40 S.-Afr. Tydskr. Dierk. 1994, 29(1) Plio-Pleistocene Hyracoidea from Swartkrans Cave, South Africa

McMahon, C.R. Mammal Research Institute. University of Pretoria, Pretoria, 0002 J.F. Thackeray Department of Palaeontology and Palaeoenvironmental Studies, Transvaal Museum, P.O. Box 413. Pretoria, 0001

Received J February 1993; accepled 15 July 1993

Swartkrans Cave, an important Pleistocene hominid site in the Sterklontein valley, has yielded abundant fossil hyracoid remains. Two extinct taxa, Procavia antiqua and P. rransvaalens;s, have been previously reported to occur at the site in deposits postdating 1,8 million years before the present (B.P.). The extant taxon, P. capens;s, has been reported in the most recent deposits, thought to include the Terminal Pleistocene. However, statistical analyses of teeth suggest that P. antiqua and P. capensis are conspecific. P. transvaalensis differs from P. capensis by being larger and more hypsodont. The coexistence of these taxa in palaeoenvironments of the Sterkfontein valley is attributed to dietary differences: P. transvaalensis being dependent primarily on grass foliage in grassland habitats, P. capens;s being a generalist, feeding on a range of different kinds of vegetation in rocky habitats. Hypoconid-metaconid distances, previously used by Broom to distinguish taxa, are shown to be related to individual age. Crown height data of molars are used to generate age distributions, and incisors are used to obtain sex ratios, Predation on adults 01 P. transvaa/ensis, Irom populations associated with a sex ratio biased towards males. may have contributed to the decline in relative abundance and ultimate extinction of this species towards the end of the Pleistocene. Predators can have included leopards. black eagles and/or hominids.

Swartkrans is 'n belangrike grot in die Sterkfontein vallei, met waardevolle 10ssiele uit die Pleistoseentydperk.. Bene en tande van twee uitgestorwe dassiespesies, Procavia ant;qua en P. transvaalens;s, word in afsettings van minder as 1,8 miljoen jaar voor hede aangetref. Fossiele van die huidige spesie, P. capensis. is in die mees onlangse afsettings gevind, moontlik uit die eind·Pleistoseen. Statistiese tandanalises het egter aan die lig gebring dat P. antiqua en P. capensis dieselfde spesie is. Die maaltande van P. transvaalensis is grater en langer as die van P. capen sis. Die gelyktydige bestaan van hierdie twee spesies in die paleo-omgewing van die Sterkfontein-vallei word aan 'n verskil in eetgewoontes toegeskryf: P. transvaalensis is hoofsaaklik op grasagtige voedsel aangewese, terwyl P. capensis op algemene plantegroei in rotsagtige omgewings leef. Metings van spasies tussen tandkrone, voorheen deur Robert Broom geneem om spesies te onderskei, het .

) eerder die ouderdom van individue aangetoon. Hoagie van tandkrane kan ook aangewend word om 9

0 ouderdomsverspreiding te bepaal. Die vorm van snytande word gebruik om geslagte van mekaar te 0

2 onderskei. Predasie op volwassenes van P. 'ransvaa/ensis uit populasies waar die vroulike diere in die

d minderheid was, mag tot die uiteindelike uitsterwing van hierdie spesie aan die einde van die Pleistoseen e t gelei het. Roofdiere kon luiperds. swart arende en hominide ingesluit het. a d ( r e Swartkrans is an important Pleistocene site in the suggested that P. anliqua was ancestral to modem dassies. h s i Sterkfontein valley, well known for its hominid fossils

l There is, howevcr, a major problcm with Broom's analysis. b (Auslralopili7ecus and early lIomo) in deposits dated Apart from not compiling metric data, he did not consider u P

between about 1,8 and 1,0 million years before the present individual age and tooth wear in relation to variables such as e h (B.P.) (Brain 1981; Bmin el al. 1988). The site also contains hypocone-metacone distance which he had used to distin­ t

y abundant hymcoid remains which may have been brought to guish P. anriqua from P. capen"is. Churcher (1956:499) had b the site by various agents of accumulation, including in fact noted similarities between these two taxa, but begged d e t leopards, black eagles and/or hominids. The principle aim of the question of their synonomy. It is therefore relevant to n a this study is to examine all specimens of Procavia that have reconsider the taxonomic status of P. antiqua from r g been recorded from Swartkrans, and to determine how many Swartkrans. e c species are actually represented at the site. Panicular If it is possible to demonstrate that P. anriqua and P. n e attention is given to specimens previously attributed to P. c capen.,is are conspecific, it would still be necessary to i l [extinct, larger than

anriqua; P. transvaalensis P. anliqua

r consider evidence for the coexistence of allcasl two species e and with P. obermeyerae as a junior synonym (Broom 1936; d of Procavia at SwartJ

n Churcher 1956)]; and P. capensis (extan~ compamble in

u Iransvaalensis and a smaller taxon, previously atuibulcd to size and morphology to specimens attributed to P. anriqua). y P. anliqua, comparable in size lo P. capenJis. In this paper a Sex mtios and age distributions are potentially useful for w the differences in dentition that could possibly relleet e purposes of assessing possible factors that can have t differences in diet arc looked at, thereby helping to account a contribulCd to the extinction of P. Iransvaalensis. G Broom (1934) had attempted to distinguish P. anriqua for the coexistence of two species of Procavia in the t e Sterkfontein valley in the Pleistocene.

n from the extant form, P. capensis, on the basis of crown i b height of molars, the distance between hypocone and meta­ a Materials and Methods S cone cusps, and the bunodont condition for the protocone. y

b He claimed that these were primitive characteristics, and Modem specimens of P. capensis, measured for com para- d e c u d o r p e R s. Afr. 1. 7-"01. 1994. 29(1) 41

tive purposes, were made available from collections at the sexing hyraxes. Males are characterized by having first Transvaal Museum. For purposes of this study, we accept incisors which arc triangular in cross-section. whereas those the taxonomy of extant hyracoid species as reviewed by of females are more rounded (Thomas 1892). The dichoto­ Meester, Rautenbach, Dippenaar & Baker (1986), who my between triangular and rounded incisors in specimens regard P. capensis as the only extant species of Procavia, attributed to P. anliqua and P. transvaalensis is assumed to whereas Bothma (1971) accepted five. All of the fossil reflect sexual differences. as in P. capensis. Minimum material used in the present study comes from the Swart­ numbers of individuals (MNI) were calculated for species in krans cave in the Sterkfontein Valley, within IO km north of each member by counting the number of upper incisors in Krugersdorp (Brain 1981; Brain e/ al 1988). each assemblage, and dividing by two in order to take left Brain has recognized five depositional unil5 at Swart­ and right sides into account. krans. Members I, 2 and 3 are thought to have been deposi­ Studies of P. capensis have also shown that degree of ted between about 1,8 and 1,5 million years B.P. Member 4 tooth wear and eruption sequences can be used to estimate dates to the Late Pleistocene and has stone artefaCI5 individual age (Millar 1971; Fairall 1980; Fourie 1983). attributable to the Middle Stone Age. However, Member 4 Fouric's non-linear relationships between age and crown contains few faunal remains. Member 5 is thought to include height arc certainly useful for purposes of studies of the Tenminal Pleistocene, and has been dated to about specimens attributed to P. capensis, but difficulties arise II ()()() B.P. by the radiocarbon dating technique (Brain el when attempting to apply such relationships to extinct taxa al. 1988). All the other members have been dated using such as P. lransvaalensis which is larger than P. capensis. faunal remains, compared to securely dated East African Although Fourie's equations cannot be used to obtain relia­ sequences (Vrba 1975; Brain el al. 1988). P. capensis was ble age estimates for both taxa, crown height and wear reported by Brain el 01. (1988) to occur only in Member 5. patterns can be used to place specimens within relative age P. anliqua was documented in Members I, 2 and 3, together classes. with P. Iransvaalensis which has been found in Member 5. No hyraxes were recorded in Member 4. Results Unfortunately the type specimen of P. anliqua from Table 1 presenl5 the number of upper incisors (left and Taung was reported missing after it was described by right), and minimum numbers of individuals (MNI) for male Broom (Churcher 1956). In its absence, a well-preserved cranium of another specimen, St 105 from Sterkfontein (fonmerly P. roberlsi, a junior synonym of P. anliqua), is used here as a substilUte for the missing type. St 106 from Uitkomst (also known as Gladysvale) has been listed as the . ) type specimen of P. obermeyerae, but as a junior synonym 9 M. 0 for P. transvaalensis (Shaw 1937; Churchcr 1956), it is used 0 2 as a reference specimen for the latter. St 105 and St 106 arc d e housed at the Transvaal Museum. t a Churcher (1956:482) drew attention to morphological d ( differences between P. IransvaaJensis and P. capensis, r e

h notably to the occurrence of double-rooted milk canines s i Figure 1 M' of P. capensis, indicating hypocone (Hy) and l with complex crowns in at least two specimens of the larger b melacone (Me) cusps, used by Broom (1934) for taxonomic u taxon. However, in this study attention is focussed on metric

P purposes. data (mesiodistal and bucco-lingual diameters) for purposes e h

t of making distinctions between P. transvaalensis and other

y specimens of Procavia. Tooth dimensions were measured b Table 1 Numbers of first upper incisors

d according to criteria given by von den Driesch (1976). All of male and female fossil dassies e t measurements were recorded in millimctrcs. For taxonomic

n attributed to P. antiqua from Members 1, a purposes attention is focussed on Ml or M2, since the cha­ r 2 and 3, P. capensis from Member 5 g racteristics initially used by Broom (1934) for distinguishing and P. transvaalensis from Members 1, e c P. anliqua are best found in uppcr molars. n 2, 3 and 5 at Swartkrans; minimum e c Broom (1934) used hypo-metacone distance in M' and numbers of individuals (taking right and i l

r M' (Figure 1) to distinguish P. anliqua from other dassies, left incisors into account) indicated in e

d but remarkably he never published metric data. In this study parenlheses n

u we provide measurements of hypocone-metacone distances P. allljqua P. Irafl.SWlaiefl.Sis y together with data on crown height in the same specimens. a w On account of small sample sizes for each member, data Memocr Males Femalc~ Males Female~ e t for Members I, 2 and 3 were combined in some instances to a 35 (IS) 16 (8) 6 (3) 2 (I)

G fonm one early Pleistocene sample, for comparison with

t 2 15 (8) 9 (5) I (I) 0(0) e Late Quaternary material from Member 5. 9 (5) n 3 II (6) 3 (2) 3 (2) i b a Sexing specimens and age estimation P. capensis P. traf1SWlaiensis S 5 23 (12) 21 (II) 3 (2) 2 (I) y First maxillary incisors of P. capen sis provide a means of b d e c u d o r p e R 42 S.-AfT. Tydskr. Dicrk. 1994,29(1)

and female specimens previously atlTibuted to P. anliqua, P. individuals for each taxon arc placed in age class I and the transvaalensis and P. capensis in each member. Males oldes1 in age class 10. The results arc ploUed as histograms consistendy outnumber females with the exception of a in Figure 3 (combining data from both M' and M'). The small sample of P. transvaalensis from Member 3, in which there arc equal numbers of males and females (Table I). Table 2 presents means and standard deviations for M' Table 3 Minimum numbers of individuals of dimensions of modem and fossil specimens, ploued in hyraxes in age classes based on wear patterns Figure 2. described by Fourie (1983); age classes of P. Table 3 lists the number of individuals within six age transvaalensis not directly comparable to those classes, based on wear patterns previously descrihcd by of P. antiqua and P. capensis on account of size Fourie (1983). Most specimens attributed to P. capensi .. and differences P. anliqua are represented in age class 2 (young indivi· P. a1l1iqWJ P. capul.fis P. fralUvaalensis duals). By eonlTasl, most of the individuals of P. transvaal· ensis are represented in age classes 4 and 5 (older Oass I (young) 2 0 0 individuals). Oass 2 16 4 2 Crown height measurements of Ml and M' have been Oass 3 7 2 Oass 4 placed within arbitary age classes, in which the youngest 8 5 Oass 5 I 0 5 Oass 6 (old) 0 0 0

Table 2 Means and standard deviations of M' Totals 34 6 14 dimensions (mesiodistal length and buccolingual width) for specimens attributed to P. antiqua from Members 1, 2 and 3 (combined sample) and Member 5; P. capensis from Member 5; mocern specimens of P. capensis; and (a) PROCAVIA ANTIOUA/CAPENSIS (n=62) P. transvaalensis from Members 1, 2, 3 and 5. 24 Measurements in mm 22 W U Z 20 I Mesiodi~taJ lenglh Ruccolingual width W II: 18 ~,, '1 II: s.d. mean s.d. n :::l ,. me'" U U 14 P. allliqua a w 12 Members 1-3 8,31 1,19 7,74 1.25 29 .

) 10 P. capen.su I-" 9 '"Z 0 Member 5 7,88 0,68 7,30 0,51 17 W 8 0 U

2 Modem s.ample 8,51 0,30 7.87 0,52 23 II: W • d Member 5 and Il. 4 e t

a modem sample 8,24 0,58 7,63 0,60 40 2 d

( P. rTCUlSYoo/ellSjs r

e (Members 1-5) 10,32 0,70 9,53 0,66 18 h s i l b (b) PROCAVIA TRANSVAALENSIS (n=22) u 24 ~

P 22 e W e E p. transvaalensis U h Z 20 ~ t

W y Members 1 and 2 '''-''.....1 II: 18 - II: b :::l ,. d ~ :: ~ U e U t ~ 0 14 n e W 12 i a r 6 8.5 L g ~ "P. antiqua" 10 [rl I "I- , e ...--~------Members 1,2,3 '"Z I c en 8 :- W 8 - , , , n II: U I e 7.5- II: c ~ W

i • C

l Il. g; III 41 r u.. 7 I I, e 2 j- I d o II I: Ii n ::I: 6.5 J_I l- u Member 5) e 1 2 3 4 5 6 7 8 9 10 y a ~ 7 8 • 10 RELA TIVE AGE CLASSES w e LENGTH OF UPPER SECOND MOLAR (mm) t Figure 3 Histograms of age distributions hased on crown height a

G Figure 1 Mean and standard deviations of M2 lengths and M2 measurements in Ml and M2 of (a) 62 molars attributed to P. t e widths for' modern P. capensis (n = 23). fossil P. anliqua from antiqua and P. capensis; (b) 22 molars attributed to P. n i Members 1. 2 & 3 Swartkrans (n =' 29). fossii P. capenJis from Iransvaaiensis. Age distributions are expressed in relative age b a Member 5 Swartkrans (/'I ::::: 17). and P. Iransvaa[ensiJ from classes ranging from I (youngest individuals in assemblage) to 10 S

y Members 1 & 2 Swartkrans (n = 18) (the oldest individuals). b d e c u d o r p e R s. Afr.J. 7m\. 1994,29(1) 43

results confirm the observation that P. an/iqUlJ and P. to P. Iransvaalensis (Table 2) is significan~y larger than the capenJiJ are represented by higher frquencies of relatively mean value obtained for modem specimens of P. capensis (I young individuals. = 5,91; df. = 40; p = 0,05), However, the mean value for Figure 4 shows relationships betwccn crown height and modem specimens of P. capensis is not significantly hypocone-metacone distance (HMO) of upper molars (M' different from mean values for speci,mens attributed to P. and M') of P. antiqua, P. capensis, and P. lransvaalensis antiqua (I = 0,65; df. = 51; p = 0,05). Morphologically, all from Swankrans. These figures show linear relationships of the specimens from Swankrans attributed to P. GntiqUlJ, between the two variables, reOccting the fact that HMO as well as St 105 (a specimen used for reference purposes in increases with individual age. this study) arc indistinguishable from modem P. capensis. The following regression equation has been obtained for As shown in Figure 3, there is also a striking similarity the relationship between crown height (x) and the hypo­ between measurements of P. antiqua and P. capensis when cone-metacone distance (Y), for M' and M' of specimens crown height is examined in relation to hypocone-metacone previously identified as Procavia antiqua, from Members I, distance, a variable which Broom (1934) had previously 2 and 3 at Swartkrans: used to distinguish the two as distinct species. Clearly, HMO varies as a function of age. For HMO to be at all y = -D,549x + 5,670 (r =0,8; n = 36) Equation I. meaningful it should be analysed in conjunction with crown The following equation was obtained for modem specimens height. of P. capensis: Analyses of HMO and crown height in this study serve to show that there is no marked difference between the y = -D,564x + 6,669 (r 0,8; n = 74) Equation 2, relationship of these two variables in specimens attributed to The following equation was obtained for P. anliqua from P. antiqua and P. capensis. By contrast, there arc striking Members I, 2, 3 and P. capensis from Member 5 (combi­ differences when data for P. transvaalensis are compared ning samples): with measurements obtained from specimens attributed to P. capensis or P. an/iqUlJ. These results suggest that specimens y = -057Sx+ 5,821 (r = 0,8; n = 62) Equation 3. attributed to P. antiqUlJ and P. capensis could have belonged The relationship obtained for specimens of P. lransvaaiensi.'l to populations of the same species, distinct from those of P. from Members 1, 2, 3 and 5 is as follows: transvaalens;s. The range of variation in crown height of P. antiqua from y = -D,776x + 9,635 (r = 0,9; n = 23) Equation 4. Members I, 2 and 3 is about 7 mm (ranging between approximately 1 mm for the oldest individuals to 8 mm for DiScussion the youngest individuals at Swarikrans), corresponding . )

9 Morphologically, specimens from Swartkrans attributed to closely to the known range of variation in modem P. 0

0 P. transvaalensis arc indistinguishable from St 106, here capensis, The range in both P. an/iqUlJ and P. capensis is 2 used as a reference specimen for this species. A I test similar to the range of variation in crown height found in P. d e t indicates that the mean length of M' of specimens attributed IransvaaienJis (3-10 mm). The implication is that, by taking a d ( r e h s

i P. antiqua l

b LJ MEMBER 1

u 7 MEMBER 2

P E

e .§ o MEMBER 3 h UJ t

0 6 P. capensls y z b 6 MEMBER 5

..: t- d e !!! 5· P. transvaalensls t 0 [J n <:;' MEMBER 5 UJ <, 0 ~ + a

r Z g 0 4 - 0 e " " , 6 , c " ..: 'h 6 n t- + , e UJ ) c ::;; i , l

UJ r Z e 0 v d u + 0 0 2 - n 0 u Q.

y >- a :I: 1- w e t a G

0 2 4 6 8 10 t e CROWN HEIGHT (mm) n i b a Figure 4 Relationship between hypocone-metacone distance and crown height of M2, for P. anJiqua from Members J. 2 & 3 at S

y Swartkrans; P. capensis from Member 5; and P. tra,uyaalen.~js from Members 1,2,3 and 5. b d e c u d o r p e R 44 S.-Afr. Tydskr. Dierk. 1994,29(1)

relative age into account, P. Iranvaalensi.'i had tccth that ratio of 2 : 1. The relatively low proportion of females is of were more hypsodont than those of P. antiqun and P. interest in relation to age distribution data which show that capensis. this species is represented by a high proportion of adults. Hypsodonty has previously been associated with a Predation on adults of P. tran!ivaalensis in populations with specialization towards grazing (for example, equids and relatively low abundances of females could perhaps have a!celaphines are highly hypsodont and both are essentially meant that this species had a higher probability of extinction grazers). The case for P. transvaalensis being a hyracoid than the smaller Procavia antiqua I capen sis. that fed primarily on grass leaves would scem to be In all members, P. antiqun I capensis is relatively more supported by strontium:calcium ratios in bones of this abundant than P. transvaalensis. The ratio of P. antiqua I species from Swankrans (Sillen 1992). capen sis to P. transvaalensis (based on incisors) is circa Variability in dental morphology of extant hyracoid 6: I in Members I, 2 and 3 (combined sample), increasing genera has been studied by Hahn (1934) and Allaert., van to circa 8 : I in Member 5. This shift suggests an increase der Audenaerde & van Neer (1982). They noted that in relative abundance of P. capensis at a time when popula­ Procavia is more hypsodont than lIeterohyrax and tions of P. transvaalensis were in a state of decline. Dendrohyrax. This is not surprising considering the fact that Heterohyrax and Dendrohyrax arc browser.;. Of interest in Conclusions this study is the observation that the extinct species of It is concluded that P. antiqua and P. capensis are conspcd­ Procavia represented at Swartkrans, P. transvaalensis, is fic, the latter having precedence according to international distinctly more hypsodont than the extant species P. rules of zoological nomenclature. P. transvaalensis remains capensis. Earlier hyracoids such as Saghatherium from the a valid taxon, distinct from the smaller species at Swartkrans Oligocene are likely to have been browsers associated with which may now be referred to as P. capensis. wooded environments. Perhaps the hyposodont condition of The co-existence of two species of Procavia in the early P. lransvaalensis is a late specialization associated with the Pleistocene is under.;tandable in view of tbe observation that opening up of grassland environments during cool intervals P. transvaalenJis had hypsodont dentition, suggesting that it in the Pleistocene. fed primarily on grass leaves, a possibility supported by Differences in age distributions of Procavia at Swankrans slronlium:calcium ralios in bone samples of this species can be addressed in the context of questions concerning (Sillen 1992). Since P. capenJis is known today to be a cave taphonomy. The fact that P. transvaalensis is represen­ generalist, fceding on leaves of bushes, shrubs, tubers, roots ted by a relatively high proportion of adults is of interest in as well as grass (Skinner & Smithers 1991), the two species relation to the possibility that it was preyed upon primarily could have co-existed without undue competition for the by leopards. This warrants attention since such carnivores same resources. .

) will prey upon hyraxes if larger prey arc not available Analyses of fossil Bovidae from Swartkrans (Vrba 1975; 9

0 (Thackeray 1990; Pienaar 1969); in areas in the western Brain et al 1988) provide some indication of the nature of 0 Cape in the absence of large ungulate prey, P. capensis can 2

Pleistocene paleoenvironments at Swankrans. The bovid d form up to 80% of the vertebrates taken by leopards e faunal assemblages are generally dominated by alcelaphines t (Norton, Lawson, Henley & Avery 1986). If given a choice a which are indicative of open grassland conditions. Such d

( between two hyracoid species, one might expect leopards to environmenL, would certainly have suited P. transvaalensis r of the larger taxon, e have preyed preferentially on individuals if it fed primarily on grass foliage, as suggested here from h

s and of these they could have preferentially taken adults. i

l its hypsodont dentition.

b This could account for the high proportion of adults of the

u The extinction of P. transvaalensiJ toward. the end of the larger species, P. transvaalensis, in the early Pleistocene P Pleistocene, apparently some time around 11 000 years e deposits at Swankrans. In support of this suggestion, h

t B.P., is of interest since this is the period when large

mandible fragments of P. tranJvaalensis show the kind of y ungulates became extinct on a global scale (Martin 1%7; b damage that Brain (1981) has documented in a study of

d leopard feeding behaviour. Klein 1974). Remarkably it is the larger of the Procavia e t speeies, evidently a grazer, that beeame extinct. Of the n There is no reason to discount the possibility that leopards a ungulates that died out in southern Africa, it was grazer.; r may have preyed on P. capensis. However, was the species g

(many of them large) that became extinct at the end of the e also preyed upon by black eagles? This possibility deserves c Pleistocene (Klein 1974; Thackeray 1981). Perhaps P. ca­ n attention since it is known to be an important food item of e

c pensis, as a smaller hyracoid species and more of a genera­

i these large birds of prey (Ginn, Mcllleron & Millstein l list, had an adaptive advantage over P. transvaalensis. r \989), and at least some fossil hyrax from Swartkrans have e Age distributions and sex ratios obtained from this d damage of the kind known to be associated with the feeding n analysis of Hyracoidea at Swankrans are of potential value u behaviour of black eagles which open up the foramen

y magnum of hyrax crania with their talons (Brain 1981). for understanding the factors contributing to the extinction a of P. transvaalensis and the survival of the smaller species, w Black eagles may have found it easier to prey upon young e t rather than on older individuals of the small species; this P. capensis. Although it cannot be determined with certainty a

G could aceount for the relatively high proportion of young that the sex ratios of hyraxes represented in the fossil faunal t e individuals of P. antiqua I capensis represented at assemblages rencet the ratios in past populations, there is n i Swartkrans. some suggestion from the early Pleistocene samples that b a In the case of early Pleistocene samples from Members I, males outnumbered females by a ratio of 2 : 1. In the case S

y 2 and 3, males of P. transvaalcnsis outnumber females by a of P. transvaalensis, there is also evidence that adult indivi- b d e c u d o r p e R S. AfT. I. 1..001. 1994.29(1) 45

duals were sclected by one or more predators. Predation on Procavia capensis. S. Afr. 1. 7.001. 15: 745-780. adult individuals of P. transvaalensis from populations with FOURIE. LJ. 1983. The population dynamics of the rock hyrOJ< relatively lower frequencies of females could have contribu­ Procavia capensis (Pallas, 1766) in the Mountain 7..ebra ted to the decline and ultimate extinction of this species. National Park. Un pub), Ph,D. thesis, Rhodes University, In a recent study of mitochondrial DN A in modem Grahamstown. hyraxes from various regions of southern Africa. Prinsloo & GINN. PJ .. MCILLERON. W.G. & MILLSTEIN. P. LE S. 1989. Robinson (1992) found genetic differences between samples The complete book of Southern African birds. Struik of P. capensis. in the absence of morphological differences. Winchester, Cape Town. They suggested that two taxa diverged approximately 2 HAHN. H. 1934. Die Familie der Procavidae. Z. Saugelierk 9: million years ago. I""ving two relict populations: one in the 207-358. Karoo ncar Beaufon West and the other in Malawi. KLEIN, R.G. 1974. A provisional statement on the tennina1 Consequent migrations nonh and south arc believed to be Pleistocene mammalian extinctions in the Cape Biotic 7..one responsible for the extant populations of Procavia (Prinsloo (southern Cape Province, South Africa). S. Afr. Arch! Soc. & Robinson 1992). The present study has shown that at least Goodwin Series 2: 39-45. two species of Procavia did in fact exist in the Transvaal MARTIN. P.S. 1967. Pleistocene overkill. In: Pleistocene within the last 1.8 million years before the present. It would extinctions: the search for a cause, (eds) Manin. P.S. & be extremely interesting if analysis of DNA in fossils Wright. H.E. pp. 75-120. YaJe University Press, New Haven. previously attributed to P. antiqua and P. transvaalensis MEESTER.I.A.1 .. RAUTENAACH. I.L .. DIPPENAAR. NJ. & from Swankrans could shed funher light on genetic and BAKER, C.M. 1986. Classification of southern African morphological variability in Procavia. mammals. Transvaal Museum Monograph Np. 5. MlLLAR, R.P. 1971. Reproduction in the rock hyrax (Procavia ACknowledgements capensis). Zool. Afr. 6: 243-261. NORTON. P.M .. LAWSON. A.B .. HENLEY. S.R. & A VERY. We would like to thank C.K. Brain. E. de We~ E. Herold. I. G. 1986. Prey of leopards in four mountainous areas of the Plug. T. Robinson. and V. WaL~on for their help in this south-western Cape Province. S. Afr. J. WildJ. Res. 16: 47-52. study which was supported in pan by an FRD grant awarded PIENAAR. U. de V. 1969. Predator-prey relationships amongst to J.F.T. Two anonymous referees kindly provided helpful the larger mammals of the Kruger National Park. Koedoe 12: comments. 108-187. PRINSLOO. P. & ROBINSON. T.1. 1992. Geographic References mitochondriaJ DNA variation in the rock hyrax, Procavia ALLAERTS. W .. VAN DER AUDENAERDE. T. & VAN capen.fis. Mo/. BioI. £vol. 9: 447~56. .

) NEER. W. 1982. Dental morphology and the systematics of SHAW, 1.C.M. 1937. Evidence concerning a large fossil hyrax. 9

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Belg. 112: 217-225. SILLEN, A. 1992. Strontium·calcium ratios (SrlCa) of d

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