skulls.

So far as can be judged from the description of the colour and pattern, this leopard does not differ very appreciably, if a t all, from the leopard of Tanganyika and Kenya in those respects, except that the range of variation is greater. The range does not, however, seem to be greater than in the skins from Mount Elgon, which in that particular are transitional between the two types. Allen who believed, I think quite wrongly, that all leopards vary in colour and pattern as much as his series of chui from the north-eastern Belgian Congo, took no account of those characters in his diagnosis of the race but trusted solely to the greater average length of the skulls of the males. As has been shown, the average in the case of the females is also greater. The females also appear to be larger in bodily dimensions. The large size of the skull of chui is of special interest in association with the extension of this leopard to the west of the Nile into the watershed of the Welle (Uele) River, where no doubt it meets and blends with the typical West African leopard described below as leopardus. A comparison of the tables of measurements of the skulls of these two types shows close resemblance between those of the adult males. I can indeed, as stated below (p. 578), find no differences of moment between the skul1,in the British Museum, from the Upper Welle (text-fig. 5 ) , which I assign, on the information supplied by Allen, to chui, and the large skull of leopardus from Cette Cama in the Gabun. Such differences as exist in width are probably due to the Upper Welle skull being younger.

(Pl. 111.) PANTIIERA PARDUS RUWENZORII Camerano. Felis pardus ruwenzorii Camerano, Bol. Mus. Zool. Anat. Torino, xxi.

no. 545, p. 1, 1906 ; i d . in Abruzzi’s I1 Ruwenzori ; Zool. i. p. 89, pl. 1, 1909 ; Allen, Bull. Amer. Mus. Nat. Hist. xlvii. p. 257, 1924 (Panthem).

Locality of type.-Bujungolo, Ruwenzori. This race, based upon a skin and skull from Mt. Ruwenzori, was diagnosed

by Camerano as distinguishable from other African leopards by three main characters : its general dark brownish olivaceous ground-colour, the relatively large size of the rosettes on the back, flanks, and more particularly on the thighs, some of them enclosing small black spots ; and by the shortness of the tail, which is bushy at the end, and is less t h a half the length of the head and body, the latter measuring 5 ft. and the tail 2 ft. 3 in.

This description was taken apparently from the mounted specimen which Camerano figured. Judging from this figure I should say the tail is of normal length, its alleged relative shortness being due to the very obvious stretching of the head and neck in the stuffed specimen. The photograph suggests that the coat is full and shows that the rosettes are, as usual when large, separated by a network of the ground-colour

The skull, of which a figure was given and of which the recorded memure- ments are reproduced in the table (p. 571), shows no distinctive features, as Camerano stated.


The Museum is indebted to the Hon. M. Hachisuka for the donation of a flat skin of a leopard which in its olivaceous brown colour agrees closely with the description of ruwenzorii. It came from the Birungu Range, about 6OOO ft. alt., to the northeast of Lake Kivu.

Text-figure 5.

Dorsal view of skull of young adult male Leopard, identified 88 Paiathera pardun chui, from the Upper Welle.

The coat is longish and thick, carrying abundance of underwool.

x t.

The hair on the back is about 18 mm., on the belly from 22 to 28 mm., on the base of the tail 25 mm., and a t its end up to 32 mm. The colour is very dark, darker than that of any leopard I have seen, apart from " black " leopards,


the interspaces and centres of the rosettes showing hardly a trace of brightness of hue under ordinary light, although under reflected light, which brings out the orange-buff annulation of the tips of the hairs, the pelage assumes a decidedly brownish hue. The dark tint of the flanks passes a t a low level into the dirty white of the belly, which is the same colour as the throat and the insides of the limbs. The limbs externally are like the flanks with the paws greyish olive. The tail is mostly black above, irregularly streaked and patched with brownish buff in its basal half, with grey distally, the lower side being white, especially at the end. The rosettes are of average size, not so large as in the type of ruwenzorii.

The skin is unsexed, but from its dimensions I take it to be that of an approximately adult female or of a young male. The tail is of normal length, about two-thirds the length of the head and body.

In 1907 (Proc. Zool. SOC. p. 783, text-fig. 205), Lydekker published a figure and description of a skin from Uganda, in the British Museum (Stanley Tomkins, no. 7.11.20.1), which he supposed to represent suahlica Neumann. The skin does not agree with any recorded examples of that race and it is not " rufous fawn " in tint, as Lydekker said. There is scarcely any brightness about the ground-colour, which is tawny olive on the back and fades quickly to creamy or buffish grey on the flanks, and to clean white below, the legs and paws being mostly greyish white, with scarcely a trace of tawny wash. The rosette's are very sharply defined against the pallid ground-colour, and are quadrangular or polygonal in outline, with tolerably straight unbroken edges, and narrowly separated by the reticulated pattern of the ground-colour ; some of them are " jaguarine," as Lydekker stated, enclosing, that is to say, one or two small black spots, the rosettes on the thighs being what are called " solid," i. e., having the centres as black as the rims.

The dried undressed skin measures head and body 4 ft. 6 in., tail 2 ft. 1 in., total 6 ft. 7 in. The tail, therefore, which seems to be complete, is relatively short, less than half the head and body, as Lydekker detected. But when pegged out for drying, the skin of the tail was manifestly stretched sideways and widened, and this has had the effect of shortening it to a certaiii extent, probably by a few inches, making it, on this supposition, about half as long as the head and body, which is short as compared with normal leopards.

I am unable to place this skin, for which, unfortunately, no precise locality is known. The coat is much shorter, the ground-tint much paler, and the rosettes much smaller than in typical ruwenzorii. It probably came from some low-lying part, of Uganda ; and its identification must be left for the present in abeyance.

Another leopard recorded from this region was named Felis pardus centralis by Unnberg (Sv. Vet. Akad. Handl. Bch. Iviii. no. 2, pp. 49-55, 1917). It was represented by the skin and skull of a subadult male from Kabare to the east of Lake Albert Edward. It has large " jaguarine" rosettes, larger perhaps than in the type of ruwenmii, the ground-colour forming a reticulated pattern. " very pale," the '' yellowish buff '' back lightening to " maize yellow " on the flanks and thence to white on the belly. The stripped skin measures: head and body 5f t . , tail about 2 it. 94 in., the tail, that is to say, although described by Gnnberg as short, is well over half the length of the head and body.

Judging from the description of the colour this leopard closely resembles rather pale examples of the ordinary East African form. The rosettes are certainly much larger than in any Kenya or Tanganyike skins I have seen ; but Heller had a :kin from Mt. Meru, close to Kilamanjaro, which, on account

It is unlike that of any leopard I have seen.

But this colour was described

THE LEOPABDS OF AFRICA. 57 1

tumnzorii.

Kabare, d ad.. . . . . . , 9 3 +

Bunyoro Bugoma,

Ruwenrori, d ad. .# 9'0 ----- (central is).

--.-__-__

d Yg. 1 8.0

Kigezi, d old . _ . . . . !:- ituren8is.

- _ - ~ -----

I

Niapu, d ad. . . . . . .' 10.2 Gamangui, d a d . .I 100 Medje, ,-J ad. . _ . _ . . 8.8 Gamsngni, d ' 9.0

Poko, d ad .._. ... . _ . 9*8+

aubod. Poko, 0 ad . __ . . . . ... 7.7 Akenge, $? ad. . . . ' 7% +

7'4+ Niapu, Q ad. ......' 7.4 + Akenge, Q ad. ... I 7.0

N i a p ~ , 9 4. ...... I I

of the large size of the rosettes, he assigned to ruwenzmii, a skin which Hollister identified as an aberrant example of suahelicu.

Clearly the type of centralis cannot be classified ; but since that name is inadmissible owing to preoccupation, as Cabrera pointed out, further discussion of its affinities would be profitless. I cannot, however, agree with Cabrera that i t may be identified with iturensis of Allen. It is much too light in tint, and no example of iturensis has as yet been seen with so bold a pattern.

The skull of which the dimensions recorded by Lijnnberg are given below, shows no distinctive features. Of the two skulls in the British Museum to the east of the lakes, one came from Bugoma, 3750ft. alt. (J. Jardine, B.M. no. 31.4.1.10). It is that of an immature male with teeth exactly the same size as those of the type of ruwenzurii; without the skin it is impossible to identify it. The other was obtained from a native by Capt. Phillips in Kigezi, S.W. Uganda (B.M. no. 28.7.6.1). It was sent with the skin of a spotted Hyzna, the two representing the " Nandi Bear," and was referred to and figured in my article on that mythical beast (Nat. Hist. Mag. ii. pp. 162-169, upper fig.). It is that of an old male with worn teeth and the whole of the back shot away. It is a very large skull, probably as long, when entire, as the big skull of chud from Paradje recorded by Allen. The mandible at ali events is a little longer than that of the male from the Upper Welle, entered in the table of measurements of chui as 11 in. in total length. The teeth also are large. It is the biggest leopard skull recorded from the east of the Rift Valley, but is not identifiable without the skin.

The available dimensions of the four skulls from the east of the Rift Valley Lakes are entered in the following table together with those of the next race.

8.4 - 1 5.7- 5'6

----

-- 7.3 50

-: - 6.6 I----

9.3 6.3 + 9 + 6 0 - 6 4 + 76+ - 8 + 5'5+

6.8 4.8- 6.8 + 5.0

Tn English inches. 1 In millirn.

1%

1.7 __

--

1.7-

1.6t I.?+

1'6+ 1.5-

1.6-- 1*6+ 1%+ 1.7- 1.6.-

1.2 2.0 25

1.8 26 27 __--- -

1.7+ 2'6- 26+

1.6+ %4+ 26+ 1.7 24- 27+

1*5+ 2.2- 24+ 1*4+ 2*2+ 26+

1*2+ 1.8 22+ 1*2+ 1*8+ 24+ 1,1+ 1*7+ 23- 1.3- 1*8+ 22+ 1.2- 1*7+ 23+

__--

Zygom. width.

-_

Maa- toid width -- -

- --

3.2

- --

4 + 3 8 + 4 -

3.7 + 3.5 - 3.5 + 3.2 31 +

-

--

Waist tt. ax. Upper '

width. width. width. carnal

25 - _ -

UPpe enme

16

21 --

- 21- 19 19 16 + 16 + 12 + 12 - 12 + 14 - 13 +

18

21 --

20 - 20 t 19 t 18 t 18 -

15 16 + 16 + 16 17 -

572 &R. R. I. POCOOg OR

PANTHERA PARDUS ITURENSIS Allen. Panthera pardus iturensis Allen, Bull. Amer. Mus. Nat. Hist. xlvii. p. 259,

Distribution.-The Ituri Rain-Forest District of the Belgian Congo. Allen, who examined 20 specimens, skins and skulls, from Poko, Akenge,

Niapu, Medge, and Gamangui in the above-mentioned area, described this race as similar to P. pardus chui, but smaller and darker in general coloration, with the pelage shorter and thinner. In cranial measurements, he added, it is intermediate between P. p . suahelica, the smallest, and P . p . chui, the largest, of the East African leopards with which he was acquainted, the average total length of five adult male skulls of swhelica being about gin., of iturensia about 9.6in., and of seven adult male skulls of chui about 10.4in. Allen also published a valuable series of photographs of skins exhibiting considerable variation in the size, shape, and spacing of the rosettes ; and he repeated that “ the coloration averages much darker and the pelage is distinctly shorter and thinner than in chui or suuhelicu, in conformity with the different environ- ments of the three forms.”

It was a misfortune that Allen had no examples Qf the common West African leopard (P. pardus leopardus) wherewith to compare iturensis. He dismissed leopardus from consideration on the grounds that its typical locality is Senegal, which is not only far remote from the Ituri Forest Region but in a very different environment. The Senegal leopard, nevertheless, judging from a skin in the British Museum is, as explained below and as stated by Buffon, like the leopard of Guinea and other coastal forested areas of tropical West Africa.

Through the kindness of the Directors of the American Museum of Natural History, and Mr. H. E. Anthony, the Curator of Mammals, I have been able to examine three of the skins and two of the skulls of P. pardus iturensis. The skins are of particular interest in exhibiting greater individual variation in general colour than I have observed in any series of leopard skins from one district. They probably cover the range in tint and pattern referred to by Allen in his description of this race.

pls. 41 & 55-64.

They may be briefly described as follows :- (1) Adult 6 from Medge, June 16-24, 1914 (no. 52032). General colour

dusky, olivaceous grey, with scarcely a trace of the bright ochreous or fulvous tint seen in P. pardus swhelica. The centres of the normally spaced rosettes are darker than the interspaces, the tint of which gradually blends with the dullish white hue of the underside and of the inside of the limbs. The coat is short, smooth, and sleek.

This skin, which was figured by Allen on pl. 56, in its pattern and general coloration is scarcely distinguishable from the skin from Senegal (Winwood Reade, 13.5.9.2), below referred to, but is shorter coated and sleeker. It is also unmistakably similar to two young female skins from Bessiadzi and Sekondi on the Gold Coast. Judging from this skin from Medge there is not a particle of evidence that P. pardus iturensis differs from the common West African leopard P. pardus leopardus.

A very dark skin, darker than the ope from Medge, partly because the closer packing of the rosettes narrows the interspaces and partly because the centres of the rosettes and the interspaces are washed with a deeper brownish orange tint. The coat is a little longer. The skin is emphatically darker than skins of typical East African leopards in its darker orange-brown wash due tQ the tips of the brighter oohraceous hairs, both within the rosettas and on the

(2) Adult 0 from Niapu, Dec. 15th, 1913 (no. 52026).


interspaces, being more conspicuously fuscous apically. Allen’s figure of this specimen on pl. 57 shows the larger, darker-centred rosettes reducing the interspaces to a narrow reticulated pattern, the particulars in which, apart from the deeper browner colour, this skin differs from the male from Medge. The plate also very clearly shows the longer coat of the Niapu example. This feature was not referred to by Allen, who wrote of the specimens of iturensis as if they were all alike in having the coat quite short and smooth. Since the female from Niapu was shot in the middle of December 1913, and the male from Medge in the middle of June 1914, the difference in the length of coat is probably seasonal. The point of interest, however, is that the female from Niapu does not differ appreciably in length of coat from leopards from Tanganyika in the British Museum.

On the evidence apparently of the skull, Allen altered the symbol to “ 0.” The skull, which was sent to me, certainly has all the characters of a female skull. The collector, nevertheless, may have made no mistake, since there is evidence from several sources that male leopards now and again fail to develop the cranial characters typical of that sex.

This skin, figured by Allen on pl. 59, differs markedly in colour, not only from the female from Niapu, but from the male from Medge. It is a richly coloured leopard quite normally spotted and would pass anywhere for suahelica. The only difference I can detect between i t and examples in the British Museum from Tanganyika Territory and Mt . Elgon is that it is the merest shade darker on the back, owing to the tips of the ochreous hairs being slightly more infuscate. But it unmistakably more closely resembles skins of suahe2ia.z than it resembles either of the other two skins from the Ituri district associated with it.

One was procured by Dr. Cuthert Christie, who tells me he shot the animal either a t Poko or Medge to the south of Niangara in the marginal forest area. It is of considerable interest as being in several respects intermediate between the two above-described specimens from Niapu, the typical locality of iturensis. The skin is thick-coated and decidedly less brightly coloured, less tawny yellow than the female from Niapu (no. 52031)) and is altogether darker than the skins I assign to suahelica. But it is not nearly so dark as the female from Niapu (no. 52026), the wider, more brightly coloured interspaces making the whole skin paler. The dried undressed and unsexed skin measures : head and body 4 ft. 2 in., tail 2 ft. 94 in. The second is a mounted specimen from the Ituri forest (R. Ward, B.M. no. 22.12.19.2). It is not so bright as the paler of the two skins from Niapu, nor quite so dusky as the example from Medje. It is gradational between typical iturensis and the West African leopard I identify as leopardus.

The measurements of the skulls of iturensis in the table (p. 571) show considerable variation in the size of male skulls and much less variety in that of feFale skulls. The table also indicates that the average size of skulls of this race is less than that of chui in males, very much less in females, that it is about the same in the two sexes as in the Kenya-Tanganyika leopard, not greater, as Allen claimed ; and that it is noticeably less in males than in the West African leopardus, and approximately the same in females.

THE LEOPARDS OF TROPICAL WEST AFRICA. The material of leopards from this area of Africa, in the British Museum,

is comparatively scanty, consisting, with one exception, of a few skins without skulls, and a few skulls without ~kins.

This skin, it may be added, was labelled I ‘ 6 ” by the collector.

(3) Adult 6 from Niapu, Feb. l l t h , 1914 (no. 52031).

Two skins in the British Museum are assignable to this race.

574 a. R. I. POOOOK ON

The skins by their colour and localities are very definitely assignable to two types, a darker forest form, for which I adopt, as in 1907 and 1909, the name leopardus, and a paler savannah form, which I believe to represent Cabrera’s reichenozoi.

The skulls have a special interest and are dealt with under a separate heading.

PANTHERA PARDUS LEOPARDUS Schreb. Le LBopard, Buffont Daubenton, Hist. Nat. ix. pp. 153, 169-170, pl. xiv.

1761. Felis leopardus Schreber, Sllug. iii. pp. 387 & 586, pl. 101, 1777 (based on

Le Gopard of Buffon, whose plate is reproduced) ; Erxleben, Syst. Reg. Anim. p. 509, 1777 ; Cuvier & Geoffroy St.-Hilaire, Hist. Nat. Mamm. i. pt. 20, 1820.

Felis pardus leopardus Pocock, Proc. Zool. SOC. 1907, pp. 675 & 1038 ; id. op. cit. 1909, pp. 204, 209, figs. 22-23 ; Cabrera, Bol. SOC. Esp. Hist. Nat. xvii. p. 479, PIS. 16& 17, fig. 2, 1918.

(Text-fig. 6.)

Felis pcmilura Valenciennes, C.R. Ac. Sci. Paris, xlii. p. 1036, 1856. Pantherapardus iturensis Cabrera, Bol. SOC. Esp. Hist. Nat. xxviii. pp. 93-94,

Locality of type.Senega1. Distribution.-Forested district of West Africa from Senegal to Spanish

Guinea. Buffon and Daubenton described the colour of this leopard of Senegal

and Guinea, of which they had seen many skins, as more or less darkish tawny on the back and flanks and whitish below, the spots all much of a size, being considerably smaller than those of the panther. The skins measure: head and body about 4 ft. ; tail 2 ft. to 23 ft. Schreber, who gave the name Felis leopardus to this leopard, described it as “ brownish yellow,” and Erxleben’s diagnosis runs “ corpore fusco, macuh subcoadunatis nigris . . . maxime approximatis.”

F. Cuvier and Geoffroy St.-Hilaire published a description and coloured plate of an example from Senegal, showing that the colour is greyish brown, much darker than the Algerian leopard which they also figured, and entirely lacking its bright yellowish hue. The rosettes, however, are tolerably large and spaced.

Writing of this race in 1907, I referred to specimens imported to the Zoological Gardens from Sierra Leone and Ashanti, and described them as very uniformly coloured and of a peculiar dusky shade produced by the olive- tawny tint of the interspaces and the approximation of the black spots. I have since seen several additional examples from the coastal districts of tropical West Africa, all exhibiting the same dusky hue quite distinct from the brighter yellowish tawny characteristic of most East African and Indian leopards.

Since Buffon referred more particularly to Senegal in his description of this leopard, Cabrera indicated that district aa the type-locality of the race.

In the British Mpseum there is a skin from Senegal obtained by Winwood b a d e (B.M. no. 63.5.9.2). It is that of a rather small unsexed specimen measuring head and body 3f t . 9 + h . , tail 2f t . 2in. ; total 5 f t . lliin., with smallish close-set spots, andof the decidedlydarkish-olivaceoua or brownish- tawny hue referred to above. There are also the skins of a young female from Bessiadzi on the Gold Coast and of a still younger male from Sekondi. Apart from minor variations these three skins are similar in colour and pattern. They completely bear out Buffon’s statement that the leopards of Senegal

1928 (not of Allen).


and Guinea are alike. C o n h a t i o n of this, so far as the darkish olivaceous- tawny huc is concerned, is supplied by an adult male from the Gold Coast, now living in the Zoological Gardens and presented to the Society by Capt. Muciie in 1922. This animal, however, has considerably larger rosettes than those of the skins mentioned above, resembling in this respect the Senegal specimen figured by Cuvier.

Text-figure 6.

Dorsal view of the skull of an old male Leopard (Panthern pardus leopnrdzra) from French Guinea. x +.

Evidmce that the leopards of these districts of West Africa may attain a large size is supplied by two entries in Rowland Ward’s ‘ Recordu,’ 1928, p. 484. One from the Gold Coast was 8 ft. 2 in. from nose to tail-tip ; another, from Sierra Leone, was 7 ft. 11 in. Both were measured before skinning. The very large skull of the Sierra Leone specimen is referred to below in confirmation of the opinion I published in 1907 and 1909, that leopards similar to those of Senegal and Guinea are characteristic of the forested districts of West Africa, at least as far south aa the Gaboon.

57 6 M R . R. I. POCOCK ON

There is also in the British Museum the skin of an immature male from Luluabourg in the Kasai District of the southern Belgian Congo, to the south of the river, collected by Phre R. Callewaert (B.M. no. 27.12.21-21). The rosettes are small, close-set, and more or less broken up. In a dull light the tint is darkish and very similar to the skin from Senegal ; but in a good light it is seen to be ycllower and altogether brighter. But i t is not nearly so bright as typical Kenya and Tanganyika skins and, on the whole, perhaps comes nearest to the yellower skins of the Ituri leopard (iturensis).

In 1909 I described some skulls from Cette Cama in the Gaboon and assigned them to leopardus. This identification was adopted in 1918 by Cabrcra, who had similar skulls from Spanish Guinea ; but in 1928, as a result apparently of Allen’s comments in 1924 on the Senegal leopard, which was unknown both to him and to Cabrera, the latter dropped the name leopardus for the Spanish Guinea leopard and identified it as iturensis of Allen, a view which is not, in my opinion, correct.

Cabrera discussed at some length the probability, as he considered it, of the West African forest leopard extending from the west coast across Central Africa to the Ituri, whence Allen’s specimens came, and thence across the Divide to Kabare in Uganda, whence came the leopard described in 1917 by Lannberg as Felis p . centralis. But even if Cabrera’s opinion on this point be correct, there is an older title than iturensis for the leopard of Spanish Guinea known to Cabrera, namely pa?cilura, given by Valenciennes in 1866 to a specimen from the French Congo, a district geographically so near and in physical features so similar to Spanish Guinea as to preclude the idea that the leopards are subspecifically distinguishable.

PANTHERA PARDUS REICHENOWI Cabrera. Panthera pardus reichenozoi Cabrera, Bol. SOC. Esp. Hist. Nat. Madrid,

xviii. p. 481, 1918 ; Allen, Bull. Amer. Mus. Nat. Hist. xlvii. pp. 260-263, 1924.

Locality of type.-Yoko in the savannah district of the Cameroons to the north of the River Sannaga.

Cabrera described this race as a small form, with a long tail ; the ground- colour is “ very vivid ’’ above and pure white below, the general tint being somewhat stronger [? brighter or darker] than the ochraceous buff of Ridge- way. The rosettes are very compact [? close-set], irregular, tolerably largc and centrally darker than the ground-colour.

This leopard, which inhabits the herbaceous meadow-land of Yoko, was separated by Cabrora from a race, referred to leopardus, found on the borders of the forest in the northern part of Akonolinga to thc south of the River Sannaga. This marginal forest form has a proportionately shorter tail, the spots more widely separated and the inferior parts of the extremity of the tail only a little paler than the rest of the pelage and inclining to golden or yellowish white. Judging from the description this form does not agree with the typical West African forest leopard which I identify as leopardus, a dusky olivaceous buff leopard with the uderparts and the lower side of the tail clean white. It3 identification may be left in abeyance ; but there is no doubt that the open country Yoko form, reichenowi, differs from the dusky forest leopard in the brightness of his hue, which approaches ochraceous buff, thus resembling the typical East African leopard commonly known as swthelica.

In the British Museum there are two skins of West African leopards which may be provisionally, a t all events, assigned to re i chemi . They differ markedly from the ordinary dusky West African type, leopardus, in their paler, brighter,


yellower tint, although neither is so richly tinted as average specimens of suahelica, nor apparently as the type of reichenowi.

One of the skins, presented by Rowland Ward (B.M. no. 7.12.12.2)) came from the Rornu country to the south of Lake Chad, where the environment is no doubt very similar to that of Yoko, though possibly less fertile and more desert. The rosettes are smallish, mostly annuliform and well spaced, with the centres only appreciably darker low down on the pale flanks. The hind legs are nearly white, the fore legs faintly washed with buff, and there is very little black a t the end of the tail.

The second skin has no definite locality, but it was obtained by Alexander Gosling (B.M. no. 7.7.8.60), who collected in the Lake Chad district. It is very similar to the Bornu skin in colour, but is a little paler, and has the rosettes more irregular and less annular in shape.

The general colour is a rather pale sandy fawn.

THE SKULLS OF WEST AFRICAN LEOPARDS. Of the skulls enumerated in the subjoined table, the first three are those

described in my paper of 1909 quoted above. They are no longer available for examination and remeasurement, the dimensions given being taken from the previous record. They were large skulls, the first and largest on the list, of which I figured the ventral side, being until 1024, the largest recorded leopard skull. They were muscularly well developed and had narrow mesopterygoid fossae, that of the largest being exceptionally narrow. A considerably smaller, but otherwise apparently very similar, 6. skull is the one from Rio Muni (Spanish Guinea) figured by Cabrera ; and there is no doubt that Cabrera was perfectly correct in assigning tkis skull and two others from Spanish Guinea, like it, but a little larger, to the same race as the Cette Cama series. Evidence that this potentially large-skulled race extends at least as far north as Sierra Leone is furnished by the skull above referred to from that locality, which, according to Ward’s entry, was l l $ in . in total length, slightly longer, that is to say, than the largest of the Cette Cama skulls I measured.

For this race Cabrera originally adopted, and I think rightly, the name leopardus which I applied to it. Subsequently, as explained, he substituted iturensis for leopardus. But the average length of the five west coast skulls, above enumerated, is only a fraction under 104 inches, that is to say, it exceeds by almost 1 inch the average length of five adult male skulls of iturensis given by Allen, and is the same as the average of seven adult male skulls of chui from the Welle district of the north-eastern Belgian Congo, also measured by Allen. These data point to the conclusion that the west coast leopard is larger than the Ituri leopard, and shares with the Algerian and Welle region leopards the distinction of being the largest existing races of this animal, a conclusion borne out by the only skull of the Welle race known to me and referred to above as practically equalling in size and resembling in structural characters the largest of the three Cette Cama skulls I described.

The average of the five Gabun and Spanish Guinea skulls is scarcely affected by the inclusion of the sixth on the list, brought by G. L. Bates from the French Congo (B.M. no. 5.11.27.14), which is a little shorter than the Rio Muni skull. It is unsexed, but is unquestionably that of a male ; and is interesting from two points of view. In the first place, its district is the same as that of the type of pcecilura described by Valenciennes. In the second place, although old with scarcely a trace of sutures remaining, it is remarkable for its small sagittal and occipital crests. These two characters give its cranial portion a resemblance to that of a female leopard, although its general dimensions

578 MR. R. 1. POCOCK ON

and the size of its teeth, particularly of the canines, mark i t as a male (text- fig. 6, p. 575). It is, in fact, a male skull which has failed to acquire on the cranium the indications of excessive development of the temporal muscles characteristic of all typical skulls of adult and old male leopards. To illustrate this point I have figured the dorsal aspect of this skull for comparison, with that of the adult male skull from Upper Welle (text-fig. 5, p. 569), which in size, proportions, and other features is to all intents and purposes like the large skull from Cette Cama assigned, with the Belgian Congo skull *, to leopardus. It may be noted, too, that the French Congo skull, showing no trace of sutures on its cranial portion, is a good deal older than the Welle River skull, which has the fronto-parietal, interparietal, and squamoso-parietal sutures unclosed, and had probably not reached its full length, although its sagittal and cjocipital crests are exceedingly well developed.

In general aspect and size the Belgian Congo skull is intermediate between the five skulls preceding i t and the four following i t on the list, which have the characters of female skulls.

The one from Kasai belongs to the skin from that district above referred to. It is that of an immature male and was correctly sexed by its collector. The teeth, it may be noticed, are considerably smaller than those of the French Congo skull, and only a little longer than those of the skulls marked as females ; my reference of this skull to leopardus is provisional.

The three skulls from Yoko, Mamfe, and the Yobe River were sexed by their collectors as males, and possibly correctly, but I have entered them interrogatively as females on the evidence of their size, shape, and dental dimensions t, the smallness of the canine tooth being particularly noteworthy. The skull from the Yobe River, on the northern border of Nigeria, was collected a t Gorgoram (F. W. A. Migeod, B.M. no. 23.12.10.1). It is strongly built and fully adult, with all the cranial sutures closed. From its locality I provisionally assigned it to reichenowi. It is, however, practically indistinguishable from the skull from Oban, to the north of Old Calabar in Southern Nigeria (P. A. Talbot, B.M. no. 12.10.28.71), which, although unsexed by its collector, I regard as a female and, from its locality, as probably ascribable to leopardus.

Theskull from Mamf6 in the Cameroons(Dr. Dyce Sharp, B.M. no. 27.12.16.1) is hardly full sized, the upper cranial sutures, including the median frontal, being still open, although the basioccipital is closed. I have been unable to ascertian whether Mamf6 is in the forested or the savannah district of the Cameroons. The determination of the leopard must therefore be left in abeyance.

The skull from Yoko, to the north of the Sanaga River in the Cameroons, belongs to the type of reichenowi described by Cabrera as a male on the testimony of the collector, Dr. Reichenow. Trusting this determination, Cabrera came to the conclusion that three West African skulls similar to it in size and shape were also those of males. One of these was the small skull from Cette Cama, which in 1909 I identified as hpardus and probably aa a female. The others were two skulls from Spanish Guinea coming from the same locality as thk large skulls he had recorded from that Colony. From these assumptions he inferred the existence of two races of leopards in West Africa, living side by side in the same country, one (leopardus) with a large

Despite the structural differences between thiR skull from the French Congo and those from Spanish Guinea, evidence of their subspecific identity ia supplied by the fact that it was procured by Bates when his headquarters were on the Benito River, which passes through Spanish Guinea.

t In male leopard skulls with female characters which I have seen from other localities, the size of the upper canine tooth is unaffected by the general suppression of male characters,

\

Som

e m

easu

rem

ents

of S

kulls

of W

est

Afr

ican

Leo

pard

s (P

. p. l

eopa

rdus

and

rei

chew

wi)

.

Max

. w

idth

.

25

2.7

+ 2.

5 +

-

-

2.4

2.1

1.9 -

1.9 -

2.0

1.9 -

~

._

Loc

ality

and

sex

. U

pper

ca

rnal

.

26 +

26 +

25

27 +

27

27

25

22 -

23

24

24

23

24

----

~~~~

~ ~

I- i Cte

C

ama,

Gab

wn,

d d. ..

...

,, I,

..

dd

..

....

,, 1,

..

d ad. .

.. S

pd

ah

Gui

nea,

d a

d.

....

....

..

Rio

Mnn

i, d

ad.

....

....

....

....

...

Fren

ch C

ongo

, d

ad. .

....

....

...

Kae

ai, S

. Con

go, d

imm

. C

ette

Cnm

a, G

aboo

n, $

? ad

. , Sp

anish

Gui

nea,

9 a

d ....

......

...

....

...

....

Obe

n, S

. Nig

eria

, 9

ad.

.I 1 ..

....

....

Y

obe,

N.

Nig

eria

, ? 9

nd.

. Yok

o, C

amer

oons

, P $2

ad

....

....

Yam

fB, C

amer

oons

,? Q a

d.

....

. ..

....

..

wid

th.

1.6

+ 1.

9 -

1-6

1.7 -

1.7-

1.7

1.8

1.6

1.5

.t

1.6

+ 1.

7

1.5

1.7

-

I

wid

th.

1.7

+ 1.

9 -

1.7

+ 1.

7

1.7

+ 1 '5

1.4

1.1

+ 1.

3-

1.4

1.5

1.2

1 *3

Tot

al

leng

th.

11.1

10

7

10

5

10.1

96

93

8.

1

1.5

76

-

--

-

-

79

'

7.6

7.1

1 5

-

7.1

48

.In E

nglis

h in

ches

.

3.1

3.0

7'0

7.1

6.0

3.8 -

5.3

3.5

4.5

: -

48

I -

l-

In m

illim

.

Low

er

carn

al.

-

-

-

-

19

20

17

-

16

16

18

16

16

~

W

m

cn 4

P

580 M R . R. I. POCOCK ON

well-developed skull, the other ( r e i c h e m i ) with a small poorly developed skull. Criticising this conclusion in 1924, Allen maintained that the cranial characters Cabrera relied upon are sexual, and that the skull and skin constituting the type of r e i c h w i belonged to different animals, the skin being male and the skull female. I accept Allen's view that the Cette Cama and Spanish Guinea skulls are those of females ; but for the following reasons I am doubtful about his verdict on the skull of reichnowi :-

(1) There is evidence from other quarters, India, Abyssinia, and S . Africa, that occasionally the skulls of adult male leopards are arrested in development at the " female )' stage through which they all pass.

(2) The male skull from French Guinea, although oldish, has the sagittal and occipital crests small and like those of the female skuIl, thus differing from the skulls of typical male West African leopards.

(3) Three collectors independently sent to Europe from West Africa three skulls ticketed with the male sign which resemble female skulls in all their characters ; and the localities whence these skulls came are not far apart, two being in the Cameroon district, the other in Northern Nigeria.

(4) Although collectors admittedly make mistakes now and again in sexing specimens, the coincidence of three such mistakes being independently made in a comparatively small area would be curious. Cabrera's view, therefore, that reichemi is a race of leopards in which the skull of the male is arrested in development cannot, in my opinion, be summarily dismissed, as Allen dismissed it, until additional material proves it to be wrong.

THE LEOPARDS OF NORTHERN RHODESIA. From this district of Africa I have seen a good number of skins transitional

in colour and pattern between those occurring to the north and south of it. They are too variable to define and are left unnamed. The following are some notes upon them.

N'dola District.-The skin of a probably adult male, presented by Dr. Gordon Lancaster to the British Museum (no. 21.7.18.6). It is, unfortunately, a good deal rubbed and has lost half the tail, and in its present condition is like the skin of a Jaguar (Panthers onca) in colour and pattern, the rosettes being fewer and larger, with contained black spots, than those of any African leopard skin hitherto depicted (Pl. IV.).

A young 9, quite different from the last in its small close-set spots, and in colour, pattern, length, and texture of coat hardly distinguishable from the 9 skin from the Mohango Drift, Damaraland, recorded below under P . p . shtridgei. It is a little smaller, the flesh measurements being : head and body 2 ft. 6 in., tail 1 ft. 11 in., and as in that specimen the milk dentition is retained, although the permanent teeth are a little more advanced. The canine and upper carnassial, cut out of the bone, are a little smaller. It is of interest to record that this little leopard, despite its dependence on the milk-teeth, " had just killed a good-sized goat, when shot."

Musa River, Kafue District.-Dec. A male skin obtained by Dr. Gordon Lancaster and kindly lent to me by Mr. G. C. Shortridge. The pattern is normal and the colour rather pale and the coat is longish and rough, but not so rough as in the skin from Upington in the southern Kalahari referred to below.

Seshehe District.-Dec. The same history as the last specimen, from which it differs in having the coat rather softer and shorter, and the rosettes larger and better defined.

N'dola (G. C. Shortridge (no. 21.7.18.7)).-Skin and skull. April 28.

THE LEOPARDS O F AFRICA. 68 1

Zygom. w'dth*

5'3 58 4.6 4.4+

--

Mumbwa District, 3800 ft., on the right and left banks of the KafuB. An interesting series of some half a dozen skins and skulls procured from natives by Capt. C. R. S. Pitman. The skins vary a good deal in tint and length of coat. Most of them are tolerably typically coloured " savannah " leopards, rich buff on the back, paler buff on the flanks, with the rosettes of normal sue and spacing. Others are richer tinted and altogether darker animals, ochreous, even rusty ochreous on the back and in the centres of the rosettes, recalling iturensis. One of the paler types, a specimen about two-thirds grown, has a considerably longer coat than the adults. A still younger individual, about one-third grown and darker tinted, has the coat long and wavy. An example from Broken Hill, 4600 ft., in Rhodesia is not so brightly coloured as the others above referred to, owing to the interspaces and the centres of the rosettes being more speckled with fuscous. It is an immature specimen with a full rough coat, and in its duskier hue approaches P . p . leopardus.

Kalene Hill.-Skin and skull of a very nearly full-sued male presented by Dr. Walter Fisher (no. 19.1.27.1). The coat is full, thick, and longish without being rough ; but the colour and pattern are very abnormal and semi-albinistic, the ground-tint being pale cream buff with the normally sized and spaced rosettes varying from pale to chocolate brown.

The following are the skull measurements of some North Rhodesian specimens :-

~~

In English inches.

Mas-

width

3.3 3.8 3.0

toid

-

Max. width.

2.2 2'2 1.8 1.9 1.8 1.9

Upper carnal

26 25 24 24 22 24

,, Q ad. ... - I 4 6 3.0

,, p a d .... - 5 I -

. Waist width.

1.5 1.6 1.6 1.8- 1.5 -- 1.6

-_ Upper Lowei canine. carnal

15 20 -_-- 16 - 12 l5b 11 17 114 16 12 17

Int. orb.

width ---'----- Kalene Hill, d ad. . 8.8 Mnmbwa, 6 snbad.. 9.2

,, $? ad. ... 7-5 ,, $? yg.4. 7.3

1.4 1.5 1.3 1.2 1.1 1 -4

8.1 8.3 6.9 -

I I

The male skulls measured are insufficient to supply reliable data ; but those of the females agree very closely with the skulls of P . p . ituremis.

THE LEOPARDS OF SOUTH AND SOUTH-WEST AFRICA. As stated above, the typical East African " Yellow " leopard, P. p . fu,au

(=smhelica), extends as far south as Nyasaland. To the south of the Zambezi, a t all events westwards, a slightly paler form sets in and this reaches its palest phase towards the coastal districts of the south-western part of the continent. The only South African leopard bearing a distinctive name is a peculiar variety occurring near Grahamstown. This was described as Felis pardm var. m&n- d& by Giinther ; and Allen, in his comparatively recent revision of African leopards, appliedmelanoticacomprehensively to the leopard of the Cape Province, which he declared, " differs in size and character of pelage from the more tropical forms." This statement, although made without citation of authority, is borne out by my observations on the available material in the British Museum.

38*

582 M R . R. I. POCOCK ON

PANTHERA PARDUS MELANOTICA Gynther. Felis pardus Trimen, Proc. Zool. SOC. 1883, p. 535. Felis pardus rnelanotica Gunther, Proc. Zool. SOC. 1885, p. 243, pl. xvi. ;

id . op. cit. 1886, p. 203, fig. ; W. L. Sclater, The Fauna of South Africa, Mamm. i. p. 37, text-fig. 10, 1900 (cited as var. of F . pardus).

Felis pardus melanosticta (errore) Lydekker, The Game Animals of Africa, p. 430, 1908.

Typical and only recorded locality ; near Grahamstown. In the typical example of this leopard, preserved in the British Museum

(no. 85.2.28.1), the pelage is smooth and sleek, but thickened with underwool. The rosettes, characteristic of normal leopards, are everywhere disintegrated into a multitude of small spots or specks, mostly so close

The solid spots in most areas are also similarly broken up. Here and there the rich ground-tint is distinguishable, and is particularly in evidence on the sides of the neck, the shoulders, the outside of the fore limbs, and the upper side of the tail, where the black speckling is comparatively not very con- spicuous. The specks are stronger and more concentrated dorsally from the crown to the loins, making these areas, especially the back and loins, to all intents and purposes black. On the cheeks and sides of the neck the colour and pattern are normal. The throat, the insides of the limbs, and the hinder part of the belly are white, with small spots, the fore-part of the belly, where it passes into the chest, being washed with buff. The skin, which is unsexed and apparently unstretched, measures : head and body 3 ft. 11 in., tail (imper- fect a t the end) 2 ft. 6 in. ; total 6 ft. 5 in.+.

This skin, sent to the British Museum by Mr. F . Bowker, is labelled “40 miles N.E. of Grahamstown.”

A second specimen from Collingham, near Grahamstown (N. Abraham, B.M. no. 86.7.15.1), is much darker than the type, the back and flanks being completely black, and the pale ground-colour mostly obliterated everywhere ; but the outsides of the thighs show a zigzag pattern of confluent solid spots, set off by whitish interspaces and the cheeks, throat, belly, and insides of the limbs are white. The coat is a little longer, but is smooth, sleek, and not rough.

A third specimen from Grahamstown, figured but not described by Mr. W. L. Sclater, apparently resembles very closely the example from Collingham, so far as i t is possible to judge. The specimen recorded by Trimen, on the contrary, seems to have been like the type. It came from ‘‘ Bucklands, near Koonap.”

The only skull I have seen which from its locality can with some confidence be assigned to melanotica is preserved in the King Williamstown Museum, and was kindly lent to me by Mr. Shortridge. It came from East London (Miss Rossiter), and on the label is the statement that leopards have long been extinct in that neighbourhood. It is the skull of an adult male and W e r s very noticeably from all the other South African skulls I have handled ‘in having the frontal area low and flat and the occiput elevated. Hence the dorsal profile from the tip of the nasals to the occipital crest is much less convex. It is muscularly well developed, with a moderately high sagittal crest, and the temporal ridges well advanced and strongly concavo-convex between the postorbital processes. The bone is compact and solid. There is no history of the skull, and its unusual shape suggests that it may have belonged

* Not “ widely separated ” RE Lydekkur said.

(Text-figs. 7 & 8, a ) .

packed * as to obscure the deepish ochraceous tawny hue of the spaces betwe P


to a leopard reared in captivity. There is, however, no trace of the sponginess of the bony tissue commonly, but by no means always, associated with skulls of the larger F e l i b reared under such artificialconditions.

But according to the evidence the mutations are, or were, locally dominant, and since there are no other skins from Grahamstown available for examination, I think it wiser to restrict the name nzelunotica to this local type and not extend it

There is no doubt that the skins above described are mutations.

Text-figure 7.

Dorual

to the rest of the leopards of the Cape Province as Allen .suggested. At all events, i t is certain that the ground-colour of the typical example is much richer and darker than that of the leopards from Damaraland and Rhodesia, which are considered next.

Apart from the skulls from Damaraland recently received from MI. Short- ridge, there are in the British Museum only three South African skulls. These were collected many years ago and probably came from the Cape Province to the south of the Orange River. One is the skull of an adult male labelled


Cape (Molyneux), B.M. no. 87.4.25.1). It is a perfectly normal skull except for the smallness of the upper carnassial. Another is the defective skull of a fully adult female which, so fir as its measurements indicate, closely resembles adult female skulls of East African leopards, but is decidedly narrower on the average. 'It is labelled S . Africa (Burchell, B.M. no. 115 d ) . The third is also a defective female skull, tolerably closely agreeing with the last, but it is considerably younger and would probably have exceeded it in dimensions. It is merely labelled Dr. A. Smith (B.M. no. 45.7.3.1). But since it was received with a number of other mam- malian remains ticketed " Cape of Good Hope," there is very little doubt that it came from that district.

These three skulls are not distinguishable by any important characters from those of the larger Damaraland form procured by Shortridge, and described below.

The teeth are about the same size.

PANTHERA PARDUS SHORTRIDQEI POC. (Text-fig. 8, b.) Panthera parduo shortridgei Pocock, Abstr. Proc. Zool. SOC. 1932, no. 347,

Locality of type.-d ad. Skin and skull : Gangongo, 3560 ft. alt. on the Okavango River some 120 miles above the Okavango swamp in Western Caprivi, Damaraland.

Distribution.-Damaraland, Bechuanaland, and the Kalahari to the Orange River in the south and eastwards through Rhodesia to the Transvattl.

Ducription of type .Shot on Sept. 29th. F u r thick, but not noticeably rough. General colour a pale greyish or pallidly olivaceous buff with no bright yellowish or orange tint anywhere, rosettes of average or rather small size, their centres at most slightly darker than the ground-colour. Limbs tinted with a pale wash of buff to the paws. Tail with white dominant at the end.

Two additional examples were shot on the Okavango River, an adult d a t Diwai, about 3400 ft., near the Bagane Drift, on Aug. 21st, and a young female a t the Mahango Drift, about the same altitude, on Aug. 9th. Both these localities are close to tne Okovango Marsh. The male is very like the type, but a little more brightly coloured, with more buff in the interspaces and in the centres of the rosettes. The young female is decidedly brighter than either of the males, owing to the interspaces of the back and the centres of the rosettes being more rusty ochraceous. The coat, too, is much longer androugher. This specimen in general hue comes nearest to the typical East African leopard, usually known as suahelica ; but is not so well coloured on the flanks and limbs and has more white on the end of the tail.

The tlat skin of a male, without measurements or skull, from Karakowisa, half way between the Okovango and Grootfontein, about 4500 ft., shot on April 22nd, is practically identical with the skin from Diwai.

An adult male from Sandfontien in the Gobabis District of the Kalahari on the borders of Bechuanaland, 4100 ft., shot on Nov. 24th, has the coat full and soft, but not so rough as in the preceding specimens. It i R a decidedly pale skin, but the ground-colour is somewhat brighter, with more buff in the pelage, than in the type. The difference in the texture of the coat is probably seaaonal.

A young female, also from Sandfontein and shot in November, haa the coat full and soft, but longer and rougher than in the adult male from this locality,

p. 33.

Under side and inside of limbs white.


. . . . . . . . . . . . . . . . .

Sandfontein, d oldish . . . . . . . . . . . 4 4%

Mohango Drift, $) young . . . . . . 2 10%

the difference being probably a question of age. The colour is pale, the back and the outsides of the limbs being merely washed with pallid buff.

A purchased skin of an immature female from the Keetsmanshoop district, closely resembles the young female from Sandfontein in its pallid tint, but has the pelage noticeably longer.

In their pallid coloration, recalling that of the leopards of Persia and of the Red Sea littoral, these two specimens differ strikingly from the comparatively brightly coloured female from the Okovango River.

The following flesh measurements were recorded by Mr. Shortridge :-

2 5#

1 11%

Total.

ft. in. 7 1 4

G 11

6 10

4 9)

When this table is compared with that of the Kenya leopards compiled by Hollister, it will be noticed that the Damaraland males are a little longer in the head and body, but shorter in the tail. I suspect this discrepancy is due to a difference of opinion t19 to the point where the tail begins. At all events the total lengths of the Damaraland leopards is less, considerably less, than the average length of Kenya leopards including those in Wards list, an average estimated to be about 73 ft. (see p. 560).

The skull of the type of this race from Gangongo closely resembles in size and shape the skull of leopardus from the French Congo, but the muzzle is not so deep and the sagittal crest is larger. A sketch from the lateral aspect is subjoined (text-fig. 8 ,b , p. 586).

Judging from the teeth, which show considerable wear of the cusps, it is that of an oldish animal. The skull from Diwai is slightly longer and has the brow between the postorbital processes noticeably higher with the sagittal and occipital crests rather less well developed. The skull from Sandfontein is a little narrower than that of the type, and is very similar to the male skull labelled “Cape.” The Olifants Kloof skull is shorter in total length and broader in the waist than the one from Sandfontein. The one from Babi- babi, 52 miles to the east of Gobabis, is not so old as the preceding, as attested by the open median frontal and interparietal sutures. The waist is exceptionally narrow for a subadult skull. The Ovamboland skull, although smaller than the last, is old. The waist is narrow and long, and the temporal crests are as strongly arched on the forehead as in the East London skull. In size it is nearly midway between the largest skull from Okavango and that of the adult male from Central Kaokoveld. The point is of interest considering the localities where the three skulls were procured.

The only example of this sex ia that of a young specimen with complete milk dentition and a t most the points of the second set, still covered by the gums, showing. Cutting the carnaasiatls and upper canine out of the bone showed them to be as large aa in the two female S. African skulls mentioned above.

No adult female skulls were collected.

586 MR. R. I. POCOCE ON

The following skins appear to represent this race :- 6. Kakamar, 2300 ft., in the Upington District, north of the Orange River

in the southern Kalahari. A skin, without measurements or skull, kindly lent to me by Mr. G. C. Shortridge. The general hue is pale and the coat is quite long and rough, a little longer and rougher than that of the young female from the Mohango Drift above described.

A pale skin closely resembling the one from Grootfontein. The skull presents no points

Text-figure 8.

8. Sebakwi River, S. Rhodesia (F. C. Selous, 19.7.15.33).

a. Side view of abnormally ahaped ekull of adult male Leopard, proviaiondls identified a8

b. The eame of normally shaped ekull of type of Shortridge's Leopard (P.pardus shortridgei) from Panthem pardue melanotica, from East London.

Gangongo, Damamland. X b.

to distinguish it from those of P. p. shortridgei. It has the 8ame condylo- basal length as the skull from Diwai, Okavango.

I am also indebted to Mrs. Cooper for a brief inspection of two skins obtained by Major Coop& a t Umtali, which apparently represent this race ; and its eastward extension is attested by a mounted specimen in the Museum, shot by St. Hamilton on the Sabi River in the North-eastern Transvaal, which presents no features justifying its separation from P. p. shortridgei.

The following table gives the meaaurements of the skulls tentatively assigned to P . p. melanotica and of those of P. p. shortridgei from Damaraland and of P. p. puella, from the same country, described below.

1 L

ocal

ity a

nd s

ex.

I

P. p

. sho

rtri

dgei

.

Diy

ai, O

kava

ngo,

d a

d.

....

....

G

ango

ngo,

d a

d.

....

....

....

....

.

Sand

font

ein,

d a

d ...

....

....

....

...

Olif

ant's

Klo

of, d

ad.

..

....

....

B

abi-

babi

(G

obab

is),

d sn

bad.

. ...

;G

"xz?lkT

-

Eas

t L

ondo

n,

ad.

....

....

....

.

I Cape,

d a

d.

......

......

......

.....

S. A

fric

a (B

urch

ell)

, 0 a

d. .

....

I C

ape

of Q

ood

Hop

e (A

. Sm

ith)

,

9 im

m.

8.8

1.7

1.6

1.6

1.6

---

Tot

al

leng

th.

-_- 8.

9

9'1 -

-

9.2 -

91

9.1

8.5

+

I

1.7

' 2.3

1.4

! 1.9

I 1.3-

1.8+

1.4

, 2.2

~

-

P. p.

puo

lla.

Oka

rosa

ve, C

. Kao

kove

ld, 6 ad

.. K

ovav

es, S

. K

aoko

veld

, Q

imm

..

In E

nglis

h in

ches

. In

mill

im.

1 I

8.1

8.0-

Con

d. b

as. i Z

ygom

. le

ngth

. i

wld

th.

_-_

_I--

1 1.7

, 1.4

1 -2

1.4

+ 1.5

1.1

6.0

8.2

8.1

I 5.7

I 4.6

+ -

-

1 4.7

1 --_.--

8.5

~ 5.9-

8.3

I 6.0

2.0

2.1

1.7

1.6

+

Mar

rtoid

w

idtb

. - .-

3.7

3.8 -

-

---

3%

3-7

8.2

1 5.6

8.0

82

1 5.4

8.0-

1 5.4

6.1

I 4.2

1 7.4

I 5'1

I 7.1

5'0

1 6'3

1 4.3

6.2

1 4.1

-------

I

3.5

3'5

35

3'5 2.8

_-

3'2

+ 3.3

+ 2.9

2.9

Upp

er

carn

al.

---

27

23

23

23

---

28

26

25

25

24

---

25

24

20

20

I

Upp

er

cani

ne.

1s

16

13

13

_--

17

17

15

16

Low

er

carn

al.

_- -

17

, 20

16

16

17

14

15

10

10

18

18

14

15

588 ME. R. 1. POCOCK Od

PANTHERA PARDUS PUELLA POC. Panthera pardus puella Pocock, Abstr. Proc. Zool. SOC. 1932, no. 347, p. 33. Locality of type.-Okorosave, 4533 ft. alt., in Central Kaokoveld. Distributian.-Kaokoveld District of Damaraland. Coat of fully adult 6, shot at Okorosave, July 17th in midwinter, short

and smooth, very much as in typical examples of Kenya and Tanganyika leopards, shorter and smoother than in any of the examples assigned to P . p . slwrtrzdgei. Colour and pattern as in the examples of shortridgei from Diwai on the Okavango and Karakuwisa nr. Grootfontein. Immature female from Kaoko Otavi, 4566 ft., shot July 27th, resembling the male in colour and pattern, but with the coat a little longer and rougher, perhaps owing to its being immature; the pelage about the same as in the male examples of shortrzdgei, dated April, August, September, and November, but not so long or rough as in the young female from Mohango Drift.

(Text-fig. 9.)

Measurements in the flesh of the two examples recorded above are :-

Locality and sex. 1

Headand I 1 body. 1 Tail. Total. ---

ft. in. f t . in. ft. in. i Okorowve, oldish 2 4* 6 1 I

1 I Ksoko-Otsvi. $2 imm. 3 4 2 5 % 5 9 t

I

Mr. Shortridge labelled the male skin “ Typical of the smaller long-t,ailed hill race,” This description is borne out by the measurements, which show that this leopard is smaller than shortrzdgei, but has a comparatively longer tail. The difference in size between the sexes is also comparatively slight.

Four skulls of this small Kaokoveld leopard were procured, but only one is adult, namely that of the male from Okorosave. It is considerably smaller, especially in condylobasal length, than the skulls above considered from districts farther to the east and is poorly developed muscularly, the sagittal crest being low and the waist wide and short, as in typical female leopards. The carnassial teeth are as large as the smallest corresponding teeth of the larger Damaraland form, but the canine is a little smaller.

The skull of an immature male from Kovares in S. Kaokoveld is a little smaller. There are two immature female skulls, both with complete unworn dentition. It will be noticed that the teeth are very considerably smaller than those of the female from the Mohango Drift, Okavango. Moreover, the smaller of the two skulk from Kaoko Otavi is in condylobaaal and man- dibular length only very slightly longer than the Mohango Drift skull. ‘Never- theless it is much older, since its permanent teeth are fully erupted, whereas in the Mohango skull they are still deeply imbedded in the bone. Clearly the Mohango leopardess would hare grown into a considerably larger animal than the one from Kaokoveld.

From the table of measurement.e of tho skulls of P . p . puella (above) it may be seen that in the female the teeth are as small rn in the Somaliland dwarf leopard P . p . nunopardus, although the skulls are larger. The teeth of the male, on the contrary, are considerably larger than in that race.


SUMMARY OF THE RACES ADMITTED IN THIS PAPEE. 1. P. p. panthcra Schreber. Algeria and Morocco. From want of material

the characters of this ram are at present iindefinable. On the evidence is is a large leopard, but varieble in pattern and colour. Possibly more than one dist nguishable race inhabits the district.

A large, pale grey thick-coated leopard apparently most nearly allied to the Persian race (P. p . saxicdm), but differing in certain cranial characters and in its less luxuriant coat.

2. P. p. jarvisi, subsp. n. Sinai.

Text-figure. 9

Dorsal view of ekull of the type of Panthcra pardw puslla, an oldish male exhibiting female characters.

3. P . p. antinorii De Beaux. Erythrzea. A comparatively small short- coated leopard with the normal secondary sexual characters of the malo skull undeveloped as in nanopardw, from which it apparently difFers in its brighter eolouring and larger teeth at least in the typical specimens.

A large, very dark, dusky leopard addicted to melanism, the back frequently black. Skull like that of the large leopards of the lowlamb to the south.

4. P. p. admta POC. Highlands of Southern Abyssinia.

590 IR. R. I. POCOCK ON

5. P. p . nunopardus Thos. Italian Somaliland for the most part. A very small, pale, short-coated leopard with small teeth and the skull of the male apparently without well-marked secondary sexual characters.

6. P. p . brockmani, subsp. n. British Somaliland for the most part. Larger and on the average brighter coloured than numpardus, intermediate in size and coloration between i t and the Kenya race, which it resembles in the development of the normal secondary sexual characters in the male skull.

Kenya Colony, Tanganyika Territory, and Nyasaland. Resembling P . p . adusta in size, but much paler and richer coloured. Larger than P. p . brockmani and better tinted, the colour and pattern on the whole nearly uniform, so far as known, throughout the districts mentioned.

8. P. p . adersi, subsp. n. The Island of Zanzibar. Distinguishable from the mainland form by its smaller more numerous rosettes, apparently shorter tail, and by some cranial characters.

Egyptian Sudan. Not sufficiently well known to diagnose, but apparently a large leopard, very variable in size and pattern and closely resembling, if not the same as, the race next described.

The Lado Enclave and savannahs of the Belgian Congo north of the Ituri rain-forest-area. Evidently nearly resembling the typical East African form fusm (suah&m), but with a larger skull and much more variable in colour and pattern than in typical Tanganyika specimens.

11. P. p . ruwenzorii. Ruwenzori and Bivungain the Lake districts of the Divide, a t high altitudes. A brownish olivaceous leopard distinguishable from the lowland forms of adjacent districts by its darker hue, longish thick coat, and bushier tail.

A smaller leopard than P. p . chui, but, like it, very variable in colour, the brightest specimens a shade darker than P. p . fusca (swthelica), the greyest hardly distinguishable from P. p . leopardus.

West Africa from Senegambia to Spanish Guinea, a t least along the coastal forest district. A large dull-coloured leopard of a dusky, greyish olivaceous buff hue, with the pattern typically composed of smallish close-set rosettes, rarely large. Skulls of males, on the average, as large as in P. p . chui, or larger.

14. p. p . reichenowi. Savannah districts of the Kamerun and adjoining districts of Central Africa. Not well known, but apparently differing from p. p . leopardus in its brighter more tawny colour.

15. p. p . shortridgei. Damaraland and eastwards to Rhodesia south of the Zambesi. Intergrading with the East African P . p . fusca (swthelica), but a t least in its typical western form paler in tint and with a better coat.

16. p. p . puelh. Kaokoveld district of Damaraland. Distinguishable from p. p . shortridgei by its smaller size, the suppression of the secondary sexual characters of the male skull, and the unusual smallness of the teeth in the female.

17. p. p . melunotica. Grahamstown ; range elsewhere unknown, the name being restricted to the variety with disintegrated, multiplied spots.

These races intergrade with intergradation of environment. The following are the principal environmental colour variations :-

1. The '' savannah '.' or " veldt " type inhabiting scrub or bush country, yellowish-tawny leopards represented by the common East African form p . p . fwm (=smblica) , apparently by the West African P. p . reichemi

7. P . p . fusca (=suahelica).

9. P. p . pardus.

Skull unknown. 10. P . p . chui.

12. P. p . iturensis. The Ituri rain-forest area of the Belgian Congo.

13. P. p , leopardus.


and by the south African P. p . s h t r i d g e i , which, however, in the western portion of its range in Damaraland approaches the next.

2. The desert type occurring in arid, rocky, or sandy districts where the annual rainfall is comparatively slight and the vegetation sparse, pallid " stone- coloured," buffish grey leopards represented by P . p . jarvisi of Sinai and P. p. nunopardus of parts of Somaliland.

3. The Mountain type occurring a t high altitudes ; very dark tawny brown or deep olivaceous greyish leopards represented by P . p . adusta of the High- lands of Abyssinia and by P . p . ruwenzwii of Ruwenzori and other mountains of the Lake District. On Mt. Elgon the leopards show an approach to this type, being on the averitge darker and richer tinted than those of the lowlands of East Africa.

4. The Forest type, inhabiting the tropical rain-forest area of West Africa and represented by P . p . leopardus, dusky leopards not so dark as the mountain type, but darker and noticeably less richly tinted than the scrub or bush leopards.

5. Possibly also to the effects of the environment must be ascribed the '' dwarfing " of such forms as P . p . nunopardus of Somaliland, P . p . antinorii of Erythraea, and P . p . puella of Damaraland, in which the males fail to acquire the marked cranial modifications characteristic of that sex.

EXPLANATION OF THE PLATES.

PLATE I. Mr. Hayne's skin of the Sinai Leopard (Pantheta pardus jaruiui).

PLATE 11. Skin of the type of the Zanzibar Leopard (Pantheta pardus adersi) .

PLATE 111. Skin of a Leopard, identified as Panthcra pardus rutcenzorii, from the Birunga Rmge.

Photograph kindly supplied by the Hon. H. Hachisuka.

PLATE IV. Skin of an unidentilied Jaguar-like Leopard ticketed N'dola, N. Rhodesia, and given to

Mr. 0. C. Shortridge by Dr. Gordon Lancaster.

Documents

26. The Leopards of Africa