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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 77 +#2Band

December 1978 Desember

Part 2 Deel

ADDITIONS TO THE REVISED. LIST OF PRESERVED

MALTERVAL OF THE: EXTINCT CAPE, COLONY

QUAGGA AND NOTES ON THE RELATIONSHIP AND

DISTRIBUTION OF SOUTHERN PLAINS *ZEBRAS

Ree RAW

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na beskikbaarheid

van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

1, 2(1-3, 5-8), 3(1—-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9),

6(1, t.-p.i.), 7-4), 8, 911-2, 7), 10(1-3),

11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33

Kopieregnavrae na die Suid-Afrikaanse Museum

ISBN 0 908407 57 2

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

ADDITIONS TO THE REVISED LIST OF PRESERVED MATERIAL

OF THE EXTINCT CAPE COLONY QUAGGA

AND NOTES ON THE RELATIONSHIP AND DISTRIBUTION OF

SOUTHERN PLAINS ZEBRAS

Ray Be AU

South African Museum, Cape Town

(With 9 figures)

[MS. accepted 7 September 1978]

ABSTRACT

Previously unpublished photographs and data of two mounted skins of the extinct

quagga (Equus quagga quagga) and a further relic are added to the list of preserved material.

Photographs of the destroyed K6nigsberg quagga are reproduced.

Examination of plains zebra (Equus ‘burchelli’) skins from southern Africa and comparison

with the true Burchell’s zebra (Equus ‘burchelli burchellii’) and the quagga (Equus quagga

quagga) suggest that all belong to one species (Equus quagga) and are forms which are dominant

in certain areas within the continuous distribution, rather than subspecies. In addition, the true

Burchell’s zebra is considered to be still extant in Zululand and possibly Swaziland, if the

generally accepted but vague range of that ‘subspecies’ is ignored.

CONTENTS

PAGE

Introduction . , : : : : ; An PAM

Additional material ; : j , : 5 AS

Notes on Burchell’s zebra and quagga ; 3.) ll

Reintroductions of zebras. ‘ : : » 41

Acknowledgements . : 3 ‘ : : 5 ala

References . : : , é ; : 2 4 44

INTRODUCTION

Since the publication of the revised list of preserved material of the quagga

(Equus quagga quagga) a photograph of and additional data on the quagga at

the Zoological Museum, State University at Kazan, Soviet Union, have now

been obtained. In addition, a mounted quagga has been discovered at the

Muséum d’Histoire Naturelle Lyon, France, and a mounted foot at the Royal

Albert Memorial Museum at Exeter, England. These new data are presented in

accordance with the method followed by Rau (1974). The photographs included

in the description of the Kénigsberg quagga (Hilzheimer 1930), which was

destroyed at the end of the Second World War, are reproduced to complete the

illustrations of mounted skins which existed into this century.

Def

Ann. S. Afr. Mus. 77 (2), 1978: 27-45, 9 figs.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

In a monograph Antonius (1951), a world authority on zebras, included the

quagga and the Burchell’s zebras in the single species Equus quagga, as had been

done earlier (Pocock 1904; St Leger 1932). This view is supported by

Rzasnicki (1951), Mohr (1964) and Klingel (1969, 1972) amongst others.

There is, however, a second school of thought which divides them into two

species, Equus quagga and Equus burchelli (Ansell 1971). As the present study

has revealed that there is no definite division between the quagga and Burchell’s

zebras, the former view is supported.

ADDITIONAL MATERIAL

KAZAN— MOUNTED SKIN

Bigsal

Catalogue number: 109

Sex: female

Locality: unknown

Date of acquisition: 1843

Remarks on acquisition: bought in Hamburg from a Mr Brandt by Professor -

Eversman for the Zoological Museum of Kazan University

Fig. 1. Kazan quagga.

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 29

History of mount: bought as mounted specimen in bad condition and remounted ;

a second remounting (‘restoring’) was performed in 1969 by M. A. Zaslavski

in Leningrad, U.S.S.R.

Description of striking features: face clearly striped; shadow-stripes on neck;

posterior part of feet (fetlock-hoof) dark; the photograph seems to show

faint striping on buttock

Measurements: head-body 2,36 m* (2,42 m)*

tail 0,54m_ (0,53 m)

ear 0,14 m_ (0,17 m)

hind foot 0,56 m (0,54 m)

shoulder height 1,29 m (1,28 m)

State of preservation: appears to be good

Further material of same individual: none

Remarks: exhibited open in gallery

References: none known

LYON— MOUNTED SKIN

Fig. 2

Catalogue number: 6102

Sex: female

Locality: Kaffraria, South Africa

Date of acquisition: unknown

Remarks on acquisition: none

History of mount: appears to be the original mount

Description of striking features: face not clearly striped

Measurements: head-body 2,10 m

tail 0,40 m

ear 0,17 m

hind foot 0,50 m

shoulder height 1,15 m

State of preservation: fair

Further material of same individual: none

Remarks: exhibited; specimen not mentioned in previous literature; V. Eisen-

mann of the Muséum d’Histoire Naturelle, Paris, discovered this specimen

early in 1975 and kindly informed the author

References: M. Philippe, ‘La zoologie au Muséum d’Histoire Naturelle de Lyon’

(in preparation)

* Measurements received through two intermediaries are not identical.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

—

Fig. 2. Lyon quagga.

EXETER —FOOT TROPHY

Fig. 3

Catalogue number: A758

Locality: Bontebok Flats, British Kaffraria, South Africa (between present

Cathcart and Queenstown)

Date of acquisition: after 1869

Remarks on acquisition: from the collection of W. S. M. D’Urban, first curator

of Exeter Museum

Measurements (hoof only): anterior height 6,0 cm

width 6,7 cm

length 8,4 cm

References: Howes & Bamber (1970)

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 31

Fig. 3. Exeter quagga foot; anterior and lateral views.

KONIGSBERG QUAGGA

Fig. 4

The KOnigsberg quagga was destroyed at the end of the Second World War

when Schloss Waldhof, about 20 km from KGnigsberg, where museum specimens

had been taken for safe-keeping, was shelled and completely destroyed by fire

(pers. comm. Prof. Dr O. Koehler (1972), director of the K6nigsberg Museum

until 1945).

The preserved material of Equus quagga quagga, including the above

specimens, now consists of:

skins—23, plus | mounted head and neck

complete skeletons—7

skulls—20, including 7 from complete skeletons; at least 3 of the mounted

skins appear to contain skulls

loose skeletal parts—2 sets of foot bones; upper and lower incisivae in

portion of jaws

other—dried connective tissue, flesh, etc., removed from skin; nasal

cartilages and palate removed from skull (in alcohol) (Rau 1974: 58)

NOTES ON BURCHELL’S ZEBRA AND QUAGGA

During February to October 1977 approximately 400 privately owned skins

of plains zebra Equus quagga (burchelli) and mountain zebra Equus zebra

hartmannae were processed at a Cape Town tannery and examined by the

author. These originated from many different localities in southern Africa.

Skins of 113 plains zebras were photographed, the initial 88 non-selectively and

sy) ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Konigsberg quagga (destroyed), adapted from Hilzheimer (1930).

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 33

the remaining 25 selectively. The proportion of the two species processed

during a two-month period (February and March) was 88 plains zebra:

62 mountain zebra.

A great number of ‘subspecies’ and even ‘species’ of plains zebra have been

described. In bestowing names to the many variations, the stripe pattern has

been the principal, and often the only, character considered (Ansell 1971). This

is certainly true in the case of the ‘subspecies’ burchelli.

The diagnostic characteristics of the true Burchell’s zebra Equus quagga

burchelli are usually given as follows: whitish, unstriped legs except occasionally

some traces of stripes on the hocks (Pocock (1897: 52) accepts some stripes on

the ‘knees’ as well); basic colour of dorsal parts white to ochre and brownish;

interspaces with shadow-stripes, which may be broad and intense, or less

conspicuous; body-stripes not meeting the ventral midline.

Equus quagga antiquorum is defined by the striping extending on to the legs

to at least the ‘knees’ and hocks, but sometimes to the pasterns.

Equus quagga burchelli (E. burchelli burchelli) is one of the few still recog-

nized as a valid subspecies. The accepted distributional range of this subspecies

included the Orange Free State, the north-eastern Cape Province, southern

Botswana and the south-western Transvaal. As zebras no longer occur in these

regions, other than reintroduced specimens, the true Burchell’s zebra is usually

considered to have been extinct since about 1910.

The locality of the type specimen on which Gray (1825) based the descrip-

tion of the subspecies is between Campbell and the Vaal River. The specimen

figured as Equus quagga burchelli by Antonius (1951, fig. 22) originated from

British Bechuanaland (Botswana) and lived at Hagenbeck’s Tierpark, Germany,

during 1907-9. However, most preserved specimens of this subspecies lack

locality data, as do those individuals that lived at menageries or zoos and are

known from illustrations and/or descriptions.

It appears, therefore, that the accepted range is based on old reports rather

than collected material, and that the true range is not known. Indeed, different

opinions as to the range of Equus quagga burchelli have been expressed. Matschie

(1894) has included Zululand while Shortridge (1934) and Eloff (1967) extend

the range to Great Namaqualand (South West Africa). The concept of the

range of a subspecies within the continuous distribution of a species is open to

question, especially in the case of fast-moving and migrating animals such as

the plains zebra. With regard to zebra distribution, Tegetmeier & Sutherland

(1895) wrote: ‘Disputing, therefore, as to whether two closely allied animals are

specifically or sub-specifically distinct is almost a waste of words. We know that

a species spread over a wide area will change according to the conditions of life,

until at last the two extremes are so diverse as to be regarded as distinct species,

but no one can say where one species ends and the other begins, for they merge

gradually into each other.’

The uncertainty about the former range of the subspecies Equus quagga

burchelli has been aggravated by the indiscriminate use of the names ‘quagga’

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

and ‘Burchell’s zebra’, and by the difficulty in distinguishing between those two

forms as well as between the true Burchell’s zebra and its northern relatives.

Accurate assessment of old documents is, therefore, practically impossible.

Pocock (1897: 42) wrote 13 years before the alleged extinction of Equus quagga

burchelli: *. .. and on account of the general application of the term Burchelli

to any or all of its subspecies, we are in ignorance of the exact area that it now

occupies in S. Africa.’

If all the mounted zebra specimens preserved in museums had precise

locality data, there would be in all probability antiquorum-types from the

burchelli ‘range’ and vice versa. Since most older mounted zebras do not,

however, have conclusive locality data, it is accepted that apparent burchelli

specimens originate from the accepted burchelli range while antiquorum

patterned specimens are thought to be from further north. Since the plains zebra

has a continuous distribution, recent interferences by man excepted, within

which certain forms or variations predominate in certain areas, it follows that,

in intergradational areas between two forms, the one form will gradually

decrease in percentage of total numbers as the area of the next form is

approached.

Despite the fact that for most of the preserved or known plains zebra

specimens, which conform basically to the description of Equus quagga burchelli,

the place of origin is not known, they are accepted as belonging to that sub-

species, as 1s shown in the figures by Cabrera (1936) (Fig. 5) and Antonius (1951).

The subspecies Equus quagga antiquorum is presently considered to include

all the extant southern forms of the plains zebra, ranging from southern Angola

and (historical) Damaraland through Botswana and Transvaal to Zululand.

Consequently, individual variation within antiquorum is considerable (Fig. 6).

Eloff (1967) stated that the three subspecies antiquorum, chapmani and wahlbergi

occur together in Zululand. Antonius (1951) had observed that in Damara zebra

and especially in Wahlberg zebra burchelli-like individuals do occur.

However, such individuals originate from outside the presumed range of

the true Burchell’s zebra and are consequently not accepted as such. Neverthe-

less, some authors are cautious. Cabrera (1936: 94), while saying that the

subspecies ‘appears to be extinct’, also states, ‘in fact, I regard as typical

burchelli all members of the species having all four legs white from the elbow

and the stifle joint, except for the occasional presence of a few short markings

across the hocks, and with the thighs free of complete and well-defined dark

stripes behind the stifle stripe’. Dorst & Dandelot (1970) list the subspecies

burchelli as ‘probably extinct’.

Amongst the extremely variable skins (many with numerous small, light

dots in stripes and shadow-stripes) examined at the tannery, were several

specimens from a Zululand game farm with a surprising combination of light

basic colour and few and/or weak shadow-stripes and considerable stripe

reduction (Fig. 7A). It is generally believed that the change in coloration of

plains zebras to the south of their distributional range is achieved by stripe-

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 35

Fig. 5. Examples of true Burchell’s zebra, Equus quagga burchelli, shown in standardized outline.

A. Type of ‘Equus burchelli paucistriatus’ Hilzheimer, 1912, at Mainz Museum. B. Animal that

lived at Jardin des Plantes, Paris, 1826. C. Type of subspecies from Little Klibbolikhonnifontein,

British Museum (Natural History) (destroyed). D-—E. Animals that lived at Knowsly Park,

about 1845. F. Animal that lived at Zoological Garden, Dresden. (Adapted from Cabrera

(1936).)

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Examples of Equus quagga antiquorum, shown in standardized outline. A. From

Rietfontein West, Munich Museum. B. Type of subspecies from Angola (Smith 1841).

C. From eastern Transvaal. D. From west of Lake Ngami. E. From Zululand.

F. From eastern Transvaal. (Adapted from Cabrera (1936).)

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA SY;

reduction and the darkening of basic colour and the occurrence of shadow-

stripes. However, the above specimens indicate that these processes do not

necessarily occur together. Indeed, there are specimens from other areas which,

in spite of more extensive striping, display dark basic colour (Fig. 8A), some

with numerous strong shadow-stripes (sometimes three in one interspace). From

the Zululand skins (Fig. 7) and additional photographs it appears that stripe-

reduction is the dominant feature in this eastern population, while to the west

on the interior plateau the darkening in both basic colour and through shadow-

stripes is more pronounced.

In terms of stripe reduction these Zululand skins certainly do not fit into

the concept of Equus quagga antiquorum (Fig. 6), but they do agree with the

description of Equus quagga burchelli. Indeed, a few are less striped than some

of the burchelli specimens figured by Cabrera (1936) (Fig. 5) and others. It

appears, therefore, that Equus quagga burchelli is, in fact, not extinct but still

survives in Zululand (and possibly Swaziland), and that its range does extend

eastward into Zululand, as Matschie (1894) stated. (The ‘myriads of Quaggas’

which Chapman (1868, cited in Antonius 1951: 96) encountered in the extreme

north-eastern Orange Free State could thus have been true burchelli, and not as

Antonius (1951: 96) considered, Wahlberg zebra.) Otherwise the subspecies

burchelli would have to be considered as part of the antiquorum complex, a

thought similarly expressed by Rzasnicki (1951) and Roberts (1951), cited in

Ansell (1971). The only other alternative would be not to recognize such

individuals (Figs 7B—D, 9B, D) as members of the subspecies burchelli. However,

had they been old mounted specimens, they would be regarded as true Bur-

chell’s zebras.

It must be noted that the farm where these specimens had been shot is a

hunting farm which was originally stocked with 300 zebras from Hluhluwe

Game Reserve, and that hunters select the most striped specimens (C. Tinley,

pers. comm.). Selective breeding is thus exercised unintentionally and this might

have ‘re-created’ burchelli, although similar specimens do occur in Hluhluwe

and Umfolozi Game Reserves, the only area in Zululand where the original

zebra population has not been interfered with through reintroductions

(P. M. Brooks, pers. comm.).

Zululand undoubtedly is an inter-gradational area between the two forms

burchelli and antiquorum. Observations made during a recent visit to that area

and Swaziland suggest that the percentage of burchelli types may be as high as

15 in Zululand, although P. M. Brooks, biologist at Hluhluwe, estimates it to be

5-10 per cent (pers. comm.).

The north-eastern and eastern parts of Swaziland also retain an endemic

population of plains zebra (T. E. Reilly, pers. comm.). There is no geographic

barrier between Zululand and the Swaziland lowveld, and it is not surprising

that some of the zebras observed there fit or approach the description of Equus

quagga burchelli, though the percentage of such specimens must be expected to

be lower than further south.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Skins of plains zebras from Zululand, 1977. A. Note stripe-reduction, few shadow-

stripes. B—-D. Note shadow-stripes on neck and stripe-reduction. (B now at South African

Museum, ZM38819.)

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 39

4 ee :

(sg)

Fig. 8. Skins of plains zebras from southern Africa. A. Precise locality unknown; note dark

basic colour of upper parts compared to lower leg. B. Precise locality unknown; note numerous

shadow-stripes and extensive striping. C. From game farm in Zululand; note dark basic colour

of upper parts. D. From southern Kaokoveld, South West Africa; note little contact between

body-stripes and dorsal midline.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Despite the uncertainty whether the ‘quaggas’ of old reports were, indeed,

the true quagga, Equus quagga quagga, the northern boundary of its range is

usually considered to have been formed roughly by the Orange and Vaal rivers.

The eastern limit might have been about the boundary between the Orange Free

State and Natal (Grout, cited in Shortridge 1934: 402). It is also generally

accepted that the ranges of quagga and true Burchell’s zebra overlapped in the

Orange Free State (Antonius 1951; Liebenberg 1964; Ansell 1971).

Antonius (1951) was of the opinion that the quagga, being essentially a form

adapted to the Karoo and possibly the most distinct of all the plains zebra

subspecies, expanded its range secondarily in a northerly direction and met and

overlapped along a broad front with the southward expanding Burchell’s zebra.

He obviously did not believe in reports about the social and reproductive

separation of the two forms, since he considered one of the preserved quagga

specimens (the more striped specimen of ‘Equus burchelli paucistriatus’ at Mainz

(Fig. 9A and Rau 1974, fig. 13) as a possible cross between the two forms

(Antonius 1951: 26).

It is difficult to accept that separate herds of the two forms existed in the

same locality. Even if they did represent separate species, as some are inclined to

believe, herds formed by members of both must be expected, since this is known

to occur in extant zebra species where their ranges overlap. For example, Equus

zebra hartmannae overlaps with Equus quagga antiquorum in South West Africa

(Antonius 1951: 82) and Equus grevyi forms mixed herds with Equus quagga

boehmi in Kenya (Antonius 1951: 82; Bartlett 1963). Even true quaggas were

seen together with the Cape mountain zebra at Geelbeckfontein, at the southern

limit of the Karoo, by Barrow (1801, cited in Antonius 1951: 82).

As shown previously (Rau 1974) the quagga and the true Burchell’s zebra

approach one another in basic colour and in stripe reduction, so much so that

some of the preserved specimens (e.g. Rau 1974, fig. 12 right) have variously

been regarded as one or other of the two. Furthermore, some of the charac-

teristics previously considered to be typical of the quagga (Rau 1974) have been

observed in the skins at the Cape Town tannery and in a specimen from Kruger

National Park. These characteristics include narrow interspaces (Smuts 1974,

fig. 1.4.5), dorsal midline being flanked by a light band on either side, and body-

Stripes not in contact with dorsal midline except sometimes via narrow bridges

(Fig. 8D).

Antonius (1951: 29) has already pointed out that the quagga had reddish

dark-brown stripes which, especially on the body, merge into the reddish basic

colour. It has previously been explained why the coloration of the quagga has

been incorrectly interpreted as light stripes on a dark background (Rau 1974: 43).

Careful examination reveals beyond all doubt that the quagga, like all the other

plains zebras, is dark-striped and that the interspaces darken progressively from

the shoulder in a posterior direction until they become equal in tone with stripes

and shadow-stripes to produce the ‘uniform’ posterior portion. In other southern

plains zebras the interspaces are also lightest on head and neck. Ansell (1971: 5)

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 4]

and others are wrong in defining the striping in the quagga as ‘whitish on dark

background’.

Since identification is therefore complex, and since it is impossible to

observe the stripe pattern of each individual when looking at a herd, the alleged

recognition of separate herds of quaggas and Burchell’s zebras in the same

locality must be treated with suspicion.

The grouping of plains zebras into subspecies is thus somewhat vague, or,

as it is expressed by Ansell (1971) ‘has never been satisfactory’. In an attempt to

sort out this confusion, Pocock (1897) pointed out that throughout the distri-

bution area of the plains zebra ‘intermediate types’ do exist between particular

forms which prevail in particular localities. In addition, as mentioned earlier,

typical representatives of the two forms occur in intergradational areas, the

closer to the area of the one form the lower the percentage of the other form.

The high degree of variability, both between geographically distant

populations and within one herd, has been attributed by Antonius (1951) to the

process of eliminating the contrasting stripe-pattern. It should be noted that

Antonius’s ‘Riickbildung der Bein- und Rumpfstreifung’ has been misinterpreted

by Eloff (1966, 1967) as being a sign of genetic pauperization.

As shown above, the gradual change from one form to another in plains

zebras appears to include, at least in exterior characteristics, the ‘separation’ into

Equus quagga and Equus burchelli. Relatively few skulls of the true quagga are

available and some of these lack sufficient data to establish their authenticity.

The fact that some skulls may have been obtained by breaking up an old stuffed

quagga skin (as has happened at Berlin, where the skull was later identified as

that of Equus caballus (Antonius 1951: 35)) is no guarantee of their identity as

Equus quagga quagga.

However, some zoologists believe that there are specific morphological

differences between skulls of the quagga and the plains (Burchell’s) zebra

(e.g. Cooke 1950; Lundholm 1951; Gentry 1975; V. Eisenmann, pers. comm.).

The recent study by V. Eisenmann (pers. comm.) has shown that in some skull

characters there is a cline from northern to southern plains zebras. In other

words, in some respects at least skulls of southern plains zebras approach those

of the quagga more closely than do northern forms. This conforms to the

well-documented cline in colour and stripe-pattern.

To conclude, it would appear that the Burchell’s zebras and the quagga

should be regarded as one species, Equus quagga. Furthermore, since the

grouping into subspecies, either by appearance or by distribution, is not clear-cut

in the case of the plains zebras, it might be preferable to refer them to forms,

types or variations prevailing in certain areas, rather than to subspecies.

REINTRODUCTIONS OF ZEBRAS

The reintroduction of animal species into areas where natural populations

have been exterminated is increasingly common. There are, however, problems

with this practice, some of which have been discussed by Greig (1977).

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

A Vi BRAN CS <i B V. BRANCO

D V BRANCO

Fig. 9. Stripe patterns and tone of basic colours of various plains zebras, shown in standardized

outline. A. Female quagga at Mainz Museum. B. Burchell’s zebra from Zululand (same as

Fig. 7B). C. Male Burchell’s zebra at Leiden Museum. D. Burchell’s zebra from Zululand

(same as Fig. 7D). E. Quagga at Tring Museum (near London). F. Type of true Burchell’s zebra,

British Museum (Natural History) (destroyed). G. Quagga at Berlin Museum. H. Male

quagga at Mainz Museum. I. Burchell’s zebra at Mainz Museum (type of ‘Equus burchelli

paucistriatus’ Hilzheimer, 1912).

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 43

Tes

ea aN H ae S Mii, | Sp &>

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Both plains and mountain zebras are amongst those South African

mammals whose ranges have been much restricted during the past three

centuries. Now, and in the recent past, zebras are being translocated from areas

where populations survive to areas which they formerly inhabited. In the case

of the plains zebras such reiniroductions have involved the movement into

certain areas of forms dissimilar to those which previously occurred there, even

though the original form, or one close to it, still exists.

As indicated above, the ‘subspecies’, Equus quagga burchelli, apparently

survives in Zululand and thus, when reintroductions of zebras into the former

range of the true Burchell’s zebra are contemplated, it would be desirable that

selected specimens from Zululand be used, rather than animals of the antiquorum

type. It would even be desirable to eliminate former indiscriminate reintro-

ductions and to replace them with individuals which are closer to the original

population. With a minimum of initial selective breeding in order to eradicate

recessive antiquorum characteristics, which are present also in burchelli-like

individuals from Zululand (an intergradational area between the two forms),

a ‘pure’ population of plains zebras resembling the former endemics could be

achieved within a relatively short period.

ACKNOWLEDGEMENTS

My sincere thanks are due to Dr Vera Eisenmann of the Muséum National

d’Histoire Naturelle, Paris, for kindly informing me of the mounted quagga

which she discovered at the museum at Lyon, France. I also thank Dr C. A.

Howes for drawing my attention to the quagga foot which is housed at the

Royal Albert Memorial Museum, Exeter, England, Dr I. Sokolov of the

Academy of Sciences, Leningrad, U.S.S.R., and Mr E. Granqvist of the

Zoological Museum, Helsinki, Finland, for being instrumental in obtaining

photographs and data of the mounted quagga in the Museum at Kazan, U.S.S.R.

I am indebted to Mr V. Branco for producing the drawings, Mr A. Byron for

some of the photographic reproductions and to Prof. G. H. Findlay, and

Drs Q. B. Hendey and P. A. Hulley for their co-operation.

REFERENCES

ANSELL, W. F. H. 1971. Perissodactyla. Jn: MEESTER, J. & SETZER, H. W. eds. The mammals of

Africa. An identification manual 14: 1-14. Washington, D.C.: Smithsonian Institution

Press.

ANTONIUS, O. 1951. Die Tigerpferde: die Zebras. Monogrn Wildsdugetiere 11: 1-148.

BARTLETT, D. 1963. Zeehorses and Zebronkeys. Animals 2: 394-396.

CABRERA, A. 1936. Subspecific and individual variation in the Burchell zebras. J. Mammal.

17: 89-112.

Cooke, H. B. S. 1950. A critical revision of the quaternary Perissodactyla of Southern Africa.

Ann. S. Afr. Mus. 31: 393-479.

Dorst, J. & DANDELOT, P. 1970. A field guide to the larger mammals of Africa. London:

Collins.

ADDITIONS TO THE REVISED LIST OF THE EXTINCT QUAGGA 45

Exorr, G. 1966. The passing of the true quagga and the little Klibbolikhonnifontein Burchell’s

zebra: which is to be next the Cape mountain zebra or Wahlberg’s zebra of Zululand?

Tydskr. Natuurwet. 6: 193-207.

Etorr, G. 1967. The phenomenon of the obliteration of stripes of the Burchell zebra with a

southerly migration. Zodn. Pretoria 8: 10-13.

Gentry, A. W. 1975. A quagga, Equus quagga (Mammalia, Equidae), at University College,

London, and a note on a supposed quagga in the city museum, Bristol. Bull. Br. Mus. nat.

Hist. (Zool.) 28: 217-226.

Gray, J. E. 1825. A revision of the family Equidae. Zool. J. Lond. 1: 241 248,

GreiG, J. C. 1977. The principles and problems of genetic conservation in relation to wildlife

in South Africa. Cape Dept. of Nature & Environmental Conservation, Research report,

September 1977: 1-55. (Mimeographed.)

Hizuermer, M. 1930. Das Konigsberger Quagga. Z. Sdugetierk. 5: 86-95.

Howes, C. A. & BAMBER, M. 1970. Rarities in a museum. Oryx 10: 326-328.

KLINGEL, H. 1969. The social organisation and population ecology of the plains zebra, (Equus

quagga). Zoologica afr. 4: 249-263.

KLuINGEL, H. 1972. Das Verhalten der Pferde. Handb. Zool., Berl. 8 (49): 1-68.

LIEBENBERG, L. C. C. 1964. Die grotere soogdiere wat vroeér dae voorgekom het in die

omgewing van die Golden Gate-Hooglandpark. Koedoe 7: 99-104.

LUNDHOL”M, B. 1951. A skull of the true quagga (Equus quagga) in the collection of the Transvaal

Museum. S. Afr. J. Sci. 47: 307-312.

MatsculE, P. 1894. Die afrikanischen Wildpferde als Vertreter zoogeographischer Subregionen.

Zool. Gart. Lpz. 35: 33-39; 65-74.

Monr, E. 1964. Von Rosshauten und Zebrafellen. Das Pelzgewerbe 15: 161-168.

Pocock, R. I. 1897. The species and subspecies of zebras. Ann. Mag. nat. Hist. (6) 20: 33-52.

Pocock, R. I. 1904. The Cape Colony quaggas. Ann. Mag. nat. Hist. (7) 14: 313-338.

Rau, R. E. 1974. Revised list of the preserved material of the extinct Cape Colony quagga,

Equus quagga quagga (Gmelin). Ann. S. Afr. Mus. 65: 41-87.

Roserts, A. 1951. The mammals of South Africa. Johannesburg: Trustees of The mammals of

South Africa Book Fund.

RZASNICKI, A. 1951. Zebras and quaggas. Annls Mus. zool. pol. 14: 203-251.

SHORTRIDGE, G. C. 1934. The mammals of South West Africa 1: 397-411. London: Heinemann.

SmitH, C. H. 1841. The natural history of horses. Edinburgh: Lizars; London: Highley;

Dublin: Curry. (The naturalist’s library [Vol. 31]. Mammalia 12.)

Smuts, G. L. 1974. Growth, reproduction and population characteristics of Burchell’s zebra in

the Kruger National Park. Unpublished D.Sc. thesis, University of Stellenbosch.

St Lecer, J. 1932. On Equus quagga of south-western and eastern Africa. Ann. Mag. nat.

Hist. (10) 10: 587-593.

TEGETMEIER, W. B. & SUTHERLAND, C. L. 1895. Horses, asses, zebras, mules and mule breeding.

London: Horace Cox.

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

noy., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text __

e.g. *... the Figure depicting C. namacolus...’; *. .. in C. namacolus (Fig. 10)...’

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by initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

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Punctuation should be loose, omitting all not strictly necessary

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book or article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’ a ieee

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

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