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VOLUME 69 PART 9 APRIL 1976 ISSN 0303-2515

OF THE SOUTH AFRICAN -
—

tae

CAPE TOWN

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.— Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
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Konn, A. J. 1960. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270.

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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 69 Band
April 1976 April
Part 9 Deel

£
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THE PLIOCENE FOSSIL OCCURRENCES IN
‘FE? QUARRY,
LANGEBAANWEG, SOUTH AFRICA

Q. B. HENDEY

Cape Town Kaapstad

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THE PLIOCENE FOSSIL OCCURRENCES IN ‘E’ QUARRY,
LANGEBAANWEG, SOUTH AFRICA

Q. B. HENDEY
South African Museum, Cape Town

(With 6 figures and 4 tables)

[MS. accepted 27 November 1975]

ABSTRACT

The Pliocene Varswater Formation in the vicinity of Langebaanweg, Cape Province, is
comprised of three main units, now named the Gravel, Quartzose Sand and Pelletal Phosphorite
Members. The lowest unit in the succession, the Gravel Member, has yielded a largely marine
fauna, including fifteen shark species, incorporated in rocky and sandy beach deposits. The
principal source of fossils, the Quartzose Sand Member, was laid down in a variety of
depositional environments in and near an estuary. The economically important unit, the
Pelletal Phosphorite Member, is fossiliferous in a relatively limited area which was situated in
the immediate vicinity of the river mouth. The fossils from the deposits overlying the Gravel
Member represent a wide variety of marine, freshwater and terrestrial invertebrates and
vertebrates. About seventy-five mammalian species are recorded, including a few belonging to
groups not previously recorded from Africa.

CONTENTS

PAGE
Introduction : : ; : 4 PALS)
Geology. 2 : ; : a) Alls)
Depositional environments . go = PDP
Fauna . , ; : : ; ; 231
Flora . 5 : ; 2 ; » 243
Dating . ‘ : : : : . 243
Conclusions : ; ; : . 245
Acknowledgements . : : . 245
References . F , ; : . 246

INTRODUCTION

Large-scale production of phosphate from deposits on the farm Langeberg
near Langebaanweg, Cape Province, was commenced in 1953 and several years
later the presence of fossils in these deposits was reported (Singer 1961). Initially
few fossils of good quality were collected, but prospecting revealed the presence
of highly fossiliferous deposits south of a small open-cast mine, ‘E’ Quarry.
During 1964 these deposits were exposed in a trench about 240 metres long,
120 metres wide and ranging in depth from 2 to 30 metres (Fig. 1). Soon after
the 1964 excavations were commenced it became apparent that the deposits
would yield well-preserved fossils in large quantities. Mining of ‘E’ Quarry (the
New Varswater Mine) has been continuous since then and although the number

215

Ann. S. Afr. Mus. 69 (9), 1976: 215-247, 6 figs, 4 tables,

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

of fossils recovered has varied from year to year, the original expectations have
been fulfilled. Certainly no other recorded African fossil occurrence of Pliocene
age has produced so large an assemblage of specimens representing so wide a
variety of species.

During 1968 the South African Museum commenced an investigation of the
fossiliferous deposits exposed in ‘E’ Quarry, this project following on from a
similar undertaking directed for a period of ten years by R. Singer of the
University of Chicago. The first phase of the Langebaanweg project had also
taken into account the mined-out occurrences at ‘C’ Quarry on the farm
Langeberg and Baard’s Quarry on the farm Muishondsfontein. The fossils from
these sites are limited in both quality and quantity and there are some still
unresolved problems relating to their geological associations.

Several publications have resulted from the second phase of the Langebaan-
weg project, including reviews (Hendey 1970a; 1973; 1974a), accounts of the
geology (Tankard 1974a; 19746; 1975), discussions on dating (Hendey 19706;
1972a; 19746), as well as descriptions of some of the fossils recovered (e.g.
Simpson 1971; Kensley 1972; Gentry 1974).

The geological study of the ‘E’ Quarry deposits, which has now been
largely completed, formed part of a broadly-based investigation of late Cenozoic
deposits in the south-western and southern Cape Province, and was undertaken
independently of the palaeontological study, although the two have been
mutually complementary.

Recently research on the fossils from Langebaanweg has decreased, although
there has been an increase in the amount of material collected. The latter
development is a direct result of changes in the mining programme. The expected
back-filling of ‘E’ Quarry (Hendey 1973) was commenced during 1974 and
collecting was accelerated in those areas to be covered by new mine dumps. In
addition, the mining company (Chemfos Limited) removed a large quantity of
fossiliferous deposit from one of the threatened areas (East Stream, see Fig. 1)
and this is being screened by the first permanent field assistants on site. The
nature of the present undertaking is being further influenced by the fact that
during 1975 mining of the last of the phosphatic deposits known to be highly
fossiliferous was commenced. Until recently these deposits were not scheduled
to be mined until about 1990.

Since its inception, the prime object of the present phase of the Langebaan-
weg project has been the recovery and identification of fossils, with the collecting
being as comprehensive as possible. The recent changes in the mining programme
have added urgency to this aspect of the undertaking since leisurely collecting
and excavation over certain areas of the mine are no longer possible and the
areas thus affected will increase with the passage of time. Many of the specimens
still in the deposits will be lost if they are not salvaged promptly. In addition,
the volume of fossiliferous deposit elsewhere is being steadily reduced by the
mining and although some is likely to remain indefinitely, technical difficulties
may make it inaccessible to further exploitation. The termination of active field

217

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA

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218 ANNALS OF THE SOUTH AFRICAN MUSEUM

work at the site in the not too distant future has now become a distinct
possibility.

The collecting of fossils could be expedited by being more selective, but as
far as possible this approach 1s avoided, mainly because it would adversely affect
that part of the undertaking concerned with the recording of the nature of the
fossil occurrences. This aspect of the investigation already suffers because of its
lower priority rating. It is hoped that eventually an analysis of specimens from
any given area in terms of species and body part representation, together with
their condition and associations, will provide information on environments of
deposition, the taphonomy of the fossils and aspects of the ecology of the area
at the time of deposition. Some information of this nature is already available.

Although more time has recently been spent on the collecting of fossils and
directly related technical matters, some progress has been made in the
palaeontological research, and the present report updates some of the
information and opinions previously recorded.

GEOLOGY

The phosphatic and fossiliferous deposits exposed in ‘E’ Quarry make up
what is now termed the Varswater Formation (Hendey 1974a; Tankard 1975).
This formation is Pliocene in age and it is underlain by the Miocene Saldanha
Formation (Tankard in press). The overlying deposits, informally termed the
‘surface bed’, are largely, or entirely Pleistocene and Holocene in age. The
Varswater Formation is comprised of three main units which were referred to
by a variety of names in earlier publications. The member names used in the
present report (Table 1) are those which are now considered most appropriate
(A. J. Tankard pers. comm.), while those of the beds are purely informal and
likely to be modified at a later date. The Gravel and Pelletal Phosphorite
Members are both unequivocally described by their names, but the name of the
Quartzose Sand Member refers to the dominant lithological element. This
member also includes horizons of carbonaceous sand and clay (the ‘peat bed’,
Fig. 1), clayey sands and silt. The non-geographic names of the members are
justified in terms of Section 3.10(d) of the South African Code of Stratigraphic
Terminology and Nomenclature (1971: 118).

Since Tankard (1975) has described the Varswater Formation in detail, the
only other comments on the geology of ‘E’ Quarry which are included here are
those which have a bearing on the palaeontology of the deposits.

The Quartzose Sand Member is the most highly fossiliferous of the three
units, while the Pelletal Phosphorite Member is poorly fossiliferous, except for
an area still exposed of the west wall of the mine (‘bed 3a’, Fig. 1). There are
only limited exposures of the Gravel Member in a few areas of the mine and
relatively little attention has been paid to this unit during the current phase of
the Langebaanweg project.

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 219

TABLE |

Stratigraphy of the ‘E’ Quarry exposures of the Varswater Formation

Over most of ‘E’ Quarry the three members of the Varswater Formation
are readily identifiable. The Gravel Member is always unmistakable, although
the sandy element of this member may on occasion have been regarded as part
of the overlying Quartzose Sand Member. Recognition of the Pelletal Phospho-
rite Member has, for the most part, not been difficult, although its lower limit
has not always been clearly defined. The problems encountered in recognizing
the lower limit of this member have been due to a variety of factors. Theoretically
the mining is cut off at the base of the Pelletal Phosphorite Member, although in
practice this is impossible and irregularities on the floor of the mine do not
necessarily reflect the nature of the boundary between the Pelletal Phosphorite
and Quartzose Sand Members. In most areas the mining has actually extended
into the latter unit where the exposed surface may be contaminated and obscured
by spillage from the excavator and slumping of deposit from the vertical mine
faces. Furthermore, the Quartzose Sand Member was truncated prior to the
deposition of the Pelletal Phosphorite and where it is very thin it has been
difficult or impossible to recognize. In certain critical areas the deposits of the
two members are superficially similar and although detailed sediment analyses
would no doubt resolve the issue in problematical instances, no such analyses
have been undertaken.

From the preceding comments it should be clear that difficulties have centred
largely on the identification of the upper and lower limits of the Quartzose Sand
Member. Added to this is the fact that this member is the most complicated unit
in the succession in terms of lithology and the variable character of the deposits
has itself led to some confusion in the past. The sometimes striking differences

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

The stratigraphy of the East Stream and peat areas of ‘E’ Quarry

in the nature of the Quartzose Sand Member are illustrated by examples of the
‘E’ Quarry succession given in Table 2.

The difficulty experienced in identifying the units of the Varswater Forma-
tion has resulted in some of the fossils collected being of doubtful provenance.
In this connection it should also be noted that before the basic three-unit
succession was recognized in 1969, little or no data on the source of specimens
was recorded and the provenance of many specimens collected before that date
may never be known. The same often applies in the case of specimens picked up
by mine workers. Although the problem of unprovenanced material is not as
serious as it might have been, it is an unfortunate complication in the
palaeontological investigation.

In the faunal lists given later, records of unknown or uncertain origin are
excluded. The source of most of the more significant specimens from ‘E’ Quarry
is well documented and all the species identified to date are represented by at
least one specimen of known provenance.

The characteristics of the units comprising the Varswater Formation are
accounted for by the sequence of events which occurred during deposition of the
formation. These events have a direct bearing on the interpretation and identifi-
cation of depositional environments (vide infra). They were summarized by
Tankard (1974a: 219) who stated that, ‘In the Pliocene a transgressing sea
pushed deltaic marsh sediments ahead of it until it reached a temporary still-
stand . . . [with the] temporarily stable conditions [allowing] a barrier bar to
build up, behind which estuarine conditions prevailed. The estuary was fed by a
river from the north-east. The final transgression reworked the older sediments.’
It was at the time of the stillstand that the Quartzose Sand Member was
accumulated, while the Pelletal Phosphorite Member was laid down during the
final transgression (Fig. 2).

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Although the geology of the Varswater Formation is now well known,
there is still scope for further detailed work. Of particular interest to the
palaeontological investigation is geological evidence indicating depositional
environments. The relationships between fossils and the deposits in which they
occur has hitherto received little attention.

DEPOSITIONAL ENVIRONMENTS

It is now generally accepted that the Varswater Formation accumulated
during a marine transgression at a time when a river discharged into the sea in
the immediate vicinity of “E’ Quarry. It follows that marine, fluvial and terrestrial
environments were then present in the area, together with some of the attendant
micro-environments peculiar to each. Theoretically it is therefore possible that
deposition of this formation took place in more than one of these environments
and also that as the sea transgressed macro- and/or micro-environmental
changes might have occurred in any given area now exposed in the mine.

Tankard (1975) has given an account of the depositional environments of
the three units comprising the Varswater Formation in ‘E’ Quarry. They were
as follows:

Gravel Member— Rocky and sandy marine beach environments.

Quartzose Sand Member—Essentially estuarine and fluvial environments,
although a peat deposit, probably representing a marsh environment,
is also mentioned.

Pelletal Phosphorite Member—A_ shallow-water environment situated

between a beach bar and beach.

The means by which the vertebrate fossils in deposits overlying the Gravel
Member came to be incorporated in these deposits has been the subject of some
dispute in the past (Tankard 1975: 281). There is in fact no single answer to this
question and it is clear that just as the depositional environments of the sediments
varied, so too did those of the fossils. The characteristics of the deposits change
both vertically and horizontally, the most noticeable and frequent changes being
in the lower levels of the succession, that is, those levels in which fossils occur
most commonly. Although no detailed study of the relationships between
sediments and fossils has yet been undertaken, it is obvious that differences in
the deposits go together with palaeontological differences.

Some observations on the environments in which the ‘E’ Quarry fossils were
laid down have already been recorded (Hendey 1974a), but it is probably worth
while at this stage to elaborate on earlier statements.

The Gravel Member

The fauna of the Gravel Member is comprised overwhelmingly of marine
species, both invertebrates and vertebrates, and the fossils of this member were
undoubtedly accumulated along a marine shoreline. The invertebrates indicate

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 223

the presence of both rocky and sandy habitats. Remains of terrestrial vertebrates
do occur, but they are invariably fragmented and rolled. This also applies to
many of the marine fossils. The condition of the fossils is evidently the result of
wave action and it differs from that of fossils from fluvial deposits in the
Quartzose Sand and Pelletal Phosphor te Members.

The Quartzose Sand Member

The situation in respect of Quartzose Sand Member depositional environ-
ments is complicated. Tankard (1975) has recognized both estuarine and fluvial
facies within this member and many of the fossils recovered from the ‘E’ Quarry
exposures must have been deposited in the river and estuary. For example, some
of the fossiliferous exposures in the eastern parts of the mine are medium to
coarse sands which are a westerly extension of the fluvial deposits referred to by
Tankard (1975: 274). On the other hand, the nature of some fossil occurrences
suggests that specimens were deposited in areas adjacent to the river and estuary
in both subaerial and subaqueous situations (e.g. floodplain and pond).

Evidence suggesting that certain of the Quartzose Sand Member fossils
were accumulated on land surfaces has been mentioned elsewhere (Hendey
1974a: 349-353). Since this discussion related to carnivore activity rather than
environments of deposition, some points concerning the latter were omitted or
insufficiently emphasized.

In certain of the Quartzose Sand Member exposures (e.g. the floodplain
deposits of East Stream) the condition of the fossils and the nature of their
occurrence contrasts with the situation where there is incontrovertible evidence
for subaqueous deposition of material (e.g. in the fluvial deposits of the Pelletal
Phosphorite Member). The fossils in the latter deposits tend to be abraded and
fragmented, while elements of single skeletons are dispersed. Exceptions to these
rules were probably specimens which had been protected by soft tissues or which
had not been subjected to prolonged transport. The fossils of subaerially
accumulated assemblages are generally perfectly preserved and damage to, or
dispersal of, specimens can usually be ascribed to carnivore activity or fires.

The fact that there are certain Quartzose Sand Member fossil occurrences
where only terrestrial species are recorded, or where they are much more
commonly represented than aquatic species, also tends to suggest that there was
subaerial accumulation of specimens. The presence of some aquatic species
could be explained by periodic inundations of land surfaces. The persistent
presence of an aquatic environment would lead to higher proportions of
aquatic species being represented and there are occurrences in the Quartzose
Sand Member where this is the case. In one such occurrence in the East Stream
area a relatively high proportion of fish bones went together with appreciable
quantities of abraded bone. These fossils were probably deposited in a channel,
a feature for which there was no other obvious evidence.

An example of burnt bone having suggested that a skeleton cannot have
been moved after it was partially burnt was given elsewhere (Hendey 1974a:

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

351). The nature of this particular occurrence was the important factor, since
burnt bone itself is not necessarily proof that there were fires over areas
presently exposed in the mine. Burnt bone might easily have been washed in
from elsewhere. There is another recently discovered example of burnt bone
which must have been in situ. This was a concentration of several hundred bones
of small vertebrates, mainly rodents and insectivores, most of which are heavily
charred. This occurrence is likely to represent the residue of a burnt owl pellet
accumulation, which cannot have been moved after burning without the bones
becoming dispersed.

A hitherto unrecorded factor which supports the theory that some subaerial
accumulation of fossils occurred concerns the presence of coprolites in the
deposits. Coprolites of at least three types have been recovered from the
Quartzose Sand Member. The first and largest type were evidently produced by
large carnivores, probably hyaenas. The second type are smaller and contain
fragmented bones belonging to small vertebrates. They were probably produced
by one or more of the smaller carnivores recorded from this member. An
account of such coprolites was recently given by Mellett (1974). The last type are
small, with no visible bone and tending to be cylindrical when not deformed.
Their source is not known but they are extremely abundant in certain areas
(e.g. East Stream).

Coprolites have been recovered only in certain parts of the mine and, except
for the smallest kind, they are nowhere common. Their condition varies, some
being remarkably fresh in appearance, while others are fragmented and distorted.
One of the ?hyaena coprolites, which was found together with four others, is
flattened, its appearance suggesting that it was trampled when fresh. Some of the
smallest coprolites, which are very fine-textured, show clear impressions made
by leaves, although they may be otherwise undistorted.

In order for faeces to be preserved intact they must almost certainly have
been dropped on a land surface and have been fairly rapidly buried thereafter.
Fresh faeces dropped in and then transported by water is unlikely to have
survived intact for long. It is also unlikely that groups of specimens would have
remained together if they had been transported and there are four instances
recorded where groups of 3, 3, 5 and 7 ?hyaena coprolites were found in close
association. These groups, together with several isolated specimens, came from
a relatively limited area in the eastern part of the mine. Unexcreted faeces could
have been carried to its final resting place while still inside a carcass, but none
of the coprolites has been found in direct association with other remains of their
possible producer. In addition, the shape of the better preserved specimens, and
the presence of leaf impressions on some, indicates that they had actually been
excreted.

Taken in conjunction the various factors referred to above are here regarded
as convincing evidence for subaerial accumulation of at least part of the
Quartzose Sand Member fossil assemblage.

The earlier reference to periodic inundations of land surfaces was intended

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 225

to indicate seasonal flooding of the river and estuary, but there is also evidence
for tidal flooding. A recently exposed deposit in the south-western part of the
mine includes an invertebrate fauna which indicates that the depositional
environment was a tidal mud flat. The nature of the deposits is in keeping with
the fauna which it contains. The limited exposures of this deposit examined prior
to their becoming temporarily inaccessible suggested that it overlies the peat bed
mentioned earlier (see Table 2).

The faunas of the peat and tidal mud flat beds are strikingly different from
one another and both differ from the fauna of more typical exposures of the
Quartzose Sand Member elsewhere. The mud flat bed is an exceptional
occurrence in this member, being the only one from which large numbers
of well-preserved invertebrate fossils have been recovered. Vertebrate remains
are rare. By contrast vertebrate fossils occur commonly in the peat bed
and in certain other exposures of the Quartzose Sand Member. As fossil-
collecting in this member has progressed, it has become apparent that while
certain vertebrate species are fairly ubiquitous, others occur only in certain
areas, while the overall representation of species and the condition of specimens
varies from place to place. While it may not yet be possible to interpret all such
evidence meaningfully, a superficial comparison of the peat bed and East Stream
faunas should serve to illustrate that different sediments go together with faunal
differences.

In this instance the sediment differences are visually striking, the black
sands and clays of the peat bed contrasting sharply with the white quartzose
sands of the East Stream area. The deposits in the latter area are here regarded
as a largely floodplain accumulation, while the peat area is thought to have been
a marsh.

The fossils of the peat bed tend to be less complete than those from East
Stream, although this may in part, or even largely, be due to the recent disturb-
ance of the peat by mining activities. This disturbance may also account for the
fact that whereas at East Stream a number of instances are recorded where
partial skeletons of individuals were preserved, nothing on a comparable scale
has so far been observed in the peat bed. A notable exception was the discovery
of the distal extremities of a sivathere fore- and hindlimb, elements of which
were found in articulation standing more or less vertically in the deposit,
seemingly all that survived of an animal trapped in the marshy deposits. At East
Stream elements of single skeletons were, with few exceptions, found slightly
dissociated from one another and they tended to lie more or less horizontally in
the deposits. This suggests that they were accumulated on a firm surface.

Although a single species of land tortoise (Chersina sp.) 1s the most
commonly represented vertebrate in both areas, there are otherwise some
marked differences in the representation of species. There are many species
recorded from East Stream which are not known from the peat bed, although
the reverse either does not apply or is at least much less obvious. Another
example concerns the pig, Nyanzachoerus, which is known from the peat bed on

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

the basis of only a few isolated teeth and bones, whereas at East Stream the
remains of at least fifteen individuals, some represented by incomplete skeletons,
have been collected. Birds, which are common in both areas, provide another
example. In the East Stream assemblage the most common species 1s a francolin,
which is a terrestrial bird, but in the peat bed it is rare, while waterbirds are
relatively more common (G. Avery pers. comm.).

Another rather curious difference between the two faunas concerns the
representation of the seal, Prionodelphis capensis. This species is not common in
either of the faunas, but at East Stream it is represented almost exclusively by the
remains of very young individuals, whereas only a few isolated teeth and bones
of adult seals are known from the peat bed. There are also differences in the
coprolite occurrences in the two areas. East Stream is one of the areas in the
mine where the smallest type of coprolite occurs in great numbers, while the
larger type with visible bone and those of ?hyaenas are rare. The only coprolites
known from the peat bed area few specimens of the type which contains visible bone.

A complete analysis of the two assemblages will no doubt provide further
and more precise examples of their similarities and differences, which are
presumably more than just fortuitous.

The potential importance of faunal analysis in determining depositional
environments is illustrated by the fact that there are obvious differences in
faunal assemblages even where deposits are superficially little different or
indistinguishable. Such differences have been observed at the same level in
apparently homogeneous deposits over distances of only a few metres. An
example mentioned earlier was the occurrence in the East Stream area of a
‘channel’ containing a relatively high proportion of fish remains and abraded
bones. A second example concerns a quartzose sand exposure in the vicinity of
the peat bed which was found on screening to contain large numbers of frog
bones, but very few terrestrial vertebrate fossils. In similar deposits elsewhere the
representation of fossils was reversed, with frogs being rare and terrestrial
vertebrates common. The presence of a pond, which left no other obvious
traces, could account for the amphibian-rich occurrence.

Although the question of depositional environments within the Quartzose
Sand Member has received only passing attention during the current phase of
the palaeontological investigation, there is evidence for fossils having accumu-
lated in estuarine, fluvial, marsh, mud flat, pond and floodplain environments.

The Pelletal Phosphorite Member

The Pelletal Phosphorite Member covers a wide area and is much the
thickest of the units in the Varswater Formation, but it is known to include
vertebrate fossils in large numbers only in a relatively limited area, exposures of
which still exist along the more northerly part of the west wall of the mine. These
fossiliferous deposits are informally termed ‘bed 3a’ and reasonably large fossil
samples have been recovered from two exposures, designated ‘bed 3aS’ and
‘bed 3aN’ (Fig. 3).

DOG),

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA

puryog ‘Jay pomodie) Seg poq pue (JY4sII pamolie) Nee paq jo

*(soor}
soinsodxs Surmoys ‘(¢L6] IsNsNYW) AlIeNYD .q, 0} UOISUd}xe A]IO}SOM MOU SUL “€ “SILT

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

The bed 3aS deposits are exposed along the lower mining face (‘bottom cut’)
of the west wall, while the bed 3aN deposits are approximately 100 metres
north-north-west on the upper mining face (‘top cut’) at elevations of between
2 and 4 metres higher. The bed 3aN deposits are close to the northerly limit of
economically recoverable phosphate and are only between 2 and 3 metres thick,
of which only the lowest 0,5 to 1 metre is fossiliferous. Some fossils have been
recovered from the uppermost levels of the Pelletal Phosphorite in this area, but
they are not regarded as part of the bed 3aN sample. The bed 3aS deposits are
also between 2 and 3 metres thick in an area sampled by controlled excavations
during 1969 and 1970. In this area fossils were concentrated in three distinct
levels spread over the lowest 1,5 metres of deposit. Bed 3aS deepens in a southerly
direction, where there may be more than three levels of concentration. The
thickness of both beds 3aS and 3aN is considerably less than the 25 metre
maximum development of the Pelletal Phosphorite Member.

In the bed 3aN area the base of this member is marked by a 0,75 metre thick
phosphatic sandstone. The surface of the rock at its most northerly exposure
appears to be fairly smooth and more or less horizontal, while the overlying
deposits are apparently not fossiliferous. A little further south the rock surface
dips markedly to the south-west and it becomes progressively more irregular.
Crevices and potholes filled with coarse sand and gravel are common and the
rock eventually becomes discontinuous. Fossils occur in abundance where the
rock surface is irregular, being concentrated in the irregularities and becoming
progressively less common upwards in the overlying finer-grained sediments.
The latter have a clay component in the more northerly exposures, but this is
absent in the southerly exposures where the deposits which immediately overly
the rock are unconsolidated sands. The ill-defined boundary between these two
types of deposit runs from north-east to south-west. Indications are that there
was a channel of fast-flowing water directed in a south-westerly direction in that
area where the rock surface is irregular and where the overlying deposits are
unconsolidated sands. The clayey-sands apparently formed the northern bank
of the channel.

The nature of the bed 3aS fossiliferous occurrences is essentially similar,
except that in this instance there is no rock horizon at the base of the deposits,
there are no clayey deposits indicating a channel bank and fossils are concen-
trated at more than one level in the deposits. The exact stratigraphic relationship
between bed 3aS and bed 3aN is not known, but the lower elevation and more
southerly situation of the former suggests that it was laid down earlier during the
marine transgression than bed 3aN. There may, however, be a direct link
between the highest of the bed 3aS levels of fossil concentration and bed 3aN.

Indications are that bed 3a was laid down in, or in the direct path of the
river which discharged into the sea most of the sediment making up the Pelletal
Phosphorite Member and that the course of the river shifted northwards as the
sea transgressed. A structure contour map of the base of the Pelletal Phosphorite
Member (Tankard 1975: fig. 3), a modified version of which is reproduced here

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 229

# U iT ‘Ss Ak Q P 0 _(N
|

(i
1
ra oe lege
' 33
yet | 40
_! 38 iE
a sal - Na
ee [4
: 6
h
0 300

“9 es nee eee) contour interval 2m
metres
10

Fig. 4. Structure contour map of the base of the Pelletal Phosphorite Member. Arrowed lines
indicate the probable courses of the river at the time that beds 3aS and 3aN were laid down.
(Adapted from Tankard 1975: Fig. 3.)

(Fig. 4), supports the suggestion that the lower course of the river was directed
towards the area where bed 3a is now exposed.

The nature of the bed 3a fossil occurrences, and particularly those of bed
3aN, provide clear evidence for deposition by fast-flowing water. Specimens
trapped by irregularities in the rock surface had protruding parts either com-
pletely abraded away or broken up and the fragments scattered in a south-
westerly direction. In many instances specimens on the rock surface had their
lower parts abraded, apparently by the coarser sediment fraction carried along
the rock surface by the flowing water. Individual elements of single skeletons
were dispersed and in some instances were traced by following connected series
of irregularities in the rock surface.

A feature of the faunas of both beds 3aS and 3aN was the large number of
specimens of the seal, Prionodelphis capensis, which are represented. Several well-
preserved skulls of this species were recovered from bed 3aN, while the several
skulls of the similarly-sized hyaena, Hyaena abronia, found in the same deposits
were not as well preserved or as complete. The more numerously represented
and better preserved remains of the aquatic carnivore suggests that deposition

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

was in a subaqueous environment and that the remains of the terrestrial species
had suffered transport over longer distances. As a general rule the terrestrial
fossils from bed 3a are more fragmented and less well preserved than similar
specimens from some of the Quartzose Sand Member deposits.

While most of the bed 3a fossils probably were deposited subaqueously, the
possibility cannot be ruled out that some subaerial accumulation of material
also took place (e.g. on the river banks and/or sand bars). A ?hyaena coprolite
was recovered from the clayey deposits of bed 3aN, that is, those deposits
regarded as having formed the northern bank of the channel at the time that the
fossils of bed 3aN were being accumulated. The smallest type of coprolite is not
uncommon in bed 3aS, but the majority of these specimens are fragmented, their
condition being in marked contrast to the generally very well preserved specimens
from the Quartzose Sand Member.

The previously stated opinion that some of the fossils from what is now
termed bed 3aS were derived from the Quartzose Sand Member (Hendey 19708:
122), may be relevant to the question of the bed 3aS coprolites. These specimens
could well be part of the derived element of the bed 3aS assemblage, having
hardened sufficiently while incorporated in the Quartzose Sand Member to
survive transport to, and redeposition in the Pelletal Phosphorite Member. On
the other hand, they were in most cases not hard enough to survive the trans-
portation intact.

While there is a strong likelihood that the bed 3aS deposits include fossils
derived from the Quartzose Sand Member, there is no evidence to suggest that
this was the case with bed 3aN. Truncation of the Quartzose Sand Member after
the 30 metre stillstand must have been confined to the early stages of the final
transgression, that is, the time when the lower levels of bed 3aS were being laid
down. The great majority of the bed 3aS coprolites come from the lower levels.
The relatively high elevation of bed 3aN virtually precludes the possibility of it
containing fossils derived from the Quartzose Sand Member.

Occasional fossils have been recovered from exposures of the Pelletal
Phosphorite Member other than beds 3aS and 3aN. Some specimens have been
collected in the west wall area from deposits overlying bed 3a, including a few
from the uppermost level. These specimens were probably also transported into
the area by the river. Their rarity may be due to the river mouth having been
some distance away when these deposits were laid down, the fossils representing
the remnants of occasional carcasses which had floated out to sea from the river
mouth. Specimens have also been collected from several different levels in the
most north-easterly exposures of the Pelletal Phosphorite Member. These
fossils are probably the remains of animals accumulated near the southern side
of the river mouth, while those of bed 3a accumulated in, ahead of and on the
northern bank of the river mouth (Fig. 4).

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 23 il

FAUNA

During the past six years large numbers of fossils have been collected from
‘E’ Quarry, most material having come from surface collecting, excavations at
random and screening of deposit, but some also having been recovered in
controlled excavations. The deposits sampled extend over an area of about
28 hectares (70 acres), with specimens having come from all the known fossili-
ferous horizons within the Varswater Formation. There is considerable variation
in the size of assemblages from individual occurrences within the deposits, while
the total assemblages from each of the three members are also of unequal size.
The fauna of the Quartzose Sand Member is the largest and best known, since it
has been exposures of this member which have been the principal focus of
attention during the current phase of the Langebaanweg project (Fig. 5). Mining
of highly fossiliferous Pelletal Phosphorite Member deposits (bed 3a) was
recommenced during 1975 so that there has recently been a substantial increase
in the amount of material collected from this member. Relatively little time has
been devoted to the collecting of fossils from the Gravel Member, exposures of
which are limited.

In terms of the requirements for taxonomic studies, many species, mainly
amongst the smaller vertebrates, are more than adequately represented in
existing collections and in such instances the addition of further material may
be of little or no significance. There are, however, many more species which are
poorly represented and in these instances there is still a real need to build up
sample sizes. Since 1969 the annual additions to the collections have always
included several new records for the site, while there has also been further
identification of material already in the collections. The ‘E’ Quarry fauna is, in
general, comparatively well known, although most of the specimens have still
to be studied in detail.

The following summary accounts of groups represented in the fauna include
references to recent new records, recent and current studies, significant additions
to previously existing species assemblages, as well as other comments on avail-
able material.

Invertebrates

Many of the invertebrate fossils recovered from ‘E’ Quarry have already
been described (Kensley 1972), this material having come from the Gravel
Member, while Tankard (1975) has mentioned other invertebrates from the
Varswater Formation. Additional material is now available, including the first
substantial invertebrate assemblage from the Quartzose Sand Member. This
material is from the tidal mud flat deposit mentioned earlier and includes
marine, freshwater and terrestrial molluscs. Many of the specimens are remark-
ably well preserved, some even retaining traces of their original colour. This
material is being studied by B. Kensley (South African Museum) and P. Nuttall
(British Museum (Natural History)).

ANNALS OF THE SOUTH AFRICAN MUSEUM

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PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 233

Lower vertebrates

No progress has been made in the identification of bony fish, amphibians
and reptiles, but P. A. Hulley (South African Museum) has completed a pre-
liminary investigation of the cartilaginous fish from the Gravel Member
(Table 3). Of particular interest was the fact that the Selachii proved more
diverse than anticipated, with the assemblage being complicated by the presence
of derived specimens (see p. 243).

TABLE 3

A provisional list of the cartilaginous fish from the Gravel Member of the Varswater
Formation, including derived material. (Identified by P. A. Hulley of the South African
Museum.)

SELACHII

Hexanchidae
Notidanus serratissimus
Carcharhinidae
Carcharhinus melanopterus
Carcharhinus limbatus
Galaeorhinus sp.
Prionace glauca
Negaprion sp. or Hypoprion sp.
Odontaspidae
Odontaspis accutissima
Odontaspis sp. B
Odontaspis sp. C
Otodontidae
Megaselachus megalodon
Carcharodontidae
Carcharodon sp.
Isuridae
Tsurus sp.
Squalidae
Squalus sp.
Squatinidae
Squatina africana
Squatina sp.
BATOIDEI
Rajidae
Raja sp.
Trygonidae
Gen. & sp. indet.
Myliobatidae
Myliobatis sp.

Birds

Although the ‘E’ Quarry birds probably constitute the largest late Tertiary
avian assemblage from anywhere in Africa, they have received only superficial
attention, with only one species, a penguin, having been positively identified
(Simpson 1971). The original study of the penguin remains suggested the
presence of a second species and this has now been confirmed by more recently
discovered material, although the second species remains unidentified (Simpson

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

1975). G. Avery (South African Museum) has identified to the family or genus
level about a dozen other birds, but there are many more which are completely
unclassified. The Quartzose Sand Member has been the source of the largest
number and the best preserved of the specimens. New material includes
incomplete skeletons of two large raptors and another belonging to a stork-like
species. The tidal mud flat deposits of the Quartzose Sand Member contain
fragments of bird egg-shell, some of which retain their colour.

Mammals

The mammalian fossils from the ‘E’ Quarry exposures of the Quartzose
Sand and Pelletal Phosphorite Members have been the principal focus of the
current phase of the Langebaanweg project. The number of species recorded
from these deposits has grown steadily over the years and approximately
seventy-five have now been positively or tentatively identified (Table 4). Only
about one-third have been described and even these include a number which are
incompletely classified, largely because of inadequacies in available material. In
some instances newly discovered material has made positive identifications
distinctly possible.

Mammalian microfauna

The updated list of “E’ Quarry mammals differs most markedly from those
previously published by including provisional identifications of many of the
small mammals from the Quartzose Sand Member. This part of the list was
provided by T. N. Pocock (Vanderbijlpark, Transvaal). Of interest is the first
record of a bat from the site. The small mammals of the Pelletal Phosphorite
Member, which are generally represented by more fragmentary material, remain
unstudied. The rodents are to be studied by Craig C. Black (Carnegie Museum,
Pittsburgh).

Primates

Although a cercopithecoid is tentatively recorded, there has been increasing
doubt about its presence in view of the continued lack of positively identifiable
material. Primates feature prominently in the late Cenozoic fossil record of
Africa and their great rarity in, or complete absence from the Varswater
Formation is a perplexing aspect of its fauna. A feature of this fauna is its great
diversity and the situation which exists in respect of so successful a group as the
primates is indeed curious. The situation of Langebaanweg in a coastal environ-
ment at the southern continental extremity is likely to be related to the rarity or
absence of this group. The Quaternary fossil record of the south-western Cape
Province is characterized by a similar dearth of primates.

Carnivores
In contrast with the primates, carnivores are an exceptionally well repre-

sented and diverse group. Fully one third of the Varswater Formation mammals
identified to date are carnivores.

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 235

TABLE 4

A provisional list of the mammals from the Quartzose Sand and Pelletal Phosphorite Members
of the Varswater Formation.

Quartzose Pelletal
Sand Phosphorite
Member Member
INSECTIVORA
Chrysochloridae
Chrysochloris sp. . : ; , A : : : < s<
Soricidae
Myosorex sp. 3 ‘ F : ; : ; ; x
Suncus sp. : : : ; : i x
Soricidae gen. & sp(p). indet. ; : 5 : : x
Macroscelididae
Elephantulus sp... ? ; F : : ; ; < x
CHIROPTERA
Vespertilionidae
Eptesicus sp. . : ‘ ‘ ; ; ; : ; x
? PRIMATES
? Cercopithecidae
Gen. & sp. indet. . : : ‘ : ; A : Xx
PHOLIDOTA
Manis sp. ; : : : ; ; ; Z . Xx
TUBULIDENTATA
Orycteropus sp. : 2 : : F é : ‘ x x
CARNIVORA
Canidae
Vulpes sp. : eee a ST eas x
Ursidae
Agriotherium africanum 3 7 ; : 3 i Xx
? Procyonidae
Gen. & sp. indet. . : 3 ! ; ; : : x
Mustelidae
Mellivora aff. punjabiensis . : x
Mellivorinae gen. & sp. indet. (af. Plesiogulo) ; x
Enhydriodon africanus . : f : x
Viverridae
Viverra leakeyi ~< x
Viverrinae gen. & sp. indet.
Genetta sp. x
Herpestes sp. A x x
Herpestes sp. B x x
Herpestinae sp. C . x
Herpestinae sp. D . x
Herpestinae sp. E . x
Hyaenidae
Percrocuta australis ; <
Hyaenictis preforfex . : ; : : ; : x
Euryboas sp. noy. . x
Hyaena abronia » <
Hyaenidae sp. B x<

Hyaenidae sp.E . : : : : ; ; : ><

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Quartzose Pelletal
Sand Phosphorite
Member Member
CARNIVORA (cont.)
Felidae
Machairodus sp. x
Homotherium sp. u Xx
Felis sp. (small) x
Felis aff. issiodorensis x x
Felis obscura : x
Dinofelis diastemata Z : 5 : : : x x
Carnivora (possibly Lutrinae) gen. & sp. indet. : : 5 x<
PINNIPEDIA
Phocidae
Prionodelphis capensis . : : : : : : x 4
PROBOSCIDEA
Gomphotheriidae
Gen. & sp. indet. . : ¢ : ; ; : ; x S<
Elephantidae
Mammuthus subplanifrons . : : : : : < ?
HYRACOIDEA
Procavia cf. antiqua : , : : , ; ‘ x ?
PERISSODACTYLA
Equidae
Hipparion sp. A. ‘ : ; ; : : : x
Hipparionsp.B . ; , : : : : : Xx
Hipparion namaquense . : : 3 : : : x
Rhinocerotidae
Ceratotherium praecox . : : F 5 : é < ?
ARTIODACTYLA
Tayassuidae
Gen. & sp. indet. . : ; : : ‘ : 3 x
Suidae
Nyanzachoerus sp(p). . : j : F ‘ : x x
Hippopotamidae
Hippopotamus sp. . : : : E : : : x
Giraffidae
Sivatherium sp. ; ; : : , : ; ; x <
Giraffa sp. : P : ; j : : : : < x
Bobidae
Tragelaphus aff. angasi . , : : : 2 : x x
Mesembriportax acrae . : ; : : é , x x
Bovini gen. & sp. indet. : , : : ; ; x x
Reduncinisp. A. 3 ; : : : ‘ : x
Reduncinisp.B. 5 : ; i : ; : x
Alcelaphini sp. A . : : : : ; : - x
Alcelaphini sp.B . : : : : : : 3 x
Raphicerus sp. : : : . : : : : x x
Gazella aff. vanhoepeni . : : : ; P : x
?2Ovibovini gen. & sp. indet. s : ; ; ; x
LAGOMORPHA

Gen. & sp. indet. . ; : 2 ‘ 3 ; F x

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA DSi);

Quartzose Pelletal
Sand Phosphorite
Member Member
RODENTIA
Bathyergidae
Bathyergus sp. 2 ' : ‘ : : : : x x
Cryptomys sp. : ‘ : é ‘ : : : x
Hystricidae
Gen. & sp. indet. . : : 4 ; ; : ? x
Muscardinidae
Graphiurus sp. : : A ; ; : } ; x
Cricetidae/Muridae
Aethomys sp. A. : : ; : : 5 ; x
Aethomys sp. B x
Mus sp. A <
Mus sp. B : : x
Rhabdomys sp. : : : x
Otomyinae gen. & sp. nov. . x
Mystromys sp. A “<
Mystromys cf. darti x
Mystromys cf. hausleitneri <
Desmodillus sp. x
Dendromus sp. x
Steatomys sp. . : ; : ‘ s , e : x
Rodentia gen. & spp. indet. : : ; 2 : ; x
CETACEA
Gen. & spp. indet. 3 ; , f : ; : : x x

By far the best represented species is the seal, Prionodelphis capensis
(Hendey & Repenning 1972). At the time that it was described, the species
assemblage was already unusually large for a fossil phocid and since then many
new specimens have been collected. P. capensis is evidently the best represented
fossil phocid in the world. New material includes several nearly complete skulls
(Fig. 6), parts of many more and hundreds of postcranial bones. Most of the
material is from the Pelletal Phosphorite Member, the newest and best specimens
having come from bed 3aN.

In view of the environments of deposition in the Varswater Formation, other
aquatic carnivores are surprisingly rare. Only one specimen belonging to the
otter, Enhydriodon africanus, is known, although the assemblage does include one
unidentified carnivore which might also have been an otter (Hendey 1974a).

Until recently Canidae were known only on the basis of a few isolated teeth
belonging to a small species and, as with the primate, there had been a growing
doubt about identification. This doubt was dispelled by the discovery of a skull
and associated postcranial bones belonging to the species. The skull has
characters which indicate that the animal concerned was a fox (Vulpes sp.). It is
known only from the Pelletal Phosphorite Member.

Another of the carnivores from this member, the bear, Agriotherium
africanum (Hendey 1972b), remains one of the rarer elements in the fauna,
although some additional postcranial bones were found recently.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Dorsal, lateral and ventral views of a skull of the seal, Prionodelphis capensis
(SAM-PQ-L 31976) from bed 3aN, Pelletal Phosphorite Member, ‘E’ Quarry. Scale in
centimetres.

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 239

Agriotherium was an unexpected addition to the ‘E’ Quarry fauna, being the
first African record of the subfamily to which it belongs. Another similarly
‘exotic’ species was recently tentatively identified, this time a procyonid. The
Procyonidae were widely distributed in Eurasia and North America during the
late Tertiary, but have not previously been recorded from Africa. The Lange-
baanweg procyonid, if that is indeed what it is, may prove to be an enduring
problem since it is poorly represented, with the only specimens having come
from an area now covered by a mine dump. The specimens in question are
fragments of the skull of an immature individual and parts of the hindfoot of an
adult.

The Mustelidae include another of the new records for the site and this
species is as exotic as the Agriotherium and ?procyonid in terms of the known
distribution of its closest relatives. The new species is a giant form whose size
approaches that of Megalictis, from the North American Miocene, which is the
largest of all known mustelids. It belongs to a group of wolverine-like carnivores
which includes Megalictis and which has previously been recorded only in
Eurasia and North America. The presence in Africa of a member of this group is
in a sense even more unexpected than the presence of the bear and ?procyonid
since, unlike them, the giant wolverines are not recorded from southern Asia,
which is the Eurasian region with the greatest faunal resemblances to Africa.

The giant wolverine and ?procyonid, which are both from the Quartzose
Sand Member, join the list of Langebaanian species that belong to groups with
an essentially Eurasiatic (and North American) record (Hendey 1974a: 61).

Apart from the otter, or otters, and the giant wolverine, the only other
mustelid known from ‘E’ Quarry is a small, poorly represented species of
Mellivora (Hendey 1974a), one additional specimen of which was recently
collected from the Pelletal Phosphorite Member.

The Viverridae is the most diverse of the carnivore families known from the
Varswater Formation. Previously only one civet (Viverra leakeyi) had been
recorded, but new material indicates that there is at least one other species as
well. The available civet material is problematical largely because it is so frag-
mentary. A large number of new specimens belonging to smaller viverrids have
been discovered. Most of the material belongs to herpestines, but a small genet
is also represented. When the herpestines were first studied (Hendey 1974a) only
two species were recognized (Herpestes spp. A and B) and they are now the most
commonly represented viverrids in the ‘E’ Quarry assemblage. The new material
includes a few specimens which apparently belong to three additional species
(Hendey 1974b: 157).

The most common of the larger terrestrial carnivores from the site are
hyaenas, which now include one additional record, an apparently new species of
Euryboas. The more advanced species of this genus were the long-legged and
sharp-toothed ‘hunting hyaenas’ which occurred in Africa and southern Europe
early in the Pleistocene (Hendey 1975). Four hyaena species had previously been
recorded, namely, Percrocuta australis, Hyaena abronia, hyaenid species B and

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hyaenictis preforfex, while a fifth, hyaenid species E, was tentatively identified
(Hendey 1974a). The status of the latter remains unresolved. Additional speci-
mens, readily identifiable with P. australis, H. abronia and Species B, have been
discovered, but H. preforfex is still only known from the remains of a single, aged
individual. A detailed study of the hyaenid material now available is warranted.
H. abronia, the best represented of the species, may prove particularly useful in
resolving the problem of whether or not the Quartzose Sand and Pelletal
Phosphorite Members are substantially different in age (vide infra).

The Felidae also include a recent new record, a small wildcat-sized species
from the Quartzose Sand Member. Although specimens in addition to those
already described have been discovered, only the false sabretooth, Dinofelis
diastemata, is reasonably well represented.

Proboscideans

The Varswater Formation has yielded relatively few proboscidean specimens
and those that have been found are generally fragmentary. The only exception
is the incomplete skeleton of an elephant from the East Stream exposures of
the Quartzose Sand Member (Maglio & Hendey 1970; Hendey 1974a: 349).
Remains of an unidentified gomphothere occur more commonly than those of
the elephant, Mammuthus subplanifrons.

Perissodactyls

Only one rhinoceros, Ceratotherium praecox, is known from the Varswater
Formation (Hooijer 1972). It is perhaps the most commonly represented of the
larger mammals from the Quartzose Sand Member, but only a few fragmentary
specimens from the Pelletal Phosphorite Member may belong to this species.

The situation in respect of the Equidae is more complex. The material from
the Quartzose Sand Member is here regarded as belonging to two species of
Hipparion, an opinion rejected by Hooijer (in preparation). A few specimens
thought to be from the Pelletal Phosphorite Member were excluded from
Table 4 because of uncertainties about their provenance. Otherwise the only
significant specimen from this member is an incomplete lower dentition from the
uppermost level of the deposits in the bed 3aN area. Hooijer (in preparation)
has referred it to Hipparion namaquense Haughton, 1932. It is notable in being
the only record of this species in the Langebaanweg area. Also no other speci-
mens from the same horizon have yet been positively identified.

The Equidae, perhaps more than any other group represented in the various
fossil occurrences near Langebaanweg, are potentially important in resolving the
problems of relative age and stratigraphic relationships of the deposits in which
they occur. For example, although the Gravel Member in ‘E’ Quarry has
produced few useful terrestrial vertebrate fossils, one interesting exception is an
incomplete equid tooth, which, together with two teeth from the same horizon
in ‘C’ Quarry, is tentatively identified with the Miocene species, Hipparion
primigenium. Hooijer (in preparation) also rejects this identification, but if the

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 241

material does indeed represent a Miocene species, it must have been reworked
from the underlying Saldanha Formation. This formation would therefore be
younger than 12,5 m.y. B.P., since before this date Hipparion was not present in
Africa, or elsewhere in the Old World (Hooijer 1975).

The Baard’s Quarry assemblage includes both Hipparion and Equus, the
latter indicating that at least part of the fauna is younger than that from the
Varswater Formation. The Hipparion material from Baard’s has recently proved
to be as controversial as that from ‘E’ Quarry, but the assemblage is here
regarded as being comprised of two species, the more commonly represented
being distinct from the ‘E’ Quarry hipparions. Earlier it had been indicated that
only one Hipparion was represented in the Langebaanweg occurrences (Hendey
1972a; 1974a), but this is now discounted. The various issues relating to the
Langebaanweg equids should eventually be resolved satisfactorily.

Artiodactyls

Perhaps the most unexpected of the recent new records from ‘*E’ Quarry is
that of a peccary. Previously the species concerned had been regarded as a
miniature pig (Hendey 1974a: 47), but additional material from bed 3aN led to
the revised identification being suggested by both A. W. Gentry (British Museum
(Natural History)) and H. B. S. Cooke (Dalhousie University) (pers. comm.).
This is the first African record of the family Tayassuidae and it is also the most
recent Old World occurrence. The nearest peccary record in both a geographic
and temporal sense is the Miocene Pecarichoerus orientalis from the Siwalik
Hills of India (Colbert 1933). The Langebaanweg peccary, which is one of the
smallest ever recorded, was the subject of a recent preliminary study (Hendey
in press).

Nyanzachoerus is the only pig which occurs in the Varswater Formation and
it is known from most of the fossiliferous exposures of the Quartzose Sand
Member, having been particularly common in the East Stream area. The species
from this member is remarkably well represented, the available assemblage being
larger than those of previously described members of this genus. Recently
Nyanzachoerus was recorded from the Pelletal Phosphorite Member for the first
time. The specimen concerned, a fragmented and incomplete skull, differs in
some respects from the Quartzose Sand Member specimens and probably
represents a second species. Although not yet studied in detail, the Quartzose
Sand Member Nyanzachoerus has already proved useful ‘in the relative dating of
the deposits (Hendey 1973), and the Pelletal Phosphorite Member species
promises to be equally useful in this respect (see p. 244).

The apparent absence of hippopotamus from the ‘E’ Quarry fauna has
previously given rise to comment, since the depositional environments of the
Varswater Formation were such that this animal might have been expected to
occur quite commonly (Hendey 1974a: 48). A few fragmentary hippo remains
were found for the first time during 1975 in the bed 3aN exposures of the
Pelletal Phosphorite Member. The fact that hippos are rare in this member and

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

that they are still not recorded from the extensively sampled Quartzose Sand
Member has yet to be satisfactorily explained.

The ‘E’ Quarry Giraffidae are fairly well represented by elements of the
postcranial skeleton, but cranial material is rare and invariably fragmented and
incomplete. Some of the best giraffid specimens, belonging to both Sivatherium
and Giraffa, have come from bed 3aN. The material is being studied by
J. M. Harris (Kenya National Museum).

One additional bovid species, a reduncine, was recently recorded from
‘E’ Quarry. Curiously, Bovidae are not as well represented at this site in terms of
numbers of individuals as they are at some of the Pleistocene fossil occurrences
in the region. In addition, there are relatively few species recorded from the
Quartzose Sand Member, which is otherwise remarkable for the diversity of
species represented. On the other hand, certain deposits tend to include
individuals of particular species in numbers which are disproportionately high
in terms of their bovid assemblages as a whole. For example, the boselaphine,
Mesembriportax acrae (Gentry 1974), is by far the most commonly occurring
bovid in the Quartzose Sand Member, while in bed 3aS it is alcelaphines which
are abundant. At least some of the Quartzose Sand Member boselaphine
remains are believed to have accumulated sub-aerially, indicating that this
species was an inhabitant of the area now exposed in the mine. Gentry (1974)
suggested that the boselaphine was an open woodland species and this sort of
habitat may well have existed in the immediate vicinity of the old estuary. The
situation in respect of the Pelletal Phosphorite alcelaphines was probably quite
different. Many, and perhaps all of the specimens were apparently washed to
their final resting-places by the river and their carcasses may have originated
upstream where the river crossed open plains, the probable preferred habitat of
the alcelaphines.

Other mammals

Other terrestrial mammals recorded from the Varswater Formation include
a pangolin, an aardvark, a dassie (hyrax) and a porcupine. All are rare. Cetacea
are more common, but are represented mainly by undiagnostic postcranial
bones. Both whales and dolphins occur, remains of the latter having been found
for the first time during 1975.

The porcupine is known only from a fragmented skull of a large and
unidentified species which is currently being studied by Judy M. Maguire
(Bernard Price Institute for Palaeontological Research). This specimen was
found in the Quartzose Sand Member shortly after publication of a comment on
the supposed absence of porcupines from the Varswater Formation (Hendey
1974a: 42).

The dassie, Procavia cf. antigua, was originally identified on the basis of
very fragmentary specimens, but more and better material is now available.
The most complete specimens are from easterly exposures of the Quartzose
Sand Member.

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 243

The aardvark is amongst the most poorly represented of the ‘E’ Quarry
mammals and until recently the same applied to the pangolin. The situation in
respect of the latter has been slightly improved by the discovery of an incomplete
skeleton, including parts of the skull, much of the tail and parts of all four limbs.

FLORA

Prior to the exposure of the peat bed the only botanical remains recovered
from the ‘E’ Quarry exposures of the Varswater Formation were some fossil
root fragments from the Quartzose Sand Member. A series of samples from the
peat bed submitted to the Institute for Environmental Sciences at the University
of the Orange Free State included one which was pollen-rich. Approximately
92 per cent of the sporomorphae belonged to one unidentified taxon (E. M.
van Zinderen Bakker pers. comm. to A. J. Tankard). This may represent a
locally abundant marsh plant. Interestingly, both tree and grass pollens were
identified and this lends support to the earlier suggestion that these vegetation
types must have been present in the area at the time that the deposits were laid
down (Hendey 1973).

During 1975 two boreholes were sunk from the floor of the mine into
deposits underlying the Varswater Formation. They intersected two peat
horizons which are evidently part of the Miocene Saldanha Formation. These
peats contain both visible and microscopic plant remains.

DATING

A Pliocene age for the Varswater Formation, with an inferred chronometric
date of 4-5 m.y. B.P., is still accepted, but there have been new developments
concerning the dating of fossils from the Gravel Member and the age difference
between the Quartzose Sand and Pelletal Phosphorite Members.

The Gravel Member is composed largely of an abraded and fragmented
phosphatic rock which is known to contain bone fragments and which is
undoubtedly pre-Pliocene in age. The rock, and other deposits with which it
may have been associated, was eroded during the early stages of the Pliocene
marine transgression (Fig. 2). The possibility that some fossils might have been
reworked from the deposits truncated by the transgression has been recognized,
but was not substantiated until a study of the Gravel Member Selachii by
P. A. Hulley revealed that some specimens are evidently of pre-Pliocene age.
The derived fossils may also include the few isolated Hipparion teeth referred to
earlier (see p. 240). The derived pre-Pliocene element in the Gravel Member
fossil assemblage is likely to make up but a small part of the assemblage as a
whole.

Still unresolved is the question of the time taken for the Varswater Forma-
tion to accumulate. There were clearly intervals of time between the deposition

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

of fossils at different levels in the succession, but so far as is known the only one
which may have been of sufficient duration to be palaeontologically significant
was that between the deposition of the Quartzose Sand and Pelletal Phosphorite
Members. The original opinion that the faunas of the two members were
broadly contemporaneous (Hendey 19705) was based on the fact that they co
have species in common and on the belief that differences between them were
due simply to a general dissimilarity in the modes and environments of
deposition of the fossils.

On the other hand, the Pelletal Phosphorite Member does postdate the
Quartzose Sand Member and it has been recognized that the time factor may
have been significant enough to be reflected in the characteristics of individual
species represented in the succession and to have influenced the overall compo-
sition of the two faunas. Previous studies on species common to the two faunas
have provided no conclusive evidence of significant evolutionary changes.
For example, the suggestion that the Pelletal Phosphorite Viverra leakeyi might
be a more advanced variety of the same species from the Quartzose Sand Member
(Hendey 1974a: 81) has still not been substantiated. In this and other instances
comparisons have been complicated mainly by inadequacies in the available
material.

The most important indication to date that the two faunas may be separated
by a substantial period in time came with the discovery of the first Nyanzachoerus
specimen from the Pelletal Phosphorite Member. The new specimen is in
certain respects more advanced than the Quartzose Sand Member Nyanzachoerus
and if there was an ancestor/descendant relationship between the two forms,
their difference in age may be of the order of several hundred thousand years.
Although comparisons with Nyanzachoerus species recorded elsewhere are
‘complicated by the uniqueness of the ‘E’ Quarry forms’, indications are that
the one from the Pelletal Phosphorite is unlikely to be less than 4 m.y. old, while
that from the Quartzose Sand dates back probably no more than 5 m.y.

Now that more material from the Pelletal Phosphorite Member is becoming
available, the potential for meaningful comparisons between the two main
faunas from ‘E’ Quarry is greatly increased.

Relative dating of the ‘E’ Quarry fossils and deposits has been based on
comparisons of some mammals with their counterparts at various localities in
East Africa. The fact that this dating is neither very secure nor very precise is
due to the small number of species which have been appropriate for such
comparisons. It is increasingly apparent that the Langebaanweg fauna is a good
deal more unique than had hitherto been supposed and although individual
species do have much in common with their contemporaries further north, they
are not always identical. If the fauna does indeed include local endemics, and
also perhaps late survivors of lineages which were already extinct elsewhere, the
faunal dating of the occurrences will become complicated and less satisfactory.

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 245

CONCLUSIONS

The ‘E’ Quarry fossil occurrences, which are the most important in the
Langebaanweg area, are significant for a variety of reasons. There are very few
recorded sites in southern Africa which have yielded vertebrate fossils of
comparable age and the assemblages from such sites are minuscule by com-
parison. Consequently the only substantial information on the nature of the
subcontinental vertebrate fauna of 4-5 million years ago is derived from the
‘E’ Quarry record. The nearest important fossil occurrences which are broadly
contemporaneous are in East Africa, about 4 500 kilometres away (e.g. Vogel
River Series, Lower Kaiso Formation, Mursi Formation of the Omo Group,
Kanapoi/Lothagam and Kubi Algi at East Rudolf). However, not even these
occurrences have produced assemblages so large and so diverse as that from
‘E’ Quarry. Elsewhere in Africa the Pliocene fossil record is either poor or non-
existent, so Langebaanweg is an especially important source of information on
the animal life of this epoch not only in a local sense, but for Africa as a whole.
It has the additional merit of being the only important Pliocene occurrence on
the continent where both marine and terrestrial faunas are represented.

Certain of the “E’ Quarry species assemblages are already impressively large
and, since collecting is continuing, they are still growing. The preservation of
specimens is generally very good, all skeletal elements are represented (and
collected) and unequivocal associations of cranial and postcranial material are
not uncommon. It should therefore ultimately prove possible to provide
comprehensive definitions of many species and this may well assist in resolving
identification problems in the smaller assemblages from other African sites.

The species diversity is remarkable by any standards and as a result the
fauna as a whole will be better known than those of many other sites where
important elements may be lacking (e.g. vertebrate microfauna, birds, aquatic
species).

The ‘E’ Quarry fauna, like many others 1n southern Africa, has so far been
dated only in a relative sense and in this respect it is more problematical than
many from East Africa. However, the Langebaanweg occurrences are amongst
the very few of any importance in southern Africa where generally acceptable
geological dating is also possible.

Perhaps the biggest drawback of the site lies with its situation at the
southern continental extremity, far from the main focus of African late Cenozoic
palaeontological investigation (i.e. East Africa). On the other hand, this may
ultimately invest it with a particular interest since, as the contemporary East
African fauna becomes better known, similarities and differences of
zoogeographic significance may emerge.

ACKNOWLEDGEMENTS

During the past two years a number of persons have contributed directly
and indirectly to the furthering of the Langebaanweg research project and | am

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

greatly indebted to them for their interest and contributions. Particular thanks
are due to Mr G. Avery, Dr P. A. Hulley and Dr B. Kensley (South African
Museum), Dr A. W. Gentry (British Museum (Natural History)), Dr J. M. Harris
(Kenya National Museum), Dr D. A. Hooijer (Rijksmuseum van Natuurlijke
Historie), Dr R. G. Klein (University of Chicago), Mr T. N. Pocock (Vanderbijl-
park) and Dr G. G. Simpson (The Simroe Foundation and University of
Arizona). Dr A. J. Tankard (South African Museum), who has undertaken the
geological study at Langebaanweg, patiently discussed the geology of the
deposits with the result that several points concerning the depositional environ-
ments of the fossils were clarified. Mr G. Benfield, the Mine Superintendent at
Langebaanweg, has been unfailingly helpful and, together with other mine
employees, has kept the project viable.

The fieldwork at Langebaanweg is financed by the South African Council
for Scientific and Industrial Research. The undertaking is aided on site in many
ways by Chemfos Ltd., a subsidiary of Samancor. The South African Air Force
has provided aerial photographs. The Wenner-Gren Foundation for Anthropo-
logical Research (New York) provided the vehicle used for the fieldwork
(Grant no. 2752-1834). The assistance of these organizations is gratefully
acknowledged.

REFERENCES

COLBERT, E. H. 1933. An Upper Tertiary peccary from India.— Am. Mus. Novit. 635: 1-9.

GENTRY, A. W. 1974. A new genus and species of Pliocene boselaphine (Bovidae, Mammalia)
from South Africa.— Ann. S. Afr. Mus. 65: 145-188.

HENDEY, Q. B. 1970a. A review of the geology and palaeontology of the Plio/Pleistocene
deposits at Langebaanweg, Cape Province.— Ann. S. Afr. Mus. 56: 75-117.

HENDEY, Q. B. 1970b. The age of the fossiliferous deposits at Langebaanweg, Cape Province. —
Ann. S. Afr. Mus. 56: 119-131.

HENDEY, Q. B. 1972a. Further observations on the age of the mammalian fauna from Lange-
baanweg, Cape Province. —Palaeoecol. Afr. 6: 172-175.

HENDEY, Q. B. 1972b. A Pliocene ursid from South Africa.— Ann. S. Afr. Mus. 59: 115-132.

HENDEY, Q. B. 1973. Fossil occurrences at Langebaanweg, Cape Province.— Nature, Lond.
244: 13-14.

HENDEY, Q. B. 1974a. The late Cenozoic Carnivora of the south-western Cape Province. — Ann.
S. Afr. Mus. 63: 1-369.

HENDEY, Q. B. 1974b. Faunal dating of the late Cenozoic of southern Africa, with special
reference to the Carnivora.— Quat. Res. 4: 149-161.

HENDEY, Q. B. 1975. Relationships of North American hyaenas.—S. Afr. J. Sci. 71: 187.

HENDEY, Q. B. In press. Fossil peccary from the Pliocene of South Africa. Science.

HENDEY, Q. B. & REPENNING, C. A. 1972. A Pliocene phocid from South Africa.— Ann. S. Afr.
Mus. 59: 71-98.

Hoover, D. A. 1972. A late Pliocene rhinoceros from Langebaanweg, Cape Province.—
Ann. S. Afr. Mus. 59: 151-191.

Hoover, D. A. 1975. The hipparions of the Baringo Basin sequence.— Nature, Lond. 254:
211-212.

KENSLEY, B. 1972. Pliocene invertebrates from Langebaanweg, Cape Province.— Ann. S. Afr.
Mus. 60: 173-190.

Mactlo, V. J. & HENDEY, Q. B. 1970. New evidence relating to the supposed stegolophodont
ancestry of the Elephantidae.—S. Afr. archaeol. Bull. 25: 85-87.

MELLETT, J. S. 1974. Scatological origin of microvertebrate fossil accumulations. — Science
185: 349-350,

PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 247

Simpson, G. G. 1971. Fossil penguin from the late Cenozoic of South Africa.— Science 171:
1144-1145.

Simpson, G. G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene
of Langebaanweg, Cape Province, South Africa.— Ann. S. Afr. Mus. 69: 59-72.

SINGER, R. 1961. The new fossil sites at Langebaanweg, South Africa.—Curr. Anthrop. 2:
385-387.

SouTH AFRICAN CODE OF STRATIGRAPHIC TERMINOLOGY AND NOMENCLATURE. 1971. Trans.
geol. Soc. S. Afr. 74: 111-131.

TANKARD, A. J. 1974a. Petrology and origin of the phosphorite and aluminium phosphate rock
of the Langebaanweg—Saldanha area, south-western Cape Province.— Ann. S. Afr. Mus.
65: 217-249.

TANKARD, A. J. 19746. Chemical composition of the phosphorites from the Langebaanweg—
Saldanha area, Cape Province.—Trans. geol. Soc. S. Afr. 77: 185-190.

TANKARD, A. J. 1975. Varswater Formation of the Langebaanweg-—Saldanha area, Cape
Province. —Trans. geol. Soc. S. Afr. 77: 265-283.

TANKARD, A. J. In press. The Saldanha Formation: a Neogene transgressive complex. — Trans.
geol. Soc. S. Afr.

6. SYSTEMATIC papers must conform with the International 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. n., sp. n., comb. n.,
syn. n., 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 (figs 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 (33.51S, 25.39E), collected by A. Smith, 15 January 1973.

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

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A. L. du Toit
Von Huene _—i but _~-F. von Huene

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 01340 8505

Q. B. HENDEY

THE PLIOCENE FOSSIL OCCURRENCES IN
‘E’ QUARRY,

LANGEBAANWEG, SOUTH AFRICA

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