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7. 68

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

VOLUME 76 BAND 76

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

VOLUME 76 BAND

HE TRUSTEES OF THE DIE TRUSTEES VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD

1978

% SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY
THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE

764

LIST OF CONTENTS

CLUVER, M. A.

The skeleton of the mammal-like reptile Cistecephalus with evidence for a fossorial
mode of life. (Published September 1978.)

CrowE, T. M.

The evolution of guinea-fowl (Galliformes, Phasianidae, Numidinae), taxonomy,
phylogeny, speciation and biogeography. (Published September 1978.)

Day, J.

Southern African Cumacea. Part 3. Families Lampropidae and Ceratocumatidae.
(Published August 1978.) - ne ;

GRIFFITH, J.

A fragmentary specimen of Saurichthys sp. from the Upper Beaufort Series of
South Africa. (Published September 1978.) :

HENDEY, Q. B.

Preliminary report on the Miocene vertebrates from Arrisdrift, South West Africa.
(Published July 1978.) : 2

HENDEY, Q. B.

Late Tertiary Hyaenidae from Langebaanweg, South Africa, and their relevance
to the phylogeny of the family. (Published September 1978.) ..

HENDEY, Q. B.

Late Tertiary Mustelidae (Mammalia, Carnivora) from Langebaanweg, South
Africa. (Published September 1978.) : ae oF i

MCLACHLAN, A. & Moore, C. G.

Three new species of Harpacticoida (Crustacea, Copepoda) from sandy beaches
in Algoa Bay, South Africa, with keys to the genera Arenosetella, ae
Leptastacus and Psammastacus. (Published August 1978.) 3 :

Moore, C. G. see MCLACHLAN, A.

RAu, R. E.

The development of Xenopus - Rose & Hewitt pone ges ae
October 1978.)

WOOLDRIDGE, T.

Two new species of Gastrosaccus (Crustacea, Mysidacea) from sandy beaches in
Transkei. (Published September 1978.) : we

Page

2A3

43

Si

299

265

329

191

247

309

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$e

VOLUME 76 PART 1 JULY 1978 | ISSN 0303-2515

po 7.68

OF THE SOUTH AF RICAN
MUSEUM

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

FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

Fiscuer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. Zen. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
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THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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(continued inside back cover)

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

Volume 76 Band
July 1978 Julie
Part 1 Deel

PRELIMINARY REPORT ON
THE MIOCENE VERTEBRATES FROM ARRISDRIFT,
SOUTH WEST AFRICA

By

Q. B. HENDEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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

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PRELIMINARY REPORT ON THE MIOCENE VERTEBRATES FROM
ARRISDRIFT, SOUTH WEST AFRICA

By
Q. B. HENDEY

South African Museum, Cape Town

(With 13 figures and 11 tables)
[MS. accepted 14 March 1978]

ABSTRACT

At least 28 vertebrate species, of which 22 are mammals, are recorded from the early
middle Miocene (c. 16 m.y. old) fossil occurrence at Arrisdrift on the Orange River in South
West Africa. The material postdates Miocene vertebrates previously recorded from the
Namib desert. The mammals include at least 3 new species (a hyracoid, a palaeomerycid and
an ochotonid), while there are at least 8 genera represented which have not hitherto been
known in southern Africa. Austrolagomys simpsoni Hopwood, 1929, is referred to Kenyala-
gomys Whitworth, 1954, and Prohyrax is placed in the Pliohyracinae, a group which apparently
had its origins in southern Africa.

CONTENTS

PAGE
MpPROG UC OU Gre he ee mt ee eh Ne Ween 1
Other Miocene vertebrate occurrences in the Namib desert . 3
Mae tossilevertebrates from Arisdrift .. . . = . «°. 8
PRPCTOlMINelOCCUEEENCE® oan) ie ee ee wee ee Oe
NCOCMVIrOMIMent “4 (aa ilsia = to od te 1 Rs ee ee | 4
PREMMOMICUOCIMEMES i. te. esa ey aes fe 6) ew 8
References 2 Reta Me icc oi eal Re ge i WO AU 38

INTRODUCTION

Until recently the only substantial information on southern African
Miocene terrestrial vertebrates came from several small fossil assemblages
collected in the southern Namib desert (Stromer 1926; Hopwood 1929;
Hamilton & Van Couvering 1977). The described material is limited in both
quality and quantity. The discovery of a new Miocene vertebrate locality in
terrace deposits at Arrisdrift on the Orange River further south in the same
region (Fig. 1) has proved an important addition to the local Miocene fossil
record (South African Journal of Science 1976; Corvinus & Hendey 1978).
The number of fossils already collected at Arrisdrift exceeds the combined
total from the other Namib desert occurrences, although the quality of the
specimens is not necessarily superior. The presence on the subcontinent of
several taxa has been revealed for the first time. These include the deinothere,
Prodeinotherium hobleyi (Harris 1977).

The fossils were discovered in a prospect pit (Pit 2 of Drill-line AD 8)
in deposits being investigated by the Consolidated Diamond Mines of South

1

Ann. S. Afr. Mus. 76 (1), 1978: 1-41, 13 figs, 11 tables.

D ANNALS OF THE SOUTH AFRICAN MUSEUM

West Africa (Pty) Ltd. Further fossiliferous deposit has since been exposed by
extending the original pit, but the limits of the occurrence have not been
established.

The material already prepared includes remains of at least 28 vertebrate
species, of which 22 are mammals (Table 1). Most have yet to be positively
identified and studies have so far been confined largely to cranial material,
which is much less common than postcranial bones. The condition of specimens
varies considerably, some being well preserved and reasonably complete, but
most having suffered post-mortem damage. The fossils occur in a poorly sorted

TABLE |
The vertebrates from Pit 2/AD 8 at Arrisdrift, South West Africa.

OSTEICHTHYES

gen. et sp(p). indet.

AMPHIBIA gen. et sp. indet.
REPTILIA
Squamata gen. et sp. indet.
Crocodilia ? Crocodylus niloticus
Chelonia . gen. et sp(p). indet.
AVES gen. et spp. indet.
MAMMALIA
Insectivora
Macroscelididae Myohyrax cf. oswaldi
Carnivora
Amphicyonidae .. Amphicyon cf. steinheimensis

Amphicyonidae or Ursidae .

?Ursidae . ?Hemicyoninae gen. & sp. indet.

?Felidae . ? Metailurus sp.

Mustelidae ? Ischyrictis sp.

indet. gen. & sp. indet.
Hyracoidea

Procaviidae Prohyrax n. sp.
Proboscidea

Gomphotherlidae . gen. et sp. indet.

Deinotheriidae Prodeinotherium hobleyi
Perissodactyla

Rhinocerotidae Dicerorhinus sp.
Artiodactyla

Suidae gen. et sp. indet.

Lopholistriodon moruoroti

Tragulidae Dorcatherium cf. pigotti

Palaeomerycidae Climacoceras sp. nov.

Bovidae gen. et sp. indet.

Pecora gen. et sp. indet.
Lagomorpha

Ochotonidae . Kenyalagomys sp. nov.
Rodentia

?Bathyergidae ? Bathyergoides sp.

Thryonomyidae Paraphiomys pigotti

indet. gen. et spp. indet.

gen. et sp. indet.

ee eS a

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 3

fluvial gravel and their imperfections are due mainly to their having been
transported by water in a high-energy environment. Only one instance is
recorded of skeletal elements occurring in articulation and, in addition to
disarticulation and fragmentation, many specimens are abraded and distorted.
Incrustations of gypsum have etched and even destroyed parts of some speci-
mens. Since the deposit incorporating the fossils is consolidated, power tools
were required during their excavation and this has caused further damage to
specimens. In spite of its shortcomings, the Arrisdrift fossil assemblage is
perhaps the most important one of Miocene age yet discovered in southern
Africa.

The purpose of the present report is to place on record some details of the
nature and number of specimens belonging to the various taxa already recog-
nized. With the exception of the deinothere teeth, none of the material has been
thoroughly studied, although such studies will be undertaken by various
authorities in the future. The geological investigation of the deposits in the
vicinity of Arrisdrift is being undertaken by employees of the mining company
prospecting the area.

The specimens discussed in this report are housed in the Department of
Cenozoic Palaeontology at the South African Museum, Cape Town. The full
catalogue numbers begin SAM-PQ., which identify the institution and depart-
ment concerned, but this lettering is omitted in the text and only the site prefix
(AD) and serial numbers of individual specimens are given. The full site
reference is Arrisdrift, Pit 2/AD 8.

OTHER MIOCENE VERTEBRATE OCCURRENCES IN THE
NAMIB DESERT

The first Miocene vertebrates from the southern Namib desert were
discovered during the First World War and were described in a series of papers
by Stromer (1922, 1923, 1924, 1926). This material came from three localities,
namely, Elisabethfeld, 38 km south of Liideritz; a borehole near Plant 4 of
the Kolonial Bergbau Gesellschaft, 20 km south of Liideritz (= Elisabeth Bay
Pan, see Greenman 1966); and Langental near Bogenfels, 80 km south of
Lideritz (Fig. 1).

Subsequently Hopwood (1929) described another small assemblage of
specimens from the same region, but his material lacks precise locality data
and was recorded as being from ‘south of Liideritz Bay’.

Little additional material was collected in the region in the decades which
followed. The South African Museum has an undescribed ruminant mandible
fragment (SAM-PQ-G 8356) from Bogenfels, which may be a synonym of
Stromer’s Langental locality. Some fragmentary material was collected by
Greenman (1966) from Fiskus and Grillental in the Elisabethfeld area. This
material is also in the South African Museum, but includes little that is
diagnostic.

ANNALS OF THE SOUTH AFRICAN MUSEUM

XPLANT 4

XxELISABETHFELD

ARRISDRIE

Alexander
)
Bay

Fig. 1. Location of Arrisdrift and other South West African Miocene fossil occurrences.

Hamilton & Van Couvering (1977) recently revisited the area and collected
more material from the various localities. They have reviewed and supple-
mented the original faunal lists and compared and contrasted a revised list
with others from early Miocene occurrences elsewhere in Africa.

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 5

The Namib fossils are generally regarded as early Miocene (‘Burdigalian’)
in age, their source is usually recorded as ‘Namib desert’ or ‘South West Africa’,
and they are treated as if they were a single assemblage. The described material
is here listed as four separate assemblages (Table 2). Since each is limited in
size and each includes unidentified or incompletely identified taxa, the tases
for comparing them to each other, and to assemblages elsewhere, are limited.
The comments which follow are confined largely to the implications of indi-
vidual taxa in respect of the age of the assemblages. Other references to this
material are included in the discussions on the Arrisdrift fossils.

There is one species from Elisabethfeld, Metapterodon kaiseri, which
Savage (1965) believed to be represented in east Africa by specimens from
Karungu and Rusinga, which are between 18 and 20 m.y. old. Savage also
recognized a second species of Metapterodon from Rusinga, namely, M. zadoki.
Van Valen (1967: 252) found ‘that the two east African species distinguished
by Savage are much more similar to each other than are the east and Southwest
African forms of “‘“M. kaiseri”’. He concluded that the east African species
are more advanced than M. kaiseri and synonymized Metapterodon with
Pterodon. Although this material is problematical, Van Valen’s opinion suggests
that the Elisabethfeld species may predate its east African counterparts. Its
age might therefore be greater than 20 m.y., that is, “‘Aquitanian’ rather than
‘Burdigalian’ in terms of the European mammal age nomenclature (Van
Couvering 1972).

Another of the Elisabethfeld species, Myohyrax doederleini, was regarded
as a synonym of the east African M. oswaldi (Whitworth 1954; Patterson 1965),
a species which has a recorded age range of 18 to 22 m.y. (see Whitworth 1954;
Walker 1969; Van Couvering 1972). An ‘Aquitanian’ to ‘Burdigalian’ age is
therefore indicated. Since a Myohyrax resembling M. oswaldi is now also
recorded from Arrisdrift, this species may have survived beyond the ‘Burdi-
galian’ (see below), and appears to be of little use for relative dating purposes.

The Elisabethfeld Propalaeoryx austroafricanus is a primitive ruminant
which is likely to be broadly contemporaneous with the Rusinga P. nyanzae
(Whitworth 1958; Hamilton 1973), but the available material of these species
is too scanty to determine possible differences of temporal significance.

The only other identified species from Elisabethfeld are Parapedetes
namaquensis and Austrolagomys inexpectatus. They are not known elsewhere,
although there are related genera, Megapedetes and Kenyalagomys, recorded
from the early Miocene of east Africa, again from the 18 to 22 m.y. period.

According to MaclInnes (1957) Megapedetes is less specialised than
Parapedetes, but he did not regard this as indicating an age difference, ascribing
it instead to different evolutionary trends on two contemporary lineages. This
interpretation raises the question of whether assemblages such as those from
Elisabethfeld can be dated in a relative sense by comparing them with east
African assemblages. If MacInnes’s interpretation of the pedetids is correct,
then it follows that seemingly primitive taxa such as Pterodon kaiseri may

ANNALS OF THE SOUTH AFRICAN MUSEUM

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MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA i

simply be conservative southern counterparts of ‘advanced’ east African species.

This problem arises again with the ochotonids. There are clear differences
between Austrolagomys and Kenyalagomys (MacInnes 1953), which may be
interpreted as indicating that the former is the more primitive (Cooke 1972).
In this instance the fact that the ochotonid represented at Arrisdrift (which
evidently does postdate both the Elisabethfeld and the east African ‘Rusinga-
type’ faunas) 1s a Kenyalagomys, may be an indication that Austrolagomys
really is a primitive and early form.

To sum up, there is some evidence to suggest that the Elisabethfeld fauna
represents a southern African equivalent of that of the European Aquitanian.
This conclusion is, however, tentative and should be re-examined if more
material becomes available and/or when the zoogeographic relationships of
east and southern African early Miocene faunas are better understood.

The Langental fauna includes five species which are known from the early
Miocene of east Africa. They are Bathyergoides neotertiarius, Paraphiomys
pigotti, Diamantomys luederitzi (Lavocat 1973), Xenochoerus africanus
(Wilkinson 1976) and Brachypotherium heinzelini (Heissig 1971). In addition,
Prohyrax tertiarius is a primitive species and apparently consistent with an
early Miocene date (see p. 33). This hyrax is regarded as one of the more
certain indications that there was some endemism in southern African faunas
during the earlier part of the Miocene and that the complication in comparing
east and southern African taxa mentioned above does have some substance.

The other identified taxa from Langental, Protypotheroides beetzi and
Pomonomys dubius, are apparently known only from the Namib desert and,
since they may be southern endemics, they may not be useful for relative dating
purposes.

The available evidence indicates that the Langental fauna dates from the
early Miocene and that it may be a ‘Burdigalian’ rather than ‘Aquitanian’
equivalent.

The Plant 4 borehole and Langental faunas have three species in common,
namely, Myohyrax oswaldi, Bathyergoides neotertiarius and Paraphiomys
pigotti. This suggests that the former may also be of early Miocene age. On the
other hand, all three taxa are, or may be represented at Arrisdrift as well (see
below), so a slightly younger age (early middle Miocene) is also possible.

The material described by Hopwood (1929) includes three species known
from the east African early Miocene. They are Paraphiomys pigotti, P. stromeri
and Myohyrax oswaldi. In addition, this assemblage includes an ochotonid
which is apparently closely related to the east African Kenyalagomys minor
(see p. 31). Hopwood’s material also includes the large myohyracine, Protypo-
theroides beetzi, which is represented at Langental. Once again an early Miocene
age is indicated. In view of the earlier comments on ochotonids, the presence of
Kenyalagomys in Hopwood’s assemblage may mean that this assemblage, or
part of it, postdates that from Elisabethfeld.

In spite of the uncertainties relating to the four assemblages, there is no

§ ANNALS OF THE SOUTH AFRICAN MUSEUM

justification for the recent practice of treating the faunas as a unit. Even if it
could be established that they are exact contemporaries, it is as well to accord
them individual status. Since the Arrisdrift fauna is younger than some or all
of those from the Lideritz—Bogenfels area, there is certainly no justification
for adding it to the Namib Miocene mixture, and for this reason alone it will
now be inconvenient and inappropriate to refer to ‘the Miocene fauna’ from
this region.

THE FOSSIL VERTEBRATES FROM ARRISDRIFT
CLASS OSTEICHTHYES

One or more species of fish are represented by a few isolated vertebrae
(e.g. AD 668, AD 672) and fin spines (e.g. AD 759, AD 779). The latter
apparently belong to catfish (Clartidae).

CLASS AMPHIBIA

A single postcranial bone (AD 811) belongs to a frog or toad.

CLASS REPTILIA

ORDER SQUAMATA

There are two snake vertebrae (AD 707, AD 1110) in the assemblage.

ORDER CROCODILIA

? Crocodylus niloticus Laurenti, 1768

The most commonly represented lower vertebrate is a crocodile, probably
Crocodylus niloticus, of which many isolated teeth (e.g. AD 71, AD 310) and
scutes (e.g. AD 335, AD 341) are preserved. Postcranial bones and skull frag-
ments are less common. The best specimens include two incomplete dentaries
(AD 344, AD 999). Crocodiles are not uncommon as fossils in east Africa and
elsewhere but have not previously been recorded from Tertiary occurrences in
southern Africa and are rare in Quarternary deposits.

ORDER CHELONIA

A few isolated scutes (e.g. AD 73, AD 832) belong to one or more species
of tortoise.

CLASS AVES

Birds are represented by a few isolated and incomplete postcranial bones
(e.g. AD 725, AD 841) belonging to more than one species.

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 9

CLASS MAMMALIA
ORDER INSECTIVORA
Family Macroscelididae

Myohyrax cf. oswaldi Andrews, 1914

A Myohyrax is the most commonly occurring small mammal in the
assemblage and is represented by many isolated teeth and mandible fragments
(e.g. AD 125, AD 1104) (Fig. 2). Most of the mammalian postcranial bones
have yet to be classified and this material probably includes specimens belonging
to Myohyrax.

Stromer (1926) recorded three myohyracines from the Namib, namely,
M. oswaldi, M. doederleini and Protypotheroides beetzi, while Hopwood (1929)
subsequently named an additional species, M. osborni, from the same region.
In the most recent review of this material, Patterson (1965) recognized only
M. oswaldi (including M. doederleini) and P. beetzi (including M. osborni). In
doing so he followed Whitworth (1954), except that Whitworth did not regard
Protypotheroides as a valid genus.

The Arrisdrift Myohyrax is apparently indistinguishable from M. oswaldi,
but the identification is qualified since the teeth are a little smaller than those

Fig. 2. Occlusal and buccal views of Myohyrax cf. oswaldi mandible (AD 971) from Arrisdrift.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

of typical east African M. oswaldi. This was one of the features which Stromer
(1926) claimed characterized M. doederleini. Whitworth (1954) found that
M. doederleini fell within the size variation observed in east African M. oswaldi,
but since Arrisdrift provides a second sample of specimens from the Namib
in which the teeth are comparatively small, there may be a taxonomically
significant mean difference between the Myohyrax from the two regions. In
addition, since examination of the Arrisdrift Myohyrax during the present
study was cursory, the material may differ from typical M. oswaldi in characters
other than size.

The Arrisdrift Myohyrax is evidently younger than any previously recorded
myohyracine (see p. 32), and this is another factor to be taken into account
when the material is studied in detail.

ORDER CARNIVORA

Although poorly represented, the only identifiable carnivores from
Arrisdrift are Carnivora rather than Creodonta. It is, however, possible that
certain non-diagnostic specimens such as isolated canines and postcranial bones
do belong to creodonts, a group whose presence is to be expected in view of
the apparent age of the Arrisdrift assemblage (see p. 35).

Family Amphicyonidae
Amphicyon cf. steinheimensis Fraas, 1885

An incomplete left mandible (AD 133) belongs to an amphicyonid (Fig. 3,
Table 3). There are also several postcranial bones which may belong to this

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Fig. 3. Occlusal and buccal views of Amphicyon cf. steinheimensis mandible (AD 133) from
Arrisdrift.

11

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA

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iD ANNALS OF THE SOUTH AFRICAN MUSEUM

species, but they were excluded from consideration.

The amphicyonids, which are sometimes regarded as a subfamily within
the Canidae (e.g. Kuss 1965), or as a separate family (e.g. Hunt 1972), were a
successful, diverse and widespread group in the Old World and North America
during the Oligocene and Miocene. They are not well known in Africa and
prior to the Arrisdrift discovery had not been recorded in southern Africa. The
taxonomy of the group is complex and, in spite of recent revisions, the identifi-
cation of specimens such as AD 133 is difficult.

AD 133 lacks the ascending ramus, the incisors and P, to P;. Of the remain-
ing teeth only the Mg is largely intact, although the salient features of the C
and P, to M, are preserved. This specimen indicates that the species was an
unspecialized, slender-jawed Amphicyon of moderate size. The P, to P3, of which
only the roots or alveoli remain, are reduced in size and more or less evenly
spaced between the C and P,. The P, was single-rooted, P, had two roots which
has coalesced at the alveolar margin, while P, was also double-rooted. The P,,
which has lost the principal cusp, has an anterior accessory cusp, a larger
posterior accessory cusp and a tiny cusp on the posterior cingulum. The M,
is a high-crowned tooth with a prominent protoconid and stout metaconid.
The talonid is sectorial, it lacks the entoconid and makes up about one-third
of the length of the tooth. The M, has a double-cusped trigonid and single-
cusped talonid. The Mj has little relief on the occlusal surface and, like M,
and Mg, is relatively narrow. _

AD 133 resembles specimens belonging to the middle Miocene A. stein-
heimensis from Europe (see Kuss 1965), and is tentatively identified with this
species. It may, however, belong to a previously unrecorded African species of
Amphicyon. The only amphicyonids recorded from east Africa are two early
Miocene species, Hecubides euryodon and H. macrodon (Savage 1965). The
Arrisdrift species differs from H. eurydon in several respects, including its
larger size. It cannot be compared with H. macrodon, which is known only
from an isolated M?.

Superfamily Canoidea (senso Savage 1977)
Amphicyonidae or Hemicyoninae gen. et sp. indet.

A largely intact, but somewhat abraded right mandible with well worn
P, to M, (AD 1520) belongs to a very large canoid (Fig. 4). This specimen
compares in size with the mandible of the largest terrestrial carnivore previously
recorded from southern Africa, namely, the early Pliocene Agriotherium
africanum from Langebaanweg, Cape Province (Hendey 1972, 1977).

In some respects AD 1520 resembles the European middle to late Miocene
amphicyonid, Amphicyon major. For example, the mandibles of the two species
are of similar overall size, while the preserved teeth of AD 1520 are morpho-
logically similar to the corresponding teeth of A. major. Like all amphicyonids,
AD 1520 lacks a premasseteric fossa. Its teeth are closest in size to those of

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA is

Fig. 4. Occlusal and buccal views of large canoid mandible (AD 1520) from Arrisdrift.

later varieties of A. major (see Kuss 1965; Table 3, this report). They do, how-
ever, differ in being relatively broad, a feature which applies particularly in
the case of P,. Although lost, the P, and P; of AD 1520 were evidently also
relatively large and, together with P,, formed a closed series.

The large size of the premolars distinguishes the Arrisdrift species from
previously recorded A. major and, indeed, from all other Miocene amphicyonids.
Apparently only in certain Oligocene species are the premolars relatively
large and in the form of a closed series (see Springhorn 1977). If AD 1520 is
indeed related to A. major, it must be more primitive than recorded representa-
tives of this taxon even though it is ‘advanced’ in terms of overall size. The
origins of A. major are obscure (Kuss 1965) and it may well have arrived in
Europe as an immigrant from Africa. The Arrisdrift species may represent the
stock from which A. major was derived.

There is, however, a second alternative which must be considered. The
Hemicyoninae, a group of Miocene ursids which share many characters with
amphicyonids, also include a very large species whose origins are obscure.
This is Dinocyon thenardi of the later middle Miocene of Europe (Hiirzeler
1944). The hemicyonines also have reduced premolars, although the reduction
is not necessarily as marked as in contemporary amphicyonids. They pre-
sumably evolved from forms in which the premolars were relatively large and

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

in this respect the Arrisdrift species may be seen as an appropriate ancestor for
D. thenardi. The lower molars of the former are a little smaller than those of
the Grive St Alban D. thenardi, and in this respect as well the Arrisdrift species
is the less specialized (1.e. more primitive).

AD 1520 does, however, differ from D. thenardi, and other hemicyonines,
in lacking a premasseteric fossa. This would not necessarily exclude it from an
ancestral role, but it does suggest an amphicyonid, rather than hemicyonine
connection.

A third alternative is that the Arrisdrift species represents the stock from
which both A. major and D. thenardi were derived. It has been suggested that
Amphicyon and Dinocyon are closely related (e.g. Matthew 1924), although
more recent interpretations of canoid inter-relationships indicate that similari-
ties between these taxa are due to parallel evolution. Nevertheless, the fact
that neither A. major nor D. thenardi have known immediate ancestors does
raise the possibility that they may have had one in common.

Finally, AD 1520 may belong to a species related to the early Miocene
Afrocyon burolleti from Gebel Zelten in Libya (Arambourg 1961). The holotype
of this species is a mandible fragment with P, to M3, which is similar in overall
size to AD 1520, but which differs in having smaller P, to M, and a double-
rooted M.. If the two forms are indeed related, then A. burolleti is clearly the
more primitive and the Arrisdrift species could still be ancestral to the European
A. major and/or D. thenardi.

In view of the uncertainties about the relationships of AD 1520, it would
be fruitless at this stage to consider the taxonomic implications of the various
alternatives mentioned above. Although unidentified, AD 1520 is still significant
in revealing the presence of a type of carnivore not hitherto known from the
Miocene of southern Africa and in suggesting that the phylogeny of similar
taxa elsewhere may require reinterpretation.

? Family Ursidae
? Hemicyoninae gen. et sp. indet.

A mandible fragment (AD 611) belongs to a carnivore intermediate in
size between the Amphicyon cf. steinheimensis and the large canoid discussed
above (Table 3). Only the anterior part of the mandibular corpus is preserved,
and of the teeth only the roots or alveoli of P, to P, and a small part of the
crown of M, are preserved. It is readily distinguished from the Amphicyon
cf. steinheimensis by its larger size and in having a deep mandibular corpus. In
the latter respect it resembles the large canoid, but the overall size difference
is more than would be expected in intra-specific variation.

Although AD 611 has yet to be positively identified, it matches in size and
other observable respects corresponding parts of the Hemicyon californicus
holotype from the Miocene of North America (Frick 1926: 34, fig. 12B). It is
larger than specimens of European Hemicyon and Harpaleocyon described by

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 1S)

Hiirzeler (1944), but since hemicyonines and other ursids exhibit appreciable
sexual dimorphism (see Colbert 1939), size differences are not necessarily a
reliable criterion for distinguishing species. Even if it could be established that
AD 611 represents a hemicyonine, it is unlikely that the species concerned could
be identified.

Hemicyonines have not previously been recorded from Africa, but
elsewhere they occur in association with some of the characteristically Miocene
taxa which have been found at Arrisdrift. The size of AD 611 suggests a middle
Miocene rather than earlier age.

? Family Felidae

? Metailurus sp.

An isolated upper canine (AD 616) apparently belongs to a large felid of
the group which includes the extinct genera Metailurus and Dinofelis. The
specimen is 16 mm long, 12,6 mm wide and has a crown height of 44 mm.

The species concerned was larger than the only Metailurus hitherto recorded
in Africa, namely, M. africanus from the east African early Miocene (Savage
1965). AD 616 compares in size with the C of M. major from the late Miocene
of China (Zdansky 1924), but differs in being slightly shorter and broader. It
also resembles the C of the early Pliocene Dinofelis aff. diastemata from Lange-
baanweg (Hendey 1974), particularly the specimen SAM-PQ-L 20685. Once
again the only difference is that AD 616 is a little shorter and broader. This
difference suggests that the Arrisdrift species was less advanced than the other
two species.

Since carnivore canines are not necessarily diagnostic, AD 616 is only
tentatively identified and is referred to Metailurus, a Miocene genus, rather
than Dinofelis, a Plio/Pleistocene genus, in view of the age of the Arrisdrift
assemblage.

Family Mustelidae
? Ischyrictis sp.

An incomplete left mandible (AD 614) belongs to a small carnivore with
a high-crowned canine, relatively narrow and high-crowned premolars and
carnassial, and a reduced M, (Fig. 5, Table 3). The M, has a small metaconid,
while the talonid is short and lacks the entoconid. The small M, and sectorial
M, talonid suggests that the relationships of AD 614 may lie with the primitive
gulonine /schyrictis and it is identified accordingly. This genus has not pre-
viously been recorded in Africa, but is known from the Miocene of Europe
and Asia Minor (see Crusafont-Pairo 1972; Schmidt-Kittler 1976).

The high-crowned teeth and relatively narrow premolars of AD 614
would be primitive characters in an Ischyrictis, and the Arrisdrift species may
prove to be an appropriate ancestor for the early Vindobonian J. zibethoides
of Europe.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

UTVTTUTIVILUTYVTTUOTOQVACETVOUUTOTTEUOCTOT LTO ETOUUTUT TT

Fig. 5. Occlusal and lingual views of ? Ischyrictis mandible (AD 614) from Arrisdrift.

Two other mandible fragments (AD 128, AD 756) apparently belong to
the same species as AD 614.

Other Carnivora

At least one species in addition to those already mentioned is included in
the Arrisdrift carnivore assemblage. An isolated canine (AD 127) belongs to a
species smaller than the ? Jschyrictis. The ‘Carnivora gen. et sp. indet.’ in the
accompanying faunal list (Table 1) refers to this specimen.

Also unidentified are two mandible fragments (AD 139, AD 773), three
canines (AD 55, AD 122, AD 214), an I? (AD 619) and several postcranial
bones, most of which are incomplete. These specimens may include some
belonging to species in addition to those listed.

ORDER HYRACOIDEA

Family Procaviidae

Prohyrax sp. nov.

By far the most commonly represented vertebrate in the Arrisdrift assem-
blage is a hyrax belonging to a group sometimes given subfamily rank, the
Pliohyracinae (Whitworth 1954). This was the most widespread of the hyrax

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA i)

groups and is known from localities in Eurasia as well as Africa. Most of the
recorded species were extremely large in comparison to living hyracoids and
some specimens have been mistakenly identified as rhinoceroses and chali-
cotheres. Later representatives were apparently aquatic or amphibious animals
(see Osborn 1899; Vekua 1972).

As here understood, the Pliohyracinae include the following genera:

Prohyrax—trelatively small; early to middle Miocene of South West Africa
(Stromer 1923, 1926; this report)

Parapliohyrax—large; middle to late Miocene of east and north Africa (Lavocat
1961; Bishop & Pickford 1975)

Pliohyrax—very large; late Miocene and Pliocene of Europe and China (Forsyth-
Major 1899; Osborn 1899; Viret 1949; Viret & Thenius 1952; Tung &
Huang 1974)

Kvabebihyrax—very large; late Miocene of the Soviet Union (Gabunia & Vekua
1966)

Postschizotherium—very large; late Miocene and Pliocene of China (Von
Koenigswald 1966; Tung & Huang 1974)

The relationships of Prohyrax tertiarius from Langental have hitherto
been obscure since it has been known only from fragmentary material, the holo-
type being a maxillary fragment with P® to M? and part of M? (Stromer 1926:
Pl. 41, fig. 33). Only limited comparisons with the abundant Arrisdrift material
are therefore possible, but there can be little doubt that the two forms are
closely related. They are probably not conspecific since the Arrisdrift material
belongs to a larger species, but they are here taken to be congeneric. Since the
Arrisdrift species is undoubtedly a pliohyracine, Prohyrax is accordingly
included in this subfamily.

The Arrisdrift species shares some characters with other pliohyracines but
it is not conspecific with any of them, the most obvious difference being its
smaller size. It is apparently closest to the east and north African Parapliohyrax
and differs appreciably from the three Eurasian genera, which are the youngest
and most highly specialized members of the group.

The Arrisdrift hyrax is here interpreted as a new species of the genus
Prohyrax, probably directly descended from the Langental P. tertiarius and a
likely ancestor of the later pliohyracines from further north in Africa and from
Eurasia.

Although the species will be dealt with in detail elsewhere, some obser-
vations on it are included here since it is such an important element in the
Arrisdrift assemblage.

More than forty individual animals of all ontogenetic ages are represented,
mainly by mandible and maxilla fragments, although isolated teeth and post-
cranial bones are not uncommon. The best specimen is a nearly complete skull
(AD 363) which lacks only the mandible, right I?, left M+, and parts of the right
M? and right zygomatic arch (Fig. 6, Table 4). The skull is slightly distorted

18

ANNALS OF THE SOUTH AFRICAN MUSEUM

i OT MN a i ny

Fig. 6. Dorsal, lateral and ventral views of Prohyrax skull (AD 363) from Arrisdrift.

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 19

TABLE 4
Dimensions of the Prohyrax skull (AD363) from Arrisdrift.
Overall length : ; : : : : : 5 4 ; ; 5 : : E 160,8
Condylobasal length . : : 2 ‘ : : ; i ; : : . 5220
Palate length along midline . : ‘ : ‘ 3 d } : : 97,7
Anterior margin of orbit to anterior marein of 1h se : é : 4 j ; : 68,0
I’ to M? length : : ; ; : ; : : : : 4 : : , : 96,9
P! to P? length : : ‘ . : : : : ; : : : 5 é 32,9
M'to M? length . : : : : ; : : : : ‘ ; ; : Sees AS
M? length ‘ ; : 5 t ; : 3 . : : : ‘ , : ‘ 24,7
Me breadth . : : é : ; : i : : : é ' : s Wee
Interorbital width . : : : : : ; : : : : d : 3657
Postorbital width . 3 ‘ : : ; : : : p ; : : , Sisk
Zygomatic width . : ; : ; : : : : , 3 : ; : : 91,0
Mastoid width : : : t ; ‘ : : ‘ ; : : 2 : 73,8
Palate width at M? : : i : : 5 : : 5 2653
Horizontal diameter of arbi : ; : : : : : : : : , é 24.3
Vertical diameter of orbit . : : ‘ : : : ; A ‘ : i ; 21,0
Ventral margin of orbit to M® alveolar margin : , ' : ; iS 7/

in places, particularly the posterior part of the braincase. Distinctive features,
some of which are characteristic of other pliohyracines, include closed orbits,
naso-maxillary fossae which lead ventrally and posteriorly into antorbital
foramina situated immediately above the infraorbital foramina, a dental
formula of 3.1.4.3 with the I? to M® series closed, a premolariform C and an
elongated M? with an additional (third) lobe situated posteriorly. The preceding
comments on the upper teeth also apply to the lowers. The mandibles lack the
fossae and fenestrae found in some other Tertiary hyracoids.

Apart from its smaller size, the Arrisdrift species is most readily dis-
tinguished from later pliohyracines by its less elevated orbits and narrower skull.

ORDER PROBOSCIDEA
Family Gomphotheriidae

Gen. et sp. indet.

Four largely intact molars and a premolar (e.g. AD 252, AD 257) belong
to at least three individuals of an unidentified gomphothere (Fig. 7). They
resemble, and may be conspecific with, specimens from Maboko in Kenya
which date back about 16 m.y.

This material was originally identified by MacInnes (1942) as Trilophodon
angustidens kisumuensis. Arambourg (1945) believed that two taxa were repre-
sented and named a new genus and species, Protanancus macinnesi, to accommo-
date some specimens. Subsequently Tobien (1973) suggested that the Maboko
material belongs to a Platybelodon (P. kisumuensis), while Maglio (1974)
referred it to Gomphotherium cf. angustidens. Recently Tassy (1977) identified
it with Choerolophodon (C. kisumuensis).

This diversity of opinion is an indication of the difficulties which exist in
identifying fragmentary proboscidean remains, and it was decided to withhold

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hi l nn Iti a i Uy Man ee ieee a i

Fig. 7. Occlusal view of Gomphotheriidae third molar (AD 257) from Arrisdrift.

even a tentative identification of the Arrisdrift species. It is, however, unlikely
to be a Platybelodon since none of the numerous tusk fragments from Arrisdrift
are of the Platybelodon type.

Even though the material is unclassified, it is important since the molars
are more advanced than those of early Miocene gomphotheres elsewhere and
they are one of the elements in the assemblage which suggest a late ‘Burdigalian’
or post-‘Burdigalian’ age for the fauna.

Family Deinotheriidae

Prodeinotherium hobleyi (Andrews, 1911)

The Arrisdrift deinothere is represented by three cheekteeth which have
been described by Harris (1977).

Other Proboscidean Material

In addition to the cheekteeth already mentioned, there are many tusk
fragments and a few postcranial bones which evidently belong to either the
gomphothere or the deinothere. They have yet to be studied.

ORDER PERISSODACTYLA
Family Rhinocerotidae

Dicerorhinus sp.

At least three individuals of a rhinoceros are represented by several isolated
cheekteeth (e.g. AD 635, AD 827) and postcranial bones (e.g. AD 251, AD 601),
the latter being mainly elements of the pes.

The metatarsals which are known are relatively long compared with those
of the living Diceros bicornis and Ceratotherium simum, which suggests that

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA

IAAL

Fig. 8. Occlusal and buccal views of Dicerorhinus M? (AD 339)
from Arrisdrift.

21

iD) ANNALS OF THE SOUTH AFRICAN MUSEUM

the species concerned was either an Aceratherium or a Dicerorhinus (see Hooijer
1966) (Table 5). The cheekteeth of African Miocene representatives of these
genera may be difficult or impossible to distinguish (Hooijer 1966, 1968a),
but two M?’s from Arrisdrift (AD 339, AD 1103) resemble those of Dicerorhinus
rather than Aceratherium in having metacone bulges and unconstricted proto-
cones (Fig. 8). These characteristics, together with the elongated metatarsals,
distinguish the Arrisdrift species from other recorded African Miocene
rhinoceroses, namely, Paradiceros, Brachypotherium and Chilotheridium
(Hooter 1966, 19685, 1971).

Several Miocene species of Dicerorhinus have been recorded in Eurasia
and Africa (Hooyer 1966), including D. leakeyi from the east African early
Miocene. The teeth of the Arrisdrift species are larger than those of D. leakeyi
and in this respect resemble the European middle Miocene D. schleiermacheri
from Eppelsheim (Table 6). The Arrisdrift metatarsals are longer than those of
D. leakeyi, which are themselves ‘remarkable for their length’ (Hooijer 1966:
178), and although the metatarsals of D. schleiermacheri are not known,
Hooijer believed that they probably ‘exceeded those of D. leakeyi in length’.
Once again a similarity between the Arrisdrift and Eppelsheim species is
indicated.

The Arrisdrift species may also be more advanced than D. Jeakeyi in having
a less prominent metacone bulge, but there is doubt as to how much significance
should be attached to variations in this feature (Hooijer 1966: 128).

The available rhinoceros material from Arrisdrift is probably inadequate
for identifying the species concerned, but it does suggest one which was not
conspecific with the early Miocene D. leakeyi and which was perhaps closer
to the middle Miocene D. schleiermacheri in an evolutionary sense.

TABLE 5
Dimensions of Dicerorhinus metatarsals from Arrisdrift.

AD251 AD249 AD253
Mt II Mt III Mt IV

Median length : : : : ; E ; ‘ : a 190 170
Proximal width . p : : ; : : 33 61 43
Proximal antero-posterior diameter ; ; : : 5 48 = —
Middle width . ‘ : : : : : ; 30 50 —
Middle antero-posterior dhanieier . : f : ‘ DS 26 —
Ratio middle width/length . : ‘ 0,17 0,26 —

All measurements approximte owing to condition of specimens.

TABLE 6
Dimensions of Dicerorhinus M*’s from Arrisdrift.

AD339 AD1103

Antero-posterior diameter . 5 ' , ‘ 5 j ? : 55,0 54,3
Transverse diameter . ! ‘ , 3 : ; : 4 5 61,0 54,1
Length of outer surface , ’ : P ; ; : , 65,9 66,3

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 23

ORDER ARTIODACTYLA
Family Suidae

Gen. et sp. indet.

A mandible fragment with two cheekteeth (AD 631) belongs to a small
suid whose identity has yet to be determined.

Family Suidae

Lopholistriodon moruoroti Wilkinson, 1976
Several specimens belong to a small listriodont pig. They include a maxilla
fragment with P* to M? (AD 136) and two isolated M,;’s (AD 135, AD 636)
(Fig..9, Table 7).

TABLE 7
Dimensions of Lopholistriodon moruoroti teeth from Arrisdrift.
Pe M! M? M? M;
] b ] b ] b ] b ] b
AD 136... ee Op dine, S83 9) Ca ae 103" 10:4 A Ost — —
1D 1135) alo — — — as —— — — 14,0 7,9
ADCs — ~~ = —— _ — — 24 orl

In an unpublished thesis, Wilkinson (1972) described and named a new
species of pig, Xenochoerus ? moruoroti, from Moruorot Hill in Kenya. He
subsequently referred it to the genus Lopholistriodon Pickford & Wilkinson,
1975 (Wilkinson 1976). The teeth of the Arrisdrift listriodont are virtually
indistinguishable from those of the Moruorot L. moruoroti and it is identified
accordingly.

According to Pickford & Wilkinson (1975) this species is present at
Moruorot (17 m.y.), Maboko (16 m.y.) and Muruyur (13 m.y.), so its presence
at Arrisdrift is taken as a further indication that this fauna is late ‘Burdigalian’
or post-‘Burdigalian’ in age.

Family Tragulidae

Dorcatherium cf. pigotti Whitworth, 1958

Two mandible fragments (AD 104, AD 262) and a few postcranial bones
belong to a small tragulid. The teeth are morphologically indistinguishable
from those of Dorcatherium and, of the African species of this genus, they are
closest in size to those of D. pigotti of the east African early Miocene (Whit-
worth 1958) (Table 8).

TABLE 8
Dimensions of Dorcatherium teeth from Arrisdrift.
P, P; M, M,

AD104 : : iy OL 24 1.95 23,0 — — --- —
AD262 : = — — — oS Mee) 9.1 6,2

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Occlusal and buccal views of Lopholistriodon moruoroti maxilla (AD 136) from
Arrisdrift.

The Arrisdrift specimens are only tentatively identified with this species
since the grounds for distinguishing poorly represented fossil tragulids are
limited. There is little variation in tooth morphology and there has been a
tendency to name distinct species in Europe, Asia and Africa on the basis of
size differences. It is by no means certain that similarly sized species on the
different continents represent different species.

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 25

Family Palaeomerycidae (senso Hamilton 1973)

Climacoceras sp. nov.

The Arrisdrift ruminant assemblage includes several fragments of antler-
like frontal appendages (ossicones) (e.g. AD 130, AD 132) (Fig. 10). The
beams of the ‘antlers’ are straight, transversely compressed, and have small
_ knobs situated at irregular intervals both anteriorly and posteriorly. They
evidently also carried some small tines, several detached specimens having
; been discovered (e.g. AD 129, AD 785). The tines are circular in cross-section
| and slightly curved. AD 648 indicates that there was bifurcation and greater
flattening of the ‘antlers’ distally. AD 483 apparently represents the proximal
part of an ‘antler’ and since it lacks a burr, the ‘antlers’ were evidently not
| deciduous. They were, therefore, not true antlers of the kind which characterize
} the Cervidae.

vat

WNIHI

9

ii

UH lti

6

mu

1

st

mu

if

£

iii

1

Fig. 10. Lateral view of Climacoceras ‘antler’ fragments from Arrisdrift: AD 648 (+ AD 763)—
distal end showing bifurcation (left); AD 130—beam fragment showing knobs (centre);
AD 129—tine (right).

The complete ‘antlers’ must have resembled those of Climacoceras africanus
from Maboko (MaclInnes 1936). Climacoceras is also present at Fort Ternan
(Gentry 1970). The Arrisdrift specimens are distinguished from east African
specimens by their larger size, the difference being of the order of 20 per cent
according to A. W. Gentry (pers. comm.). The beam circumference of specimens
from Maboko varies from 54 to 92 mm (MaclInnes 1936), while the correspond-

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

ing figures for Arrisdrift specimens are 85 to 110 mm. The longest of the Maboko
tines recorded by MacInnes measures 48 mm, whereas two of the Arrisdrift
specimens (AD 129, AD 1177) are about 60 mm long and both are incomplete.

The larger size of the Arrisdrift specimens is taken to indicate that they
belong to a hitherto unrecorded species of Climacoceras, although the morpho-
logical similarity to specimens from Maboko suggests that the Arrisdrift
species was in a comparable evolutionary state to C. africanus.

Many mandible and maxilla fragments, isolated teeth and postcranial
bones probably belong to this species. It is, however, not certain that all the
material provisionally assigned to the Climacoceras belongs only to this species.
A comparison between two of the more complete mandibles (AD 261, AD 612;
Fig. 11, Table 9) revealed differences which may be taxonomically significant.
For example, AD 612 has slightly larger teeth, an appreciably deeper mandibular
corpus, larger basal pillars on the molars and a less expanded P, metaconid.
In addition, the lingual surface of the M, third lobe is directly connected to the
second and is flanked by a small, more or less circular enamel island which
evidently corresponds to the central cavities of the first and second lobes. By
contrast, the M, third lobe of AD 261 is transversely compressed dorsally and
is connected lingually to the second lobe by a deeply indented loop in the
enamel. It also lacks a ‘central cavity’.

TABLE 9

|

Dimensions of lower teeth and mandibles AD 261 and AD 612, tentatively assigned to
Climacoceras from Arrisdrift.

P, P, M, -M, Ms;
l b l b l b b ] b
AD 261 — — 14,0 7,6 (S33) Og (PNG 12 Sia 95
A DiGi 2 ae ael Ss) ciao Wo SINE te) c.17,0 — 21.53%: CO aseee oS wt 0, 7,
AD: 261 ~~ AD Gl2
Depth of mandible below P, ; f 25,0 29,0
Breadth of mandible below P, . : al 14,5
Depth of mandible below M,_ . ; 28,5 Sie
Breadth of mandible below M; . : 13,0 15,8

Sorting the remaining mandible fragments on the basis of these criteria
was not entirely satisfactory owing to the poor condition of some specimens and
because in some instances ‘characteristics’ of one type occurred in conjunction
with ‘characteristics’ of the second. Nevertheless, AD 259, AD 263, AD 269
and AD 270 are apparently of the AD 612 type and represent at least 4 indi-
viduals, while AD 271, AD 272, AD 346, AD 356 and AD 621 appear to be of
the AD 261 type and represent 6 individuals.

The two sets of specimens are otherwise similar and amongst the shared
characteristics are a giraffoid-like orientation of the diastema region relative
to the cheektooth row, low-crowned cheekteeth, absence of P, and simple,
crescentic central cavities on the lower molars, the posterior one opening

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA DL]

lingually in early wear. Although several symphyseal teeth are known, they
do not include bilobed giraffoid-like canines.
The less numerous maxillae and upper teeth have not been closely examined.
Gentry (pers. comm.) has found that although the large ruminant teeth and

A

INNNUnI a

ET

yee

inna

Fig. 11. Occlusal and buccal views of ? Climacoceras mandibles, AD 261 (above) and AD 612
(below), from Arrisdrift.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

dentitions from Arrisdrift share several characters with the early Miocene
Propalaeoryx from Elisabethfeld and the Fort Ternan Climacoceras, they are
in some respects intermediate between the two. For example, the Elisabethfeld
Propalaeoryx is more primitive in retaining P,. On the other hand, at least some
of the Arrisdrift M,’s (the AD 612 type) have ‘central cavities’ on the third
lobes, whereas the Fort Ternan Climacoceras M,’s lack this feature. In addition,
the Arrisdrift teeth are probably less high-crowned than those of the Fort
Ternan Climacoceras, while the metastylids are developed to a degree inter-
mediate between the Elisabethfeld and Fort Ternan species.

Since the Maboko Climacoceras predates the Fort Ternan species and is
younger than the Elisabethfeld Propalaeoryx, a temporal link with the Arrisdrift
species is suggested. Unfortunately the only described teeth assigned to the
Maboko Climacoceras are three lower molars (MacInnes 1936), so the basis
for comparisons with Arrisdrift specimens is limited. The Maboko teeth are,
however, similar to the AD 261 type in size and some morphological characters,
including reduced or absent basal pillars and dorsally compressed third lobe
of Msg.

There are several complete specimens amongst the postcranial bones
tentatively assigned to the Climacoceras. They include a tibia (AD 1100),
several radii (e.g. AD 494, AD 562) and metapodials (e.g. AD 198, AD 199),
as well as elements of the manus and pes. The long bones are slender and
elongated compared with those of living bovids and cervids of similar overall
size (e.g. Damaliscus dorcas, Cervus unicolor). The housing of the extensor
tendon of the distal metatarsals shows the bovid and giraffid rather than the
cervid condition (see Whitworth 1958: 23).

Family Bovidae
Gen. et sp. indet.

Several mandible fragments (e.g. AD 103, AD 106) and postcranial bones
belong to a small ruminant (Fig. 12, Table 10). Morphologically the teeth are
perhaps closest to those of the somewhat larger Walangania africanus
(= Palaeomeryx africanus + Walangania gracilis (see Hamilton 1973)) of the
east African early Miocene. W. africanus was once thought to be a bovid, but

TABLE 10
Dimensions of the lower cheekteeth and mandibles of the Bovidae indet. from Arrisdrift.
E Ee M, M, M;
l b ] b ] b ] b l b
AD OS =: & eS 4,2 8,3 4,8 — — 904 “Gil 13.0) “6,5
AD 106 — — 8,1 4,5 9,1 6,3 9,7 6,6 = =
AD 105 AD 106
Depth of mandible below P, ! E eles 14,3
Breadth of mandible below P, . : ; 6,3 6,1
Depth of mandible below M, . ‘ ; ISS 18,7

Breadth of mandible below M3. : : 7,6 71,9

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 29

HL

Fig. 12. Occlusal and buccal views of Bovidae mandible (AD 105) from Arrisdrift.

'

Gentry (pers. comm.) now believes it could be congeneric with the primitive
ruminant Dremotherium from the European late Oligocene to early Miocene.

Gentry (pers. comm.) reports on the Arrisdrift material as follows: “The
smaller metastylids of the lower molars are more advanced towards bovids
than those of Walangania. Similarly, the weaker anterior ribs and the better
developed closure of the central cavities. The Maboko bovid fragment with
M, (see Whitworth 1958: 25, fig. 10a—c) appears to agree with the Arrisdrift
species except in not being smaller than Walangania: its metastylid is weak,
the anterior rib is not localized, and the central cavities are more enclosed within
the tooth.’

He concludes that the Arrisdrift species ‘is a bovid, although finding a
horn core is needed to be conclusive’.

Other Ruminants

The possibility of a second taxon being represented amongst the material
assigned to the Climacoceras has already been mentioned.

In addition, there are several ruminant postcranial bones which are too
small to belong to the Climacoceras, but far too large to belong to the bovid.
They include a distal humerus (AD 39), proximal metacarpals (AD 764, AD 964)
and a first phalanx (AD 895). The ‘Pecora gen. & sp. indet.’ on the accom-
paying faunal list (Table 1) refers to this material

ORDER LAGOMORPHA
Family Ochotonidae

Kenyalagomys sp. nov.

A mandible fragment with M, and M, (AD 813) (Fig. 13) and an isolated
P? (AD 1185) belong to a small ochotonid which resembles species previously

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Occlusal and buccal views of Kenyalagomys
mandible (AD 813) from Arrisdrift.

recorded from early Miocene deposits in the Namib desert and east Africa.

The first African Miocene ochotonid to be described was Austrolagomys
inexpectatus from Elisabethfeld (Stromer 1924, 1926). Hopwood (1929) recorded
a second species from the same region, namely A. simpsoni. Subsequently
MacInnes (1963) identified a second genus, Kenyalagomys, on the basis of
material from east Africa and named two species, K. rusingae and K. minor.

MacInnes made no reference to Hopwood’s species, but amongst the
characters used to distinguish Kenyalagomys from Austrolagomys (MacInnes
1953: 20-21) are two of the three characters which distinguish A. simpsoni
from A. inexpectatus (Hopwood 1929: 2). They are a deep external fold on P3
and a marked median angulation (rib) on the posterior walls of the anterior
lobes of P, to Mg. It follows that if the generic distinction is justified, and it is

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 3]

here assumed that it is, then A. simpsoni must be referred instead to
Kenyalagomys.

Judged on the basis of size, neither of the east African species is a synonym
of K. simpsoni. K. rusingae is larger and, although K. minor is similar in overall
size, it differs in having a smaller P,; and larger molars.

Another species of Kenyalagomys, K. mellalensis, was recently recorded
from the middle Miocene (c. 14 m.y.) of Beni Mellal in Morocco (Janvier &
De Muizen 1976).

The molars of the Arrisdrift mandible (AD 813) are almost identical in
size to the corresponding teeth of the K. simpsoni holotype, but the posterior
walls of the anterior lobes lack the prominent ribs which characterize Kenyala-
gomys. There is, however, a faint indication of such ribs and the teeth of AD 813
are closer to the condition in Kenyalagomys than that in Austrolagomys, where
the posterior walls are smoothly curved.

The Arrisdrift P? (AD 1185), which measures 1,6 by 3,2 mm, is similar
in size to that of K. minor, it is smaller than that of K. rusingae and longer, but
narrower than that of K. mellalensis. It also resembles K. minor in having the
postero-external corner pointed rather than rounded as in both A. inexpectatus
and K. rusingae. K. mellalensis is in an intermediate position in this respect.
AD 1185 differs from K. minor in having the posterior border more or less
straight rather than convex and in this respect resembles K. rusingae and
K. mellalensis. Visible on the occlusal surface is a deeply indented enamel fold
which resembles corresponding features in the east and north African species
of Kenyalagomys, but which is absent in Austrolagomys.

The Arrisdrift ochotonid is here regarded as a previously unrecorded
species of Kenyalagomys, whose closest relatives are K. minor and K. simpsoni.
If there is a phyletic relationship between the latter two species and the one
from Arrisdrift, then the reduced median ribs of the anterior lobes of the
lower molars of AD 813 may be interpreted as being either in an incipient or
in a vestigial state. The latter alternative is more likely in view of the probable
younger age of the Arrisdrift species relative to the Rusinga K. minor and its
apparent contemporary from the Namib, K. simpsoni. Since K. mellalensis is
more likely to be related to K. rusingae than the smaller African ochotonids,
its evolutionary state relative to that of the Arrisdrift species is not determinable.

To sum up, the Arrisdrift ochotonid is apparently a new species of Kenyala-
gomys which is more advanced than the smaller species already recorded from
the early Miocene of Africa.

ORDER RODENTIA
? Family Bathyergidae
? Bathyergoides sp.

Two incomplete lower incisors (AD 141, AD 1024) belong to a large
rodent, possibly a bathyergid. They are tentatively attributed to Bathyergoides,

By ANNALS OF THE SOUTH AFRICAN MUSEUM

a genus recorded from the early Miocene of the Namib (Stromer 1926) and east
Africa (Lavocat 1973).

Family Thryonomyidae
Paraphiomys pigotti Andrews, 1914

Two incomplete mandibles (AD 629, AD 1049) belong to a rodent which
appears indistinguishable from that described by Stromer (1922, 1926) as
Neosciuromys africanus. This taxon has since been recognized as a junior
synonym of Paraphiomys pigotti by Lavocat (1973). P. pigotti is one of the more
commonly occurring rodents in deposits of early Miocene age in both east
Africa and the Namib desert. The Arrisdrift P. pigotti evidently postdates all
previous records of this species (see below).

Other Rodents

The Arrisdrift assemblage includes many isolated rodent incisors and,
judged on the basis of size, they represent at least two species in addition to
those mentioned above.

AGE OF THE OCCURRENCE

The fauna from Pit 2/AD 8 at Arrisdrift undoubtedly dates from the
Miocene and it has been suggested elsewhere (South African Journal of Science
1976; Corvinus & Hendey 1978) that it falls within the age limits of 12 to 18
m.y. The present study has tended to confirm the older Hmit, but it has also
suggested that the 12 m.y. limit is too young.

At present the fauna can be dated only in a relative sense by comparing
individual taxa with more securely dated ones elsewhere. The non-mammalian
vertebrates cannot yet be used in this way and the comments which follow are
confined to the mammals.

On the basis of previous records only two of the twenty-two mammalian
species suggest an early Miocene date for the fauna. They are Myohyrax cf.
oswaldi and Paraphiomys pigotti, which in east Africa are recorded from deposits
ranging in age from 18 to 22 m.y. Since little has been published on the midde
Miocene small mammals of east Africa, it cannot yet be assumed that these
taxa did become extinct there 18 m.y. ago. In addition, since there is some
evidence of differences in the patterns of mammalian evolution in east and
southern Africa during the earlier part of the Miocene (see below), it is possible
that taxa such as M. oswaldi and P. pigotti survived longer in southern Africa.

Two other species, the large canoid and ? Ischyrictis sp., may also be
indicative of an early Miocene date, but in both instances doubts about identifi-
cation render them unreliable for relative dating purposes.

Five of the twenty-two species suggest a middle Miocene date, that is,
late “Burdigalian’ at the earliest, but more probably ‘Vindobonian’ or even

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 33

‘Maremmian’ (senso Berggren & Van Couvering 1974). They are Amphicyon
cf. steinheimensis, Gomphotheiidae indet., Dicerorhinus sp., Lopholistriodon
moruoroti and Climacoceras sp. nov. In addition, there are another three
species which are, or probably are, more advanced than previously recorded
African “Burdigalian’ species and would be consistent with a middle Miocene
date. They are Prohyrax sp. nov., Bovidae indet. and Kenyalagomys sp. nov.
These eight species are regarded as the most significant for dating purposes.

Four of the twenty-two species would be consistent with any age from
‘Burdigalian’ to ‘Vallesian’. They are ?Hemicyoninae indet., ? Metailurus sp.,
Prodeinotherium hobleyi and Dorcatherium cf. pigotti.

The remaining six species provide no evidence of age. They are the
unidentified carnivore, suid, pecoran and rodents.

Negative evidence also gives some indication of the probable age of the
fauna. For example, the absence of equids suggests it is pre-‘Vallesian’ (see
Hooijer 1975), while the absence of positively identifiable bovids and palaeo-
tragines suggests that it predates the 14 m.y. old Fort Ternan fauna (see Gentry
1970; Churcher 1970).

There is, in fact, no secure evidence that the Arrisdrift fauna is as young as
that from Fort Ternan, while there is good evidence that it postdates the
‘Rusinga-like’ faunas of east Africa. Consequently, the likely age limits may
be reduced to between 14 and 18 m.y., with the median estimate being about
16 m.y. This is the age of the Maboko fauna and, although it has yet to be fully
described, it includes at least six species which are conspecific with, or closely
related to species from Arrisdrift (Table 11). They are Choerolophodon
kisumuensis, Prodeinotherium hobleyi, Lopholistriedon moruoroti, Dorcatherium
pigotti, Climacoceras africanus and the unidentified bovid. This suggests that
the two faunas are, indeed, of the same order of age. On the other hand, the
deinothere and tragulid are of little use as precise age indicators, while the two
faunas do differ in certain respects. Primates and creodonts are absent or

TABLE 11

The mammals from Maboko, Kenya, and their counterparts from
Arrisdrift, South West Africa.

MABOKO* ARRISDRIFT
Primates _
Paracynohyaeonodon leakeyi Various Carnivora
Megalohyrax championi Prohyrax sp. nov.
Choerolophodon kisumuensis Gomphotheriidae indet.
Prodeinotherium hobleyi Prodeinotherium hobleyi
Aceratherium acutirostratum Dicerorhinus sp.
Lopholistriodon moruoroti Lopholistriodon moruoroti
Brachyodus aequitorialis —

Dorcatherium spp., including D. pigotti D. cf. pigotti
Climacoceras africanus Climacoceras sp. nov.
Bovidae indet. Bovidae indet.

* Bishop (1967); Hooijer (1968a); Pickford & Wilkinson (1975);
Tassy (1977); Van Valen (1967); Whitworth (1958).

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

apparently absent, at Arrisdrift and the hyracoids from the two occurrences
are at least generically distinct. These differences will be discussed again later,
but they indicate that either the two faunas were not exactly contemporaneous,
or that there were regional differences between contemporary east and southern
African faunas at that time.

The final word on the age of the Arrisdrift fauna has yet to come, but
available evidence suggests that it is early middle Miocene, with an inferred
date of about 16 m.y. before present.

PALAEOENVIRONMENT

The Arrisdrift fauna dates from a zoogeographically important period.
Andrews & Van Couvering (1975: 85-87) have discussed the ‘abrupt changes’
which occurred in the faunas of east Africa between 14 and 18 m.y. ago and
ascribe them to the development of a land bridge between Africa and Eurasia
and the consequent immigration of new taxa. According to Berggren &
Van Couvering (1974) the land bridge in question resulted from the closure
of the eastern Tethys between 18 and 20 m.y. ago. Faunal changes must also
have been experienced in southern Africa during this period, although it cannot
be assumed that they were coincident with, or that they were an exact parallel
of those in east Africa. There is, in fact, some evidence that the situation in
southern Africa did differ from that in east Africa.

Andrews & Van Couvering (1975) pointed out that during the early
Miocene the dominant hyracoids in east Africa were geniohyids and that they
were replaced in the middle Miocene (post-Fort Ternan) by procaviids. In the
Namib region the only recorded hyrax is the procaviid Prohyrax, which was
contemporary with geniohyids in east Africa (up to and including Maboko).
It follows that the procaviids are likely to have had southern Africa as their
centre of origin and that they moved into east Africa at a time when this region
was also receiving Eurasian immigrants. Even if there are changes in the classifi-
cation of the hyracoid taxa concerned, the substance of the preceding theory
remains the same since the east and southern African forms clearly belong to
different lineages, and it is the southern African one (a pliohyracine) which
had descendants in east Africa (and elsewhere) in post-early Miocene times.

Also relevant here is MacInnes’s (1957) opinion that the southern African
Parapedetes and east African Megapedetes were contemporary representatives
of different lineages since it, too, suggests that there was some independent
evolution of related taxa in the two regions. Other of the Namib rodents, as
well as the ochotonid Austrolagomys, may also have been southern endemics.

In comparing faunas, it may be unwise to emphasize the absence of certain
taxa since this may be due to sampling deficiencies or even incorrect identifi-
cation of specimens. While the apparent absence of, for example, primates at
Arrisdrift might be due to such factors, if their absence is real then it must be
palaeoenvironmentally significant. The absence or great rarity of primates at

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA 35

another southern African late Tertiary locality, namely, Langebaanweg, has
already been mentioned elsewhere (Hendey 1976: 234), while this group is
also not recorded from the other Miocene localities in the Namib.

By contrast, Andrews & Van Couvering (1975: 86) noted that during the
early Miocene of east Africa ‘there was a notable proliferation of hominoid
primates, seven species in all; and there were at least five species of prosimians’.
Primates became much less common in east Africa during the middie Miocene,
although during the 14 to 18 m.y. transition period monkeys were ‘common
at Maboko’, whereas ‘there is little evidence for their presence in the Early
Miocene environments’ (Andrews & Van Couvering 1975: 93). The primates are
thus one group which reflect the ‘abrupt changes’ referred to earlier, but in this
instance the change is not manifested in the southern African record.

The apparent absence of primates in southern Africa during the earlier
part of the Miocene is made even more remarkable by the fact that this group
features in the ‘Burdigalian’ faunal interchange between east Africa and southern
Eurasia. Thus, while a northward movement of primates from east Africa is
documented, there is no record of a corresponding movement to the south.

To sum up, there is some evidence which suggests that early in the Miocene
faunal interchange between east and southern Africa was inhibited and that at
least some related taxa evolved independently of one another in the two regions.
This indicates the existence of an environmental barrier between the two
regions and it is most likely to have been comprised of the extensive river system
of central Africa, together with the Rift Valley lakes (see Kortlandt 1972).
Even in their present form the Congo and Zambezi river systems, especially
in the region of the Congo/Zambezi divide, make up a broad and almost
continuous area of channels and marshes between east and south-west Africa.
Tectonic disturbances in central Africa during the mid-Tertiary may well have
complicated the headwater drainage patterns of these rivers and so have created
an even more effective barrier to limit the crossing of at least certain mammals.

Another possible barrier may have been that of an intervening arid and
semi-arid region. A more extended form of the present Kalahari desert would
have effectively separated east and south-west Africa from one another. This
alternative is perhaps less likely since the present aridity of the south-western
parts of Africa was apparently initiated only in the very late Tertiary
(A. J. Tankard & J. Rogers, unpublished manuscript). Prior to this the climate
and vegetation of Africa may have been of a more uniform nature.

The preceding observations suggest that in assessing the character and
composition of southern African Miocene faunas, allowance must be made
for deviations from the better documented east African pattern because
geography is a complicating factor, the full implications of which have yet to
be established.

In this connection the apparent absence of creodonts at Arrisdrift may
also be significant. Creodonts predominate in the carnivore faunas of the 18
to 22 m.y. period in east Africa and the only identified carnivore from con-

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

temporary occurrences in the Namib is also a member of this group. Similarly
the only recorded carnivore from Maboko is a creodont, while at Fort Ternan
creodonts are common if not predominant. In other words, creodonts were
an important element in the ‘Aquitanian’ to ‘Vindobonian’ faunas of east
Africa. Consequently, it is to be expected that they would not only be present
at Arrisdrift but would be more commonly represented than fissiped carnivores.

Since carnivores are less restricted by environmental factors than herbi-
vores, the dispersal of immigrant taxa may well have been rapid in spite of
barriers which impeded the movement of, for example, primates. Should it
be established that creodonts are, indeed, rare or absent at Arrisdrift, the
situation could be explained in only one of two ways.

Firstly, southern Africa was an important centre of fissiped evolution
and they superseded creodonts in this region before the same happened in east
Africa. This possibility can be dismissed in view of what is known of fissiped
origins and evolution (Savage 1977) and since it would require complete
isolation of southern Africa from east Africa during the early Miocene.

The second possibility is that Arrisdrift is younger than was indicated
previously and dates from a period when the creodonts had been largely or
completely replaced by fissipeds. The carnivores would then be the only obvious
‘advanced’ element in the Arrisdrift fauna, while those taxa which suggest
ac. 16 m.y. date would be ‘primitive’ forms which survived longer in the Namib
region than in east Africa. This interpretation would require, for example,
that early bovids and giraffids such as those found at Fort Ternan had been
prevented from spreading southwards by the hypothetical zoogeographic
barrier, whereas immigrant fissipeds had already surmounted it and become
established in southern Africa by the time that the Arrisdrift fossils were being
deposited.

Perhaps the only firm conclusion to be drawn from the preceding discussion
is that the present state of knowledge of southern African Miocene faunas
leaves much to be desired.

Some information on the nature of the environment in the immediate
vicinity of Arrisdrift at the time that the fossiliferous deposits were laid down is
suggested by both the fossils and the deposits themselves.

There is no doubt that the fossils accumulated in a river channel, the verte-
brate remains simply being an additional element in the coarse sediment
fraction of a fluvial gravel. The fact that a Prohyrax skull and other delicate
fossils were recovered from the deposits indicates that they at least could not
have been transported far in what was evidently a turbulent channel. Even
those specimens which are abraded are not seriously damaged. Thus, most of
the fossils must represent the remains of animals which lived in the immediate
vicinity or a little further upstream.

The only invertebrate in the fossil assemblage, a serpulid polychaete (cf.
Mercierella sp.), is a typically estuarine form (B. Kensley, pers. comm.) and its
presence suggests that at the time of deposition the coastline, which is at present

MIOCENE VERTEBRATES FROM ARRISDRIFT, SOUTH WEST AFRICA BY)

about 30 km away, was much closer. This, together with the fact that the
fossiliferous deposits are about 50 m above sea-level, suggests that deposition
took place during a period of relatively high sea-level. There is evidence for a
world-wide marine transgression during the Miocene, between 10 and 20 m.y.
ago (Flemming & Roberts 1973) and presumably at least part of the Arrisdrift
terrace sequence can be correlated with this event. Arrisdrift is situated in a
hilly area on the last meander of the Orange River before it reaches the flat
and low-lying coastal section of its valley, which is likely to have been inundated
during the transgression. The mouth of the river was, therefore, probably
only a few kilometres west of Arrisdrift.

The area around Arrisdrift is now very arid, with Alexander Bay at the
mouth of the Orange River having a mean annual rainfall of less than 50 mm
(Dept. of Transport 1965). The Orange River is, however, a large perennial
river and is flanked by a narrow belt of bushes and trees, although the vege-
tation becomes ephemeral a short distance away. There is little or no soil cover in
the area and aeolian sands and bedrock exposures are ubiquitous. Although
the climate is ameliorated by the proximity of the cold Atlantic Ocean, it is
nevertheless an inhospitable area capable of supporting only sparse popu-
lations of a relatively small number of mammalian species (see Shortridge 1934).

The environment at the time that the fossils were deposited must have been
very different. Although poorly preserved plant remains occur in the Pit
2/AD 8 deposits, no direct information on the nature of the vegetation is yet
available. The large mammals, particularly the two proboscideans, suggest a
densely vegetated and probably wooded environment. Harris (1975) has
suggested that both Prodeinotherium and gomphotheres preferred such a habitat.
The low-crowned teeth of the rhinoceros suggest that it, too, was a browser,
while the long limbs and low-crowned teeth of the Climacoceras indicate that
it was not adapted to grazing. Both are likely to have been woodland species.

The smaller herbivores are probably indicative of a dense undergrowth
at least in the immediate vicinity of the river. Living tragulids are forest-dwelling
browsers and their Miocene ancestors, including the Arrisdrift Dorcatherium,
probably had a similar habitat preference. The same is likely to apply to the
bovid and two suids since they resemble the Dorcatherium in both size and
hypsodonty, while the Prohyrax probably occupied a similar habitat, but with
somewhat different vegetable-food preferences.

Since later pliohyracines were aquatic or amphibious animals, early forms
such as the Arrisdrift Prohyrax may already have developed a preference for
life in water-side situations. The fact that it is the most commonly occurring
vertebrate in the assemblage supports the theory that it was a riparian species
since remains of such animals are more likely to be incorporated in fluvial
deposits than those of other terrestrial species. In this connection it is probably
significant that the other commonly occurring vertebrate at Arrisdrift is a
crocodile, which undoubtedly is a riparian species.

The picture which emerges is of a forested riverine setting, probably with

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

dense undergrowth adjacent to the river and with the sea no more than a few
kilometres away. An essentially similar environment was suggested for those
areas further north in the Namib where the early Miocene vertebrates were
discovered (Stromer 1926; Hopwood 1929). In the case of these occurrences
the contrast to the modern environment is even more striking since there are
no rivers in the area today. There is, however, ample evidence of their presence
during the Miocene.

The Arrisdrift fossils, together with the older ones from the Liideritz—
Bogenfels area, provide some of the evidence which supports the theory that
the present Namib desert is relatively young (A. J. Tankard & J. Rogers,
unpublished manuscript). Whether or not a desert existed in the area in pre-
Miocene times is still a matter of dispute, but the earlier part of the Miocene
was evidently a period of relatively high rainfall and more luxuriant vegetation.

Andrews & Van Couvering (1975) believed that during this period a belt
of lowland forest stretched across equatorial Africa and was flanked on either
side by woodlands. The evidence from Arrisdrift and the other occurrences
suggests that these woodlands extended at least as far south as the Orange River.

ACKNOWLEDGEMENTS

I am indebted to the management and staff of Consolidated Diamond
Mines of South West Africa (Pty) Ltd at Oranjemund who were involved in
the collecting of the Arrisdrift fossils and their subsequent donation to the
South African Museum. Particular thanks are due to Dr G. Corvinus who
undertook the excavation of the material, Mr J. O. Richards (General Manager)
and Dr C. G. Stocken (Chief Geologist). A generous grant from the company
has partly covered the costs of preparing the fossils.

I am also indebted to those persons who have assisted the preliminary
study of the Arrisdrift fossils. They are: Drs A. W. Gentry (British Museum
(Natural History)), J. M. Harris (Kenya National Museum), B. Kensley (South
African Museum), R. G. Klein (University of Chicago), M. Pickford (University
of London) and R. J. G. Savage (University of Bristol). Dr Gentry was especially
helpful in sharing his extensive knowledge of ruminants; he is, however, not
responsible for any errors in interpretation of the Arrisdrift material.

The preparation of the fossils has been undertaken at the South African
Museum by Mrs K. Volman and Miss T. Salinger, while Mr N. Eden provided
the photographs which illustrate this report.

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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., Syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87. ;
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holeivpe
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus...’; *. .. in C. namacolus (Fig. 10)...’

(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 but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

Q. B. HENDEY

PRELIMINARY REPORT ON
THE MIOCENE VERTEBRATES FROM ARRISDRIFT,
| SOUTH WEST AFRICA

i ia \ ‘
VOLUME 76 PART 2 SEPTEMBER 1978 ISSN 0303-2515

.
‘ee 2
i =a

OF THE SOUTH AFRICAN
~~ MUSEUM

CAPE TOWN

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

FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. Zen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Canis (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. 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.

(continued inside back cover)

wrrveawwrtltenwrvr yy

Fig. 1. Guineafowl taxa recognized in this study.

A. A. meleagrides. B. A. niger. C. Guttera plumifera plumifera. D. G. p. schubotzi. E. G. pucherani

pucherani. F. G. p. verreauxi. G. G. p. sclateri. H. G. p. barbata. I. G. p. edouardi. J. Acryllium

vulturinum. K. Numida meleagris meleagris. L. N. m. sabyi. M. N. m. galeata. N. N. m. somaliensis.

O. N. m. marungensis. P. N. m. reichenowi. Q. N. m. mitrata. R. N. m. coronata.
S. N. m. damarensis.

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

Volume 76 Band
September 1978 September
Part 2 ~+=Weel

THE EVOLUTION OF GUINEA-FOWL
(GALLIFORMES, PHASIANIDAE, NUMIDINAE)
TAXONOMY, PHYLOGENY, SPECIATION AND

BIOGEOGRAPHY

By
T. M. CROWE

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE EVOLUTION OF GUINEA-FOWL
(GALLIFORMES, PHASIANIDAE, NUMIDINAE)
TAXONOMY, PHYLOGENY, SPECIATION AND BIOGEOGRAPHY

By
T. M. CROWE

FitzPatrick Institute, University of Cape Town

(With 53 figures, 11 tables and 2 appendices)
[MS. accepted 15 March 1978

ABSTRACT

Patterns of qualitative and quantitative character variation in 1 833 museum specimens
encompassing all taxa attributed to the Numidinae are analysed to produce an hypothetical
taxonomy and phylogeny for the subfamily. Quantitatively and ecologically defined genus,
species and subspecies concepts are applied in erecting the taxonomy. A cladistic approach,
based in postulated primitive-derived character sequences, is used in developing the phylogeny.
4 genera, 6 species and 16 subspecies are recognized. Cladistic events are linked to likely
causal geological and palaeoecological events to determine a possible evolutionary chronology.
Genera are thought to have arisen as a consequence of Miocene and Pliocene radiations;
species and subspecies as a result of Pleistocene divergence. An hypothetical map of African
avifaunal zones, based on evolutionary patterns found in guinea-fowl, is offered. Comparisons
of this map with other African avifaunal maps and with distribution maps of selected francolin
taxa suggest that biogeographic patterns found in guinea-fowl reflect broad patterns found
in many African birds. An hypothesis as to the causes of relatively high species richness in
francolins is offered.

CONTENTS
PAGE
Introduction ; : : : , 5 5 . 44
Methods ; ; : j : >
Material and characters ; : ; : oee'§45
Taxonomic philosophy : 3 : : a)
Taxonomic methodology . : : : . 30
Phylogenetic philosophy . : : : Fee)
Phylogenetic Semen . : : san ee
Speciation . . : : ; : i ae
Biogeography : : : : ee
Results, discussion and conclusions . ; : - —36
Taxonomy . : : : 2 ; : S436
Genera . : : : ‘ els
Species and subspecies : : : / “356
Taxonomic summary . : ; : 95
Phylogeny . Se etl:
Primitive and derived character ‘states Be ult
Phyletic analysis . : ; : . 120
Speciation . : Be ull
Origin and derivation of the Numidinae gort2
Evolution of genera. sf l2
Evolution of species and subspecies . a 12D
Biogeography : ; : é : i b 1 128
Results . ‘ ; : : : : 123
Discussion . ; : : : 5 a 18}
Conclusions . ; 5 5 : : ~ §25

43

Ann. S. Afr. Mus. 76 (2), 1978: 43-136, 53 figs, 11 tables, 2 appendices.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE
Synthesis ; : i : : : : ; » 1126
Acknowledgements. é : ; : L130
References . ; ; : . ; ; : . 130
Appendix 1 . ; : : : : : . = 138
Appendix 2 . : : ; : : ; «| h34

INTRODUCTION

There are many reasons why an understanding of the evolution of guinea-
fowl (Numidinae; Sibley & Ahlquist 1972) should provide an insight into
patterns of avian evolution and biogeography in Africa.

|. The Numidinae are endemic to Africa, and presumably evolved and/or
radiated there from a francolin-like ancestor (Ghigi 1936; Cracraft 1973;
Olson 1974).

2. At least one guinea-fowl species has become adapted to life in each
major terrestrial African biome outside of desert, Mediterranean vegetation
and montane forest (Crowe & Snow 1978).

3. Guinea-fowl are sedentary birds (Chapin 1932a; Priest 1933; Archer
& Godman 1937; Elgood et al. 1973), and thus should be more susceptible
to local selection pressures than would be more mobile species (Ehrlich &
Raven 1969).

4. The distributions of some guinea-fowl taxa do not correlate well with
present-day vegetation and topography (Chapin 1932a). Such anomalous
distributions of plants and animals can be used to infer environmental con-
ditions and the distributions of African biomes during the Tertiary and
Quaternary (Chapin 1932a; Moreau 1963, 1966; Roberts 1975; Hamilton
1974; Axelrod & Raven 1978).

5. The genetic basis of morphological variation in several guinea-fowl
species is relatively well understood (Ghigi 1936).

6. Perhaps most importantly, guinea-fowl species are relatively well
represented in museum collections and, accordingly, lend themselves to
quantitative analysis, which is desirable in formulating precise evolutionary
and biogeographic hypotheses (Mayr ef al. 1953; F. Vuilleumier 1975).

Thus, guinea-fowl provide a simple, characteristically African system
from which evolutionary and biogeographic hypotheses may be derived. The
aims of the present study are to:

1. re-examine and revise, if necessary, the rather confused taxonomy within
the subfamily (Table 1), using a repeatable, relatively objective quantitative
methodology;

2. produce a parsimonious phylogeny based on the analysis of shared derived
character states;

3. develop models of speciation, which are consistent with the phylogeny
developed herein and the likely past geological and climatic history of
Africa;

THE EVOLUTION OF GUINEA-FOWL 45

4. suggest tentative avifaunal subregions, provinces and districts for Africa,
based on analysis of the distributions of recognized guinea-fowl taxa.
The possible familial status of the subfamily (Wetmore 1960), the adaptive-
ness of inter- and intra-specific morphological variation in guinea-fowl, and
the predictive value of evolutionary and biogeographic models based on
patterns found in guinea-fowl, will be discussed briefly, if at all, herein. These
topics will be dealt with in greater detail in future papers.

METHODS

MATERIAL AND CHARACTERS

The author examined | 833 museum specimens, including all taxa attributed
to the subfamily in Table 1. All characters investigated (Appendices 1 and 2,
Figs 2-4) appear to have a genetic basis (Ghigi 1936), and have been discussed
in previous studies (e.g. Beddard 1898; Bannerman 1930; Chapin 1932a;
Ghigi 1936; Archer & Godman 1937; Jackson 1938; Boetticher 1954;
Mackworth-Praed & Grant 1952, 1962, 1970). Therefore, they need not be
described again in detail. Characters were chosen to reflect as much of the
phenotype as possible, while eliminating the necessity of close comparisons of
specimens from different collections. Due to logistical constraints (e.g. small
sample sizes of each sex, time allotted for examination of specimens, and
variation in the detail of collectors’ notes and care in specimen preparation)
some of the quantitative characters listed in Appendix | were assessed relatively
subjectively, and data for both sexes were lumped. This introduces a certain
amount of imprecision into the analysis. But, subjective assessments were made
against a set of reference specimens, photographs or drawings encompassing
the range of observed variation. Also, the previous studies mentioned above
have stated that sexual dimorphism is absent, or relatively minor in comparison
with geographical variation in guinea-fowl. Moreover, it was felt that the
advantages accruing from considering a large number of characters for many
specimens outweighed any sampling bias due to imprecise measurement and
sexual dimorphism.

TAXONOMIC PHILOSOPHY

The taxonomic philosophy adhered to in the present study follows that
outlined by Mayr et al. (1953). A species is a group of actually or potentially
interbreeding individuals which has diverged sufficiently in allopatry to have
become reproductively isolated from other such groups. A genus is a mono-
phyletic taxon which is decidedly qualitatively distinct, and is adapted to a
particular mode of life, i.e. a generic ‘niche’.

The subspecies is much more difficult to define. Table 1 shows that much
of the ‘taxonomic variation’ in guinea-fowl systematics is at the subspecies
level. Some systematists (e.g. Wilson & Brown 1953; Moreau 1957; Selander
1971; Gould & Johnston 1972) state that the category is useless, and even

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TABLE 1
A history of the taxonomy of guinea-fowl.*

Ber as a cv re nm

Chapin Mackworth- : Crowe
Taxa (19324 and Peters Ghigi Boetticher Praed & White (this
in litt.) (1934) (1936) (1954) Grant (1952; (1965) study)

1962; 1970)

Phasidus niger x x x x x : a
elastes niger
ip elena x x x x x x 4
Acryllium vulturinum x x x x x x x
Guttera plumifera plumifera x x x x x x x
schubotzi x x x x x x x
pucherani* x x x x
edouardi* edouardi x x x x x x x
suahelica x x x
schoutedeni x x x x x x
seth-smithi x x x x x x
verreauxi* x x x x x x x
sclateri x x x x x x x
' chapini x x x x x
granti x x x
barbata x x x x x x
lividicollis x x
kathleenae® x x x
symonst x
pucherani x x< x
Numida meleagris meleagris x se axe x x x x
sa CU eS cc ae a Koa ot ae 9 a sears am emmeere— eeen E—
> v im) es ME Pad x ? ~~ a ~ ~~
x x x x x x x
intermedia x x x x x x
toruensis x x x x x x
neumanni x
omoensis x
ansorgei x
inermis
sabyi x x x x et
galeata x x x x y : .
marchei x x x % S
strasseni x x x x ~ <
mitrata x x x se &, 5 =
reichenowi x x x x y) .
marungensis x x x x Me ~ :
callewaerti x x x x Mc .
maxima x x x x
uhehensis x x
rikwae x S
frommi x
coronata x x x % e %
limpopoensis x x
transvaalensis x
papillosa x x x % e ¥
damarensis x x x % S v 2
blancout* x
reichenowi .

1 Unless otherwise specified, nomenclature follows Peters (1934).
2G. e. verreauxi = G. e. pallasi, see White (1965).
3 See White (1965).

2 Se es two taxa are considered to be conspecific in the present study, following the principle of priority the specific name recognized is
. pucnerant.

WngsaAW NVOIdsVv HINOS AHL 40 STVNNY

TMOS-VANIND AO NOILN TOA FHL

Ly

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Quantitative characters 1, 4-17. A. Lateral view of head, neck and collar of Numida
meleagris. B. Dorsal view of the same.

Fig. 3. Quantitative characters 19-22. A. Secondary remex from Numida meleagris. B. Wing
covert from N. meleagris. C. Mid-dorsal feather of the same.

THE EVOLUTION OF GUINEA-FOWL 49

/
Oa 39
oO
Jae a ™
ere(° eh
\

Fig. 4. Quantitative characters 22-26, 29-39. A. Lateral view of head, neck and collar of
Guttera plumifera. B. Mid-dorsal feather of G. pucherani edouardi. C. Spots on mid-dorsal
feathers of Guttera spp.

detrimental to the understanding of geographic variation and intra-specific
evolution. These critics maintain that:

1. characters used to delineate subspecies usually have patterns of geographic
variation which are discordant with each other and with the distribution
attributed to the taxon;

2. indistinguishable phenotypes occur in geographically isolated areas, pre-
sumably due to parallel evolution under similar selective regimes;

3. no objective degree of difference can be offered to distinguish subspecies
from slightly differentiated local populations.

In an attempt to satisfy these legitimate criticisms, Ford (1974) offers
the taxo-evolutionary subspecies concept, the concept adhered to in the present
study. This concept limits the awarding of subspecies status to geographic
ageregates of populations which appear to have undergone genetic and pheno-
typic divergence in allopatry. Past allopatric divergence may be inferred for

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

presently parapatric taxa if their distributions can be derived from concordant
character variation, and if they are separated by a zone of secondary inter-
gradation (Ford 1974).

TAXONOMIC METHODOLOGY

Since both the recognition of taxa and their assignment to taxonomic
categories (with the possible exception of the species) are ultimately subjective
processes (Mayr et al. 1953; Selander 1971; Ford 1974), it is of utmost
importance to outline the methodology underlying taxonomic decisions.

With the foregoing taxonomic philosophy in mind, the procedure used
to determine genera involved four steps:

1. a sorting of specimens into groups which consisted of members of both
sexes, and which possessed unique combinations of qualitative characters;

2. a cluster analysis of these groups according to the number. of shared
qualitative character states;

3. presentation of the clustering similarity matrix in the form of a phenogram;

4. interpretation of the matrix and the phenogram, according to the taxonomic
philosophy outlined above, to determine genera.

Once genera were identified, each was analysed separately to determine
species and subspecies. Two multivariate statistical computer programmes
were used in tandem in these analyses. The first programme, BMDP2M (Dixon
1975), was used to cluster specimens into operational taxonomic units (OTUs,
Sneath & Sokal 1973). In BMDP2M, the clustering algorithm first amalgamates
the two specimens which are most similar. The amalgamated specimens are
then treated as one case in future comparisons. This clustering algorithm con-
tinues until all specimens are grouped into one large cluster. The similarity
measure used is the Euclidean distance, and the clustering method is the
weighted average pair group method (Sneath & Sokal 1973). The printed output
of BMDP2M includes a phenogram which illustrates the results of the cluster
analysis in a hierarchical manner. OTUs were identified from examination of
these phenograms. In a phenogram, OTU status was awarded to any specimen
cluster which contained members of both sexes from several geographically
contiguous localities, and which was morphologically more distinct (i.e. linked
up with other such groups at a lower similarity) than was a group of specimens
from a single, relatively well-sampled locality within the range of an undisputed
taxon listed in Table 1.

OTUs were compared using stepwise multiple discriminant analysis pro-
gramme BMDP7M (Dixon 1975). This programme calculates a series of
linear classification functions in a stepwise manner, such that within-OTU
variance is minimized and between-OTU variance maximized. At each step in
the analysis, the character not yet entered into the functions that best separates
OTUs is included. This procedure continues until all significant (P < 0,05)
characters are included in the functions. The printed output of BMDP7M

THE EVOLUTION OF GUINEA-FOWL 51

includes a probabilistic statement as to the OTU membership of each specimen.
In the final step, BMDP7M computes canonical discriminant functions between
OTUs, and plots the first two so as to give an optimal two-dimensional picture
of the separation of the OTUs. This plot consists of a multivariate centroid
for each OTU, surrounded by a cloud of individual points corresponding to
specimens in that OTU. In this study, an OTU, or group of OTUs, was awarded
species rank if none of the component specimens were intermediate between
two OTUs (i.e. were assigned a probability greater than 0,05 of belonging to
another OTU).

Once species were identified, character variation within each species
consisting of several OTUs was analysed to determine if any component OTUs
merited subspecies status. The taxonomic procedure used in these analyses
involved four steps:

1. division of the species range into uniform areas;

2. computation of mean values and coefficients of variation (COV, Sokal &
Rohlf 1969) for each character for each area;

3. contour mapping of mean values for each character, and of the total COV
for all characters for each area;

4. interpretation of the contour maps to determine OTUs whose distributions
could be derived from concordant character variation, and whose boundaries
were circumscribed by a zone of secondary intergradation.

Subspecies rank was awarded to any OTUs whose distributions could be
derived from concordant variation in at least three characters, and which
enclosed an area of relatively low total COV bounded by an area(s) of high
total COV. Areas with relatively low variability (i.e. low total COV) were
taken to be probable ‘core regions’ for their associated subspecies. Areas with
relatively high variability (i.e. high total COV) were taken to be regions of
secondary intergradation.

The smallest uniform area that produced sample sizes that were statistically
adequate was that enclosed by a block four degrees on a side. Contour mapping
was done with the assistance of a computer, using contouring programme
GPCP (CALCOMP 1971). This programme fits an approximate contour
surface to the data using least squares polynomial analysis.

PHYLOGENETIC PHILOSOPHY

A cladistic approach (Marx & Rabb 1970; Cracraft 1972) was used to
infer phylogenetic relationships between guinea-fowl genera and _ species.
Cladistic analysis involves discernment and use of derived character states.
Derived states are those which are unique or relatively restricted to the taxa
under study, or which could be adaptive in a social or ecological context (Marx
& Rabb 1970). Shared primitive character states, i.e. those commonly inherited
from distant ancestors, cannot be used to unite more recently evolved taxa
(Cracraft 1972). Other than its consistent, relatively objective methodology,
the major advantage of a cladistic analysis is that a proposed phylogeny can be

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

refuted on clearly specified grounds. A proposed phylogeny must be modified
or abandoned if: derived characters used to produce it are shown to be primi-
tive; another interpretation of the same or different combination of derived
character states yields a more parsimonious phylogeny, i.e. requiring fewer
convergences; a fossil form is found which possesses a character suite incom-
patible with the proposed phylogeny; or if it is grossly at variance with known
biogeographic events (Cracraft 1972; F. Vuilleumier 1975).

PHYLOGENETIC METHODOLOGY

Since the presumed ancestor of the guinea-fowl is a francolin-like phasianid
(Ghigi 1936), any character state common among extant francolins or
guineafowl-phasianid hybrids was taken to be primitive. Ghigi (1936),
Mackworth-Praed & Grant (1952, 1962, 1970) and Hall (1963) were used as
sources of information on francolins. Ghigi (1936), Bourke (1967) and R. Chapin
(in litt.) were the sources of information on the phenotypes of hybrids. Any
character state unique to, or relatively common among, guinea-fowl species
was taken to be derived.

Once hypothetical primitive-derived sequences of character states were
determined, a parsimonious phylogeny was produced following methods
outlined by Cracraft (1972).

SPECIATION

There is now widespread agreement among evolutionary biologists and
biogeographers that speciation, extinction and drastic distributional shifts in
plants and animals have been common and world-wide during the Tertiary
and Quaternary (Moreau 1966; B. Vuilleumier 1971; Axelrod & Raven 1978).
As mentioned in the introduction, past geological and climatic events have
almost certainly helped to shape patterns of speciation and biogeography in
Africa. Several authors (Chapin 1932a; Cooke 1962; Howell & Bourliére 1963;
Moreau 1966; Carcasson 1964; Butzer 1967; Hamilton 1974; Axelrod & Raven
1978) have given maps of Africa depicting the hypothetical distribution of
vegetation during relatively wetter and/or drier conditions in the past. Living-
stone (1975), Hamilton (1974) and Axelrod & Raven (1978) have pro-
vided some temporal estimates for climatic and geological events which
may have caused these conditions. As a preface to this study, the various
hypothetical vegetation maps were compared, and compromise ‘wet’ and ‘dry’
maps (Figs 5-6) were drawn which incorporated salient features of each. An
hypothetical pattern and chronology for speciation in guinea-fowl were
developed by relating the phylogeny derived herein to present-day (Fig. 7)
and hypothetical past ‘wet’ and ‘dry’ vegetation maps in the light of temporal
estimates given by authors mentioned above.

BIOGEOGRAPHY

Once the taxonomy and phylogeny of a group are defined, biogeographic
hypotheses based on patterns found in that group may be formulated. Since

THE EVOLUTION OF GUINEA-FOWL 53

14 6

Tes PT
vd Sheet
é tf

XF ae AAA CAYO
a | ae KAKKAA WV. Ay ica i Coe oe

| a es rar

By merce
( aay, een
(KX)
SH sere ag x28 er '

RAR RT EYRE RHR TOGO ITIOOL gto aay
aie SYK sas easels i SERRE gi Rb ai LARK SY
Bat
MM

ORES ee en Se RRR

Ar iged

pe a A ee
fo
aoe

NY

BS

Tropical lowland forest

Forest- savanna mosaic and
deciduous savanna woodland

aA gets |
Re fet t ell |
vas rc) gee Lawl

KX] Bushveld, grassland and steppe ‘6

4
Temperate grassland
[I] Mediterranean vegetation

[J Desert and sub- desert

[e] Lakes
Swamp

log
i ae

Fig. 5. Hypothetical vegetation map of Africa during a wetter period.

32

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

boledaa tL Saannee A

LAX
KKK)

iS)
a

nei,

Chess ms Lt
ee. OR
SRR een

ran abate
ae Rn ERIN ye) hae RXR
OXY aie ROO ry) RRR RRA
YOY ERR ERT ne
ee obs one

ek: claw
e —-

elaees ee Sa

— - a ra 0

Montane and temperate forest neh a
Ae ive

[=] Tropical lowland forest

hte ae i A
(e - 1 eo
We

aan RR a4

Forest -savanna mosaic and
deciduous savanna woodland

ea Bushveld, grassland and steppe

[| Desert and sub-desert

[e] Lakes
[fa] swamp

Aaa 'a:

at ae
ry,
vil

PNT tee
km Ves canes he
aa CRT ot

Fig. 6. Hypothetical vegetation map of Africa during a drier period.

THE EVOLUTION OF GUINEA-FOWL 35

22 14 6

; ely
Ly Sepuega
Lett TT Lp
- Scie ci
ete TR
id NO,
x
i /|

VYTY VY

hy Ts roman rae AX
Oh Reh YY ne

Hannan a mi

Bet ROKR
ee

IO XXX
ce PERRIN KS 1 oe
YY)

4 RY

te

Montane and temperate forest

Bea ee ea

Tropical lowland forest

> 1g
|
|
Forest-savanna mosaic and ee |
deciduous savanna woodland [= SSIS
wy Bushveld, grassland and steppe Belew [everclear
VY Yh 16
T rassland }
emperate grass ane
(] Mediterranean vegetation
EASA AAA se
Lal Desert and sub-desert
[e] Lakes >|
km a te |
Be Swamp ———— a — 32
0 600

Fig. 7. Present-day vegetation map of Africa.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

guinea-fowl are sedentary, stenotopic animals, the distribution of a given taxon
should reflect the geographical limits and temporal stability of its associated
biome. If a taxon has a vicariated distribution, it is likely that the vicars (see
Udvardy 1969) were isolated as a result of past partitioning of its biome.
However, if a taxon shows little geographic variation, its biome probably has
not been fragmented in the past. In this study, the boundaries of recognized
genera, species and subspecies were used to formulate an hypothetical avifaunal
map of Africa. Boundaries of taxonomically homogeneous genera (i.e. with no
subspecies) were used to delimit African avian subregions in this map. In other
words, a subregion is any biome which has presented its associated guinea-fowl
taxa with a sufficiently consistent selective regime to produce a monotypic genus,
or a genus composed of monotypic species. Species boundaries were used to
delimit provinces, and subspecies boundaries to delimit districts, following
a similar reasoning used in defining subregions.

RESULTS, DISCUSSION AND CONCLUSIONS

TAXONOMY
GENERA

There are fourteen groups of guinea-fowl specimens (Table 2) which
possess unique combinations of the qualitative characters listed in Appendix 1.
These groups are termed operational genera (OG). In the light of habitat
preference information summarized by Crowe & Snow (1978), the results of a
cluster analysis of the OG (Fig. 8) suggest that there are four genera in the
Numidinae. These are labelled A—D in Figure 7, and become apparent at the
similarity level of eleven shared characters. In Table 1, genus A corresponds
to the genera Agelastes Bonaparte, 1850, and Phasidus Cassin, 1857; genus
B to Guttera; genus C to Acryllium; and genus D to Numida. Genus A must
be named Age/lastes. The following section consists of a taxonomic summary
including taxonomic conclusions, brief descriptions and mensural statistics,
and a discussion of taxonomic conclusions when deemed necessary. All recog-
nized genera are said to be largely restricted to broad ‘niches’ (Crowe & Snow
1978). The genus Agelastes is found only in dense tropical lowland forest. The
genus Guttera 1s also limited to forest areas, but inhabits riverine forest and the
forest edge as well as tropical lowland forest. The genus Acryllium is confined
to the subdesert steppe of north-eastern Africa. The genus Numida can be found
in virtually all areas of non-forested Africa outside of desert, Mediterranean
and montane vegetation.

SPECIES AND SUBSPECIES

The genus Acryllium appears to be monotypic. A cluster analysis of 43
Acryllium specimens (Fig. 9) according to 7 quantitative characters (nos 1-5,
41-42 in Appendix 2) yields only one OTU. Six individuals from Mt Kunchurro,
Boran, Ethiopia (c. 4°30’N 38°E), link to form a cluster at a level (indicated

57

THE EVOLUTION OF GUINEA-FOWL

TABLE 2

Operational genera derived from analysis of qualitative character states listed in Appendix 1.

Operational genera

Character

Sy LOSS SIM Gs Il athe a Ql Gs Ws Gs i eli celia @
UO CME SCA CO NO NEN CNN Fe Shi ret mer)
BSTC CUTE SARE ECT CAE CNC CA tat arte
Soe eee NEC EAC CAEN CD CN ONION a ST et ac)
Ose ON eet STON NECN tS a
DANATTANTRANANTANATANANANAAS Tt
CONMANANMAARAAMNANANANAA Tt
mAMATAAINANTRNAMRNMNANNANAA TST
ONMAANANANRBNTAAMNANTDAAA=sST
MAMNMNAANTMANABNMNAAAAAS Tt
FAMNTANTRANDBNMNAANANADWAAT
NANATFTAN RIAN MRUONAANTAAMANN
CU Tey UN sete ce esa en ee em erie
ee et a ee se ws i si me st OO Oe

OANNANHNORODOHAAMNANWON oO
ra es eS st et Sst

* Only in the juvenile bird.

12

14

10

13

NUMBER OF SHARED CHARACTERS

B. Guttera.

Fig. 8. The results of a cluster analysis of OG in Table 2. A. Agelastes.

C. Acryllium. D. Numida.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

1 7 6 9 2319 4 1311 2 26 4032 3 14 5 43 10 21 41 27 16 36 18 39 33 35 37 38 286 34 30 29 42 31 22 2517 15 20 8 2412

0

JONVLISIG NV3GI19N4g
w&

Fig. 9 The results of a cluster analysis of forty-three specimens of Acryllium.

by the dashed line in Fig. 9) lower than does any combination of specimens
from the remainder of the distribution of the genus. Thus, for the characters
investigated, variation in a single population is as great as that found through-
out the range of the genus. Accordingly, in agreement with all previous studies
(Table 1), one monotypic species, Acryllium yulturinum, is recognized. The
distribution of this species is plotted in Figure 10.

The genus Agelastes is composed of two monotypic species. A cluster
analysis of 49 specimens according to 11 quantitative characters (nos 1-5,
23-24, 41-44 in Appendix 2) yields two OTUs, labelled A and B in Figure 11.
The dashed line in the figure indicates the level at which eight individuals from
Kribi, Cameroons (2°50’N 10°5’E), link to form a cluster. A discriminant
functions analysis of the two OTUs reveals no intermediate specimens. There-
fore the OTUs are recognized as species. In Table 1, OTU A corresponds to

TABLE 3

Similarity matrix showing number of shared qualitative character states between OG derived
and listed in Table 2.

OG no. 1 2 3 4 5 6 il 8 9 10° If “2. See
1 — 13 5 2 2 3 3 1 2 5 5) 4 4 5)
2 — 7 3 2 4 4 2 3 5 4 4 4 5
5 — 6 6 6 6 8 9 a / a 6 i
4 — 17 14 13 14 = «13 8 8 9 9 8
5 Sy Sy ele ES ee 8 8 9 9 8
6 == 5.;./ to: > 15 6 6 6 5 6
jj — 15 12 4 4 4 4 4
8 — 15 6 6 6 5 6
9 — 7 i 7! 6 7

10 — 13 5) Assay
11 — 13 14 14
12 — 15 16
13 — 13

THE EVOLUTION OF GUINEA-FOWL 59

|
one Polen
x y wh |
K x!
wy WY) C x 16
HN

ap, ae, ae are § XXX Xx ye N
Yh werk (eos) ieee yp j x x SAA AA AR AAA AR AS ,
x xX 4 y Ko Ny Es ae fy

g h
J = i
nS x XK XK KY OOOO OOOOS . “XX vk \ x |
YOO OOOO ; xx x «x 4
Be XXX) > 40 4 x x 4 x x xX x dX X X x x ‘
‘\ KX £o,4 A> Od > KXAXKXW NX XX XIX XK KX XW) vw VW YY ¥ ¥ x. VOM |
PY ROOK ( dy eee: i ayy YY Kk x2 > —
Y q -¥ \ ; oan
YY Y , <x
v EN b
\ hp 4 ¢
K

Montane and temperate forest

Tropical lowland forest

Forest-Savanna mosaic and
deciduous Savanna woodland | .A[- --|- = -

Bushveld, grassland and steppe PNEV Nos ioc IS (Sere Oe beter ae

Temperate grassland
Mediterranean vegetation
Desert and sub-desert

Lakes

AN

Swamp pe ‘ tee: AK
0 600 1200 NU i

Fig. 10. The distributions of Agelastes meleagrides (C1), Agelastes niger (MB), and Acryllium
vulturinum (CO).

Boe Sh eh el eles

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

112 8 4 611 2 9103 5 7 23 49 17 304645 15 14 21 38 18 29 34 42 31 26 28 40 39 22 3635 19 48 47 43 27 33 37 24 25 16 4420 32 41 13

INVLSIG NVAGITONG

d

5

Fig. 11. The results of a cluster analysis of forty-nine specimens of Agelastes. A. A. melea-
grides. B. A. niger.

Agelastes meleagrides and OTU B to A. niger. The distributions of these species
are plotted in Figure 10.

The remaining two genera are taxonomically more complex than are the
first two. The results of a cluster analysis of 494 Guttera specimens according
to 24 quantitative characters (nos 1-5, 22-40 in Appendix 2) are summarized
in Figure 12A. The dashed line in Figure 12A indicates the level at which twelve
individuals from Ngayu, Zaire (1°45’N 27°15’E), link to form a cluster. Seven
OTUs are recognized. A discriminant functions analysis (Fig. 13) suggests that
there are two groups of OTUs between which there are no intermediate
individuals. These OTU groups are recognized as species. The first species,
comprising OTUs | and 2, corresponds to G. plumifera in Table 1. The second
species, comprising OTUs 3-7, corresponds to two commonly recognized
species, G. pucherani (Hartlaub), 1860, and G. edouardi (Hartlaub), 1867
(Table 1). Following the law of priority, this species must be named G. pucherani.

The two OTUs comprising Guttera plumifera partition the distribution
of that species into eastern and western portions. The distributions of these
OTUs (Fig. 14) are delineated by patterns of variation in fifteen of the sixteen
quantitative characters which vary in G. plumifera. Contour maps of variation
in these characters, and of the total COV for six areas (Fig. 15) within the distri-
bution of G. plumifera are given in Figures 16 and 17. Two types of character
variation, ‘mountain—valley’ and clinal variation, are apparent in these contour
maps. ‘Mountain-valley’ variation occurs when eastern and western OTUs
have similar values, and are separated by a transition area(s) with higher
(‘mountains’) or lower (‘valleys’) values. Characters which show ‘mountain—
valley’ variation are: bill length (Fig. 16B), wing length (Fig. 16C), tarso-
metatarsus length (Fig. 16D), wattle length (Fig. 16F), crest frontal length
(Fig. 16G), crest basal length (Fig. 17J), dorsal spot number (Fig. 17K), total
spot barbs (Fig. 17M), and total within-spot barbs (Fig. 17N). Characters
which show clinal variation are: occipital fold (Fig. 16A), wattle basal width
(Fig. 16E), crest rear length (Fig. 16H), dorsal spot size (Fig. 17L), ear patch

THE EVOLUTION OF GUINEA-FOWL 61

FONVLSIG NVAGITONA
wn

Fig. 12. The results of a cluster analysis. A. Guttera specimens B. Numida specimens.

t JTAVIYVA TWOINONVO

=6 -4 -2 O +2 +4 +6 +8 +10 +12 +14
CANONICAL VARIABLE |

Fig. 13. A discriminant functions analysis of Guttera OTUs. Circles encompass 90 per cent

of the individuals assigned to each OTU. Only intermediate specimens between non-

overlapping OTUs are plotted. B — intermediate between OTUs 5 and 6; E — intermediate
between OTUs 6 and 7; P — intermediate between OTUs 1 and 2.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Vv

||
aia

BB Montane and temperate forest

Tropical lowland forest

Forest-savanna mosaic and
deciduous savanna woodland eS as lise ere ; a | A ee mas

Bushveld, grassland and steppe MEA RS || GSE oe ae

Temperate grassland

Mediterranean vegetation

4
Desert and sub-desert

X Y ; y\ KAAAA (
a, x x \ A Vi
. } : : Sy
Xx AA A}
¥ ae ae = ty
Lakes (5 0.01, 0,0,0 Na
km i’ Sb. k [|

Fig. 14. The distribution of Guttera plumifera plumifera (W) (= OTU 2), G. p. schubotzi
(VY) (= OTU 1), and a single intergrade (x ).

Fa iO es i 8

THE EVOLUTION OF GUINEA-FOWL 63

Fig. 15. Areas used in contour map analysis of Guttera spp. Those areas marked with an ‘X’
have data for both species. Those marked with a ‘P’ have data for G. plumifera only, and
those without notation for G. pucherani only.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

14°E
A |
B J
30 28
C 233230 227 223 K
D
L 11,0
\
E Mi
34
F A a
fi
H P

i=)

Fig. 16. Contour maps of character
variation in Guttera plumifera. A. Occipital
fold. B. Bill length. C. Wing length.
D. Tarso-metatarsus length. E. Wattle

width. F. Wattle length. G. Crest frontal

length. H. Crest rear length. OTU Fig. Contour maps of character

boundaries are indicated by thick lines. mee and total COV for Guttera
plumifera. I. Crest central height. J. Crest
basal length. K. Dorsal spot number.
L. Dorsal spot size. M. Total spot barbs.
N. Total within spot barbs. O. Ear patch.
P. Spot barb blueness. Q. Total COV.
OTU boundaries are indicated by thick

lines.

THE EVOLUTION OF GUINEA-FOWL 65

(Fig. 170), and spot barb blueness (Fig. 17P). These well-defined OTUs each
have a region of relatively low total COV (c. 250) within their distributions, and
are separated by a transition area with a relatively high total COV (c.750)
(Fig. 17Q). Thus, both OTUs meet the criteria set for subspecies (see Taxonomic
methodology). In Table 1, the western subspecies (OTU 2) corresponds to
G. p. plumifera, and the eastern subspecies (OTU 1) to G. p. schubotzi. In
statistical comparisons, these subspecies differ significantly (P < 0,05; ¢ test)
in eight characters. Guttera plumifera plumifera has significantly higher values
for bill length, wattle basal width, wattle length, crest rear length, and crest
basal length. Guttera plumifera schubotzi has higher values for occipital fold, ear
patch, and spot barb blueness. Means and standard deviations for these and
other characters are given in the section below.

Approximate geographic distributions of the five OTUs comprising
G. pucherani are shown in Figure 18A. Contour maps of twenty-three characters
which vary in this species (nos 1-5, 22-31, 33-40 in Appendix 2) are given in
Figures 19-30A. In these maps, the distributions of all five OTUs are delineated
from those of their neighbours by statistically significant patterns of variation
in six to seventeen of the characters analysed. The results of all possible
pairwise statistical comparisons (f tests) between neighbouring OTUs are
summarized in Tables 4 and 5. A contour map of variation in total COV for
twenty-five areas (Fig. 15) within the distribution of G. pucherani is given in
Figure 30B. This figure shows that the distributions of all five OTUs enclose
or fall within a region of relatively low total COV (200-400). For all OTU
distributions but one, that of OTU 4, the region of relatively low total COV
is bordered by a region(s) of relatively high total COV (e.g. 600-800). Thus, all
OTUs ascribed to G. pucherani satisfy the criteria specified for subspecies (see
Taxonomic methodology). The lack of a high total COV interface between the
low COV regions of OTUs 3 and 4 is attributed to poor sampling and the small
geographic distribution of OTU 4. Assuming that G. edouardi is a synonym
of G. pucherani in Table 1, and following the law of priority, OTU 3 corresponds
to G. p. verreauxi, OTU 4 to G. p. sclateri, OTU 5 to G. p. pucherani, OTU 6
to G. p. barbata, and OTU 7 to G. p. edouardi. The geographic distributions of
these subspecies, and of intermediate populations, are shown in Figure 31.

The results of a cluster analysis of 704 Numida specimens according to
characters |—22 in Appendix 2 are summarized in Figure 12B. Nine well-defined,
and one borderline OTU are recognized. The dashed line indicates the level at
which eight individuals from the vicinity of Gassam, Senegal (14°50’N 15°20’W),
link to form a cluster. Only 704 specimens could be analysed in this cluster
analysis due to computer storage limitations. However, those specimens analysed
were chosen so as to ensure uniform sampling of the sexes and collection locali-
ties. A discriminant functions analysis including all 1 245 Numida specimens
(Fig. 32) reveals intermediate individuals between all parapatric OTUs. There-
fore only one species is recognized. Following the law of priority, this species
is named Numida meleagris.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

ast

Fig. 18. The geographic distributions of OTUs comprising A. Guttera pucherani
and B. Numida meleagris.

THE EVOLUTION OF GUINEA-FOWL 67

A

Fig. 19. Contour maps of character variation in Guttera pucherani. A. Bill length. B. Wing
length. OTU boundaries are indicated by thick lines.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 20. Contour maps of character variation in Guttera pucherani. A. Tarso-metatarsus
length. B. Wattle basal width. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 69

Fig. 21. Contour maps of character variation in Guttera pucherani. A. Wattle length.
B. dorsal spot size. OTU boundaries are indicated by thick lines.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 22. Contour maps of character variation in Guttera pucherani. A. Crest frontal length.
B. Crest rear length. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL Wi

Fig. 23. Contour maps of character variation in Guttera pucherani. A. Crest central height.
B. Crest basal length. OTU boundaries are indicated by thick lines.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 24. Contour maps of character variation in Guttera pucherani. A. Anterior crest curli-
ness. B. Posterior crest curliness. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 73

Fig. 25. Contour maps of character variation in Guttera pucherani. A. dorsal black collar.
B. Ventral black collar. OTU boundaries are indicated by thick lines.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. Contour maps of character variation in Guttera pucherani. A. Occipital fold.
B. Throat red. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL TS

Fig. 27. Contour maps of character variation in Guttera pucherani. A. Orbital red. B. Dorsal
spot number. OTU boundaries are indicated by thick lines.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 28. Contour maps of character variation in Guttera pucherani. A. Total spot barbs.
B. Total within spot barbs. OTU boundaries are indicated by thick lines.

:
— “

THE EVOLUTION OF GUINEA-FOWL Gh

Fig. 29. Contour maps of character variation in Guttera pucherani. A. Spot barb blueness.
B. Chestnut blotch size. OTU boundaries are indicated by thick lines.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 30. Contour maps of character variation in Guttera pucheranj. A. Chestnut blotch
extent. B. Total COV. OTU boundaries are indicated by thick lines. Larger numbers in
B. refer to OTU numbers shown in Figure 18A.

THE EVOLUTION OF GUINEA-FOWL 79

6 6
eo ss eo
D | | ; | | | | \ \ ' eo |
poet at Po | | Ale | + =e a \ aay
ATID (<i ! KOO) | ae | \ \ i
An) 7K eo | ae os | | \ —_\+32
R090 ee oe |

Leo > SD
, He ae ah aN ad be
606. ee x

Cie
RAR NUK OCC EKO

fox
PB Montane and temperate forest

Tropical lowland forest

Forest-savanna mosaic and
deciduous savanna woodland

Bushveld, grassland and steppe Meo. ae of ff.

Temperate grassland

Mediterranean vegetation

Desert and sub-desert

Lakes

Swamp

[4
iY
Hl
is
[ej
Ed

Fig. 31. The distributions of Guttera pucherani subspecies. G. p. pucherani (MB). G. p. verreauxi
(A). G. p. sclateri (OQ). G. p. barbata (C0). G. p. edouardi (@). Intergrades (x).

80

TABLE 4

ANNALS OF THE SOUTH AFRICAN MUSEUM

A comparison of OTU 3 for G. pucherani with OTUs 4, 5 and 7. x — not significantly
different (P < 0,05; ¢ test); + — OTU 3 significantly greater; — — OTU 3 significantly
lower; * — differences delineate OTUs in contour maps.

Character name?

Bill length A :
Wing length . ; ;
Tarso-metatarsus lengt
Wattle basal width
Wattle length

Dorsal spot size

Crest frontal length
Crest rear length .
Crest central height
Crest basal length
Anterior crest curliness
Posterior crest curliness
Dorsal black collar
Ventral black collar
Occipital fold

Throat red

Orbital red

Dorsal spot number
Total spot barbs .
Total within spot barbs
Spot barb blueness
Chestnut blotch size
Chestnut blotch extent

1 See Appendix 2 for character descriptions.

@ AIVIYVA TVIINONVD

CANONICAL VARIABLE |

OTU

Contour map
Figure no.
19A

Fig. 32. A discriminant functions analysis of Numida OTUs. Circles encompass 90 per cent
of the individuals assigned to each OTU. Only intermediate specimens between non-
overlapping OTUs are plotted. M — intermediate between OTUs 4 and 7; R — intermediate

between OTUs 4 and 8; X — intermediate between OTUs 4 and 2.

THE EVOLUTION OF GUINEA-FOWL 81

TABLE 5

A comparison of OTU 6 for G. pucherani with OTUs 5 and 7. x — not significantly different
(P < 0,05; ¢ test); + — OTU 6 significantly greater; — — OTU 6 significantly lower; * —
differences delineate OTUs in contour maps.

Character name* OTU Contour map
5 7 Figure no.
Bill length 4 x x 19A
Wing length . : ; x Le: 19B
Tarso-metatarsus lengt x de; 20A
Wattle basal width x _* 20B
Wattle length x ae 21A
Dorsal spot size . x _* 21B
Crest frontal length -* x DN
Crest rear length . ae —* 22B
Crest central height x 23A
Crest basal length x + * 23B
Anterior crest curliness x _* 24A
Posterior crest curliness x —* 24B
Dorsal black collar JLES _* 25A
Ventral black collar a = 25B
Occipital fold x +* 26A
Throat red aa aL 26B
Orbital red = a 7A
Dorsal spot number x JL 27B
Total spot barbs . x ae 283A
Total within spot barbs x -.* 28B
Spot barb blueness = au 29A
Chestnut blotch size ae = 29B
Chestnut blotch extent ae —* 30A

1 See Appendix 2 for character descriptions.

The approximate geographic distributions of the ten OTUs comprising
N. meleagris are shown in Figure 18B. Contour maps of variation in twenty-two
quantitative characters analysed for this species (nos 1-22 in Appendix 2),
and of the total COV for forty-eight areas (Fig. 33) are given in Figures 34-44.
In these contour maps, the distributions of all ten OTUs but one, that of OTU 3,
are delineated from those of their neighbours by statistically significant patterns
of variation in nine to seventeen of the twenty-two characters analysed (Tables
6-11). Results of all possible pairwise statistical comparisons (¢ tests) between
neighbouring OTUs are summarized in Tables 6-11. The distribution of OTU
3, the borderline OTU in Figure 11B, is delineated from those of its neighbours
(OTUs 2 and 6) by at most two characters (Tables 7, 9). In the contour map of
total COV (Fig. 45), the distribution of all OTUs, except again OTU 3, enclose
a region of relatively low total COV (c. 400-500) bordered by a region(s) of
relatively high total COV (c. 700-1000). Thus, all OTUs ascribed to N. meleagris,
except OTU 3, satisfy the criteria set for subspecies (see Taxonomic methodology).
In Table 1, following the law of priority, OTU 1 corresponds to N. m. sabyi,
OTU 2 to N. m. galeata, OTU 4 to N. m. meleagris, OTU 5 to N. m. somalienses,
Siw 7G to N. m. marungensis, OTU 7 to. N. m. mitrata, OTU 8 to
N. m. reichenowi, OTU 9 to N. m. damarensis, and OTU 10 to N. m. coronata.

82

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Areas used in contour map analysis of Numida meleagris.

THE EVOLUTION OF GUINEA-FOWL 83

Fig. 34. Contour maps of character variation in Numida meleagris. A. Bill length. B. Wing
length. OTU boundaries are indicated by thick lines.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Contour maps of character variation in Numida meleagris. A. Tarso-metatarsus
length. B. Wattle basal width. OTU boundaries are indicated by thick lines.

——

THE EVOLUTION OF GUINEA-FOWL 85

Fig. 36. Contour maps of character variation in Numida meleagris. A. Wattle length.
B. Wattle per cent blue. OTU boundaries are indicated by thick lines.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 37. Contour maps of character variation in Numida meleagris. A. Helmet frontal
length. B. Helmet rear length. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 87

Fig. 38. Contour maps of character variation in Numida meleagris. A. Helmet central height.
B. Helmet basal length. OTU boundaries are indicated by thick lines.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 39. Contour maps of character variation in Numida meleagris. A. Helmet thickness.
B. Cere structure length. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 89

Fig. 40. Contour maps of character variation in Numida meleagris. A. Cere structure thick-
ness. B. Collar plumage. OTU boundaries are indicated by thick lines.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41. Contour maps of character variation in Numida meleagris. A. Nape filoplume length.
B. Nape anteroposterior coverage. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 9]

Fig. 42. Contour maps of character variation in Numida meleagris. A. Nape filoplume lateral
coverage. B. Nape filoplume density. OTU boundaries are indicated by thick lines.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 43. Contour maps of character variation in Numida meleagris. A. Secondary remex
outer web vermiculation. B. Wing covert barring. OTU boundaries are indicated by thick lines.

THE EVOLUTION OF GUINEA-FOWL 93

Fig. 44. Contour maps of character variation in Numida meleagris. A. Dorsal vermiculation.
B. Dorsal spot size. OTU boundaries are indicated by thick lines.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 6
A comparison of OTU 4 for N. meleagris with OTUs 2, 5, 6, 7 and 8. x — not significantly
different (P < 0,05; ¢ test); + — OTU 4 significantly greater; — — OTU 4 significantly lower;
* — differences delineate OTUs in contour maps.
Character name? OTU Contour map
D, 5 6 il 8 Figure no.

Bill length . : < ’ ; : +* S< —* — x 34A
Wing length : : : : ‘ -- < —* —* * 34B
Tarso-metatarsus length . : 2 +* % —* — = 35A
Wattle basal width . : : fi x< < +* ae — 35B
Wattle length. : as ‘ : _ —* —* =+ aL 36A
Wattle per cent blue . ; : ‘ +* +* +* * ae 36B
Helmet frontal length ; : : + * x —* —* —* 37A
Helmet rear length . ; ‘ F +* x< — = = 37B
Helmet central height . ; ‘ +* < — —* —* 38A
Helmet basal length . : ; : < +* —* —* —* 38B
Helmet thickness : : : ‘ +* SE —* —* _ 39A
Cere structure length : : +* —* — a a5 39B
Cere structure thickness . : : +* < =e — =o 40A
Collar plumage . : ‘ ; P +* x x — x 40B
Nape filoplume length . : ; —* —* —* —* —* 41A
Nape filoplume anteroposterior

coverage . ; —* +* —* ‘< x 41B
Nape filoplume aera Coverane , +* +* +* +* +* 42A
Nape filoplume density . : : x S< x = +* 42B
Secondary remex outer web ver-

miculation . : ; : , +* o +* +* +* 43A
Wing covert barring . ‘ , : +* +* +* +* +* 43B
Dorsal vermiculation , : ‘ —* + +* +* + 44A
Dorsal spot size . : +* —* - x —* 44B

1 See Appendix 2 for ee deseanucne
TABLE 7
A comparison of OTU 2 for N. meleagris with OTUs 1, 3 and 6. x — not significantly different
(P < 0,05; ¢ test); + — OTU 2 significantly greater; — — OTU 2 significantly lower; * —
differences delineate OTUs in contour maps.
Character name* OTUs Contour map
1 3 6 Figure no.

Billlength . : : : : ; : : x _ —* 34A
Wing length . 3 : : ; —* — —* 34B
Tarso-metatarsus length : : : —* — —* 35A
Wattle basal width * x +* 35B
Wattle length ; : : : : 5 . * »< _ 36A
Wattle per cent blue . : : : : , +* = —* 36B
Helmet frontal length . ; : : —* — —* 37A
Helmet rear length . : ; ; : : : —* — — 37B
Helmet central height . —* — — 38A
Helmet basal length x — —* 38B
Helmet thickness . : : : 3 : : . * — _ 39A
Cere structure length . x — x 39B
Cere structure thickness x = x 40A
Collar plumage —* —* —* 40B
Nape filoplume length. : —* x —* 41A
Nape filoplume anteroposterior coverage x x =F 41B
Nape filoplume lateral coverage ; : x =e EG 42A
Nape filoplume density * x x 42B
Secondary remex outer web yermiculation x =P ale BA
Wing covert barring . : : é: +* + +* 43B
Dorsal vermiculation . . . .-. eae +* == 2h 44A
Dorsal spot size... : +* — — 44B

1 See Appendix 2 for nietie desiniiods:

THE EVOLUTION OF GUINEA-FOWL 95

TABLE 8
A comparison of OTU 8 for N. meleagris with OTUs 5 and 7. x — not significantly different;
(P < 0,05; ¢ test); + — OTU 8 significantly greater; — — OTU 8 significantly lower; * —
differences delineate OTUs in contour maps.
Character name+ OTUs Contour map
5) 7 Figure no.
Billlength . : : : : : ; ; ‘ : x —* 34A
Wing length . : : : ; ; ‘ : : ; +* +* 34B
Tarso-metatarsus length 5 3 ; , : : ; + * +* 35A
Wattle basal width. ; , : : ‘ . : : — - 35B
Wattle length F ; : : ; ; : ; 3 ; — — 36A
Wattle per cent blue . : ; : ; : ; f > =F —* 36B
Helmet frontal length . j : : : t : 4 e +* =a 37A
Helmet rear length . : : : ; : : 5 : +* * 37B
Helmet central height . : i : : ; 5 , : + +* 38A
Helmet basal length . : ; ; : : : : : +* +% 38B
Helmet thickness . : : : ' x A , : : _ + 39A
Cere structure length . —* _ 39B
Cere structure thickness — + 40A
Collar plumage x — 40B
Nape filoplume length +* = 41A
Nape filoplume anteroposterior coverage + x 41B
Nape filoplume lateral coverage : a _ 42A
Nape filoplume density : : : ; —* x 42B
Secondary remex outer web vermiculation : d : : —* +* 43A
Wing covert barring . : ' : : a + 43B
Dorsal vermiculation . : : : : : : ; : x = 44A
Dorsal spot size . : : : : : ‘ * — 44B
1 See Appendix 2 for seers Tere
TABLE 9
A comparison of OTU 6 for N. meleagris with OTUs 3, 7 and 9. x — not significantly different
(P < 0,05; ¢ test); + — OTU 6 significantly greater; — — OTU 6 significantly lower; * —
differences delineate OTUs in contour maps.
Character name} OTUs Contour map
3 7 9 Figure no.
Bill length at == aes 34A
Wing length . a= a5 a 34B
Tarso-metatarsus length x +* +* 35A
Wattle basal width — +* == 35B
Wattle length é ; : , ' _ x = 36A
Wattle per cent blue . : : : : 5 : +* +%* ++ 36B
Helmet frontal length . : ‘ ; 3 : + +* +* 37A
Helmet rear length ‘ — — —* 37B
Helmet central height . : : . : + x — 38A
Helmet basal length . : ; : ; ; : + +* +* 38B
Helmet thickness . Pee hae Mee: + a5 ae 39A
Cere structure length . : : : : : . + sé —* 39B
Cere structure thickness : ; + * —* 40A
Collar plumage . : : : : == * —* 40B
Nape filoplume length + +* a= 41A
Nape filoplume anteroposterior coverage : : — +* +* 41B
Nape filoplume lateral coverage : j : _ -- +* 42A
Nape filoplume density : : : x +* +* 42B
Secondary remex outer web venmiculation : ‘ — — —* 43A
Wing covert barring . ; : : — x x 43B
Dorsal vermiculation . : : : : — _ —* 44A
Dorsal spot size . : n : ; == == x 44B

1 See Appendix 2 for shanadict HeccEDHOAS:

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 10
A comparison of OTU 10 for N. meleagris with OTUs 7 and 9. x — not significantly different
(P < 0,05; ¢ test); + — OTU 10 significantly greater; — — OTU 10 significantly lower;
* — differences delineate OTUs in contour maps.
Character name* OTUs Contour map
7 9 Figure no.
Bill length . 5 : ; : 3 i : : . ; = ar 34A
Wing length . : ‘ : : : : ; ; SS = 34B
Tarso-metatarsus jensth f 3 ; ; : ; 5 ; x + 35A
Wattle basal width . : ; ; : : : : : x x 35B
Wattle length ; ; : ; : : ; A +* x 36A
Wattlespemcentblucy i) i) fy see ey a eR ee —* 36B
Helmet frontal length . f 5 ‘ ; ; ; : +* +-# 37A
Helmet rear length . ‘ : ‘ : : : : ‘ +* -+-* 37B
Helmet central height . : : : 2 : ; . : +* +* 38A
Helmet basal length . : A : : : : Emer +* 38B
Helmet thickness . : : : : 4 : : : : + x 39A
Cere structure length . : ‘ ; E : . : + —* 39B
Cere structure thickness. : ; ‘ ; : : + —* 40A
Collar featheration . : : ; : : : i melee —* 40B
Nape filoplume length : a ee a 41A
Nape filoplume anteroposterior coverage co og Dee eS a= 41B
Nape filoplume lateral coverage : : i : : : = =." 42A
Nape filoplume density ; ; : : ; — +* 42B
Secondary remex outer web vermiculation ‘ : : » ees = 43A
Wing covert barring . : : , 3 : : x x 43B
Dorsal vermiculation . ‘ : F : i i : : +* x 44A
Dorsal spot size . : WME 9 ie el —* 44B
1 See Appendix 2 for Charice ee ee
TABLE 11
A comparison of OTU 7 for N. meleagris with OTU 9. x — not significantly different (P <
0,05; ¢ test); + — OTU 7 significantly greater; — — OTU 7 significantly lower; * — differences
delineate OTUs in contour maps.
Character name? OTU Contour map

9 Figure no.
Bill length : : 3 . ; 5 : : : 4 +
Wing length . : : 2 s , ; : 3 : : + 34B
Tarso-metatarsus jeneth : ; : ' 4 : +
Wattle basal width x
Wattle length =

x

Wattle per cent blue 36B
Helmet frontal length . ; : : : ; : : 37A
Helmet rear length . é ; : : ? ; : : : —* 37B
Helmet central height . : ‘ ’ ‘ ; ; t ; : —* 38A
Helmet basal length . : : : , : : : : ; +* 38B
Helmet thickness . : A : : : : : . —* 39A
Cere structure length . : 5; : : : : —* 39B
Cere structure thickness. ‘ : : : : : ‘ s —* 40A
Collar featheration . : ; : ; ; : : —* 40B
Nape filoplume length 3 ; : . , : +* 441A
Nape filoplume anteroposterior coverage : ; : : ; +* 41B
Nape filoplume lateral coverage : 5 ; : ; ; : +* 42A
Nape filoplume density : : : : : +* 42B
Secondary remex outer web vermiculation : ; ; : : —* 43A
Wing covert barring . , Fj ‘ : ‘ : ‘ ; : x 43B
Dorsal vermiculation . : : : : ; —* 44A
Dorsal spot size . . } ' ; é ‘ —* 44B

1 See Appendix 2 for cece escape

THE EVOLUTION OF GUINEA-FOWL 97

Fig. 45. A. Contour map of total COV in Numida meleagris with OTU boundaries indicated
by thick lines. B. The approximate distribution of zones of intermediacy between Numida
OTUs. Larger numbers in A refer to OTU numbers shown in Figure 18B.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

The distributions of these subspecies are plotted in Figure 46, and zones of
intermediacy are shown in Figure 45B.

TAXONOMIC SUMMARY

In the following classification, verbal descriptions are brief, focusing on
more obvious differences and similarities with respect to the qualitative and
quantitative characters analysed. For each subspecies, reference is made to its
OG or OTU number and the appropriate page(s), tables and figures summarizing
its taxonomic relationships and quantitative comparisons with other subspecies.
If a synonymized taxon is not discussed, it must be assumed that it did not
emerge as an OTU in the cluster analyses. In other words, it was not taxo-
nomically distinct above a level attributable to individual variation in a relatively
homogenous population. Subspecies which are mentioned in Table 1, but are
not recognized or synonymized, are taken to be intergrades. Herein, an inter-
grade is a group of taxonomically indeterminable, phenotypically highly variable
populations. A population is taken to be taxonomically indeterminable if
many of its component individuals show affinities to two or more subspecies
(i.e. are classed as intermediates) in discriminant analyses. A population is
taken to be highly variable phenotypically if it has a high total COV. See the
section on Taxonomic methodology for a detailed discussion of OTUs, dis-
criminant analyses and total COV.

Genus Agelastes
Agelastes Bonaparte, 1850
Fig. 8(0GI1-2)
Agelastes meleagrides Bonaparte, 1850
Figs 1A, 8(0G1)
Agelastes meleagrides Bonaparte, 1850: 145.

Description

Small overall size; no crown, occipital, cere, nape or throat adornments;
rudimentary red gape wattles; no feathers on head or neck, skin colour of head
and neck red; collar plumage white; body plumage black with faint vermicu-
lations; tarso-metatarsus covered with imbricated scales in rows, and usually
with a well-developed spur(s); iris brown; outer margins of secondaries black
with faint vermiculations; furcula blade-shaped; abdominal plumage white.

Statistics for quantitative characters (N = 12)

Character X S.D.
BIpleMmeo tie 2a fon i. abo cube eM cele sige ea Cm eases eae 16,8 mm 17
Wine len@th . sleek. . 20. oe 2 ZO 4,8
iarso-metatansus lemeth. 2° 202°. al. 2 ee 80,9 mm 3,6

Waitt Tel OV GOIN aoe esis eh Selene’ “Gone Eek Vey ure alee ence 6,5 mm 2

THE EVOLUTION OF GUINEA-FOWL ‘09

ile us Ly Py aad

x mi A 2 ' ‘ ~ ee a pg
Seas aoe Ne aA HE Mie a nace Oh aA

: = 1s —oo
a and temperate forest

Tropical lowland forest

Forest-Savanna mosaic and
deciduous savanna woodland

| i Bushveld, grassland and steppe
:
Ey Temperate grassland
TW
Ue

Mediterranean vegetation

Desert and sub-desert

Lakes
K Mm
Swamp _— rs)
0 600

|
pk = =

Fig. 46. The distribution of Numida meleagris. N. m. meleagris (A). N. m. sabyi (s).
galeata (L)). N. m. somaliensis (O). N. m. marungensis (HB). N. m. reichenowi (VY).
N. m. mitrata (t). N. m. damarensis (A). N. m. coronata (@).

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character x S.D.
NWiattleslemetla 08 aay an ere ene eal ee ee aati mt 2,7/mm OW
Tarsalsinucrune mulches ee 0,8 0,6
ansall structune lemetine (hy yn) eo eee 4,8 mm 85)
White collar) js % pe, 0 Aa ais ie, eg oem ea KOON OR A 0,0
Facial floplamess (3.4560) ian) Man eee oe ne 0,0 0,0
Distribution

See Figure 10.

Discussion

White collar and facial filoplumes are used as quantitative characters in
cluster and discriminant analyses to demonstrate the qualitative distinctness of
this taxon from A. niger.

Agelastes niger (Cassin, 1857)
Figs 1B, 8(0G2)
Phasidus niger Cassin, 1857: 322.

Description

As A. meleagrides except: crown surmounted by a short crest of feathers;
nape covered by short, sparsely distributed, black downy feathers; face covered
with sparsely distributed filoplumes; collar and abdominal plumage black with
faint vermiculations, abdominal plumage white in juveniles.

Statistics for quantitative characters (N = 39)

Character X S.D.
Bil Memetln wees PIC voi nee ee aig omn 3 Car leew 16,9 mm igs)
Wing length ae ae A en en enue ee ())3) 0) Sclida 13
Marso-metatansus lemorh: oan eae en ee 79,2 mm 3,4
Wattlevbasale wig. "eu 2h spose cela 6,5 mm 12
Wattle lemetin “is ys cam ac ee Val sian To [ite 2,4 mm 0,6
Mansa eseeuceune mune |) eae) tee) ene IL Il 0,7
Tarsal structure length Oe te Nek a ee RR BN i 3,4 mm 2,4
Wonite collar, agus, ooo. Ue uoey eds i Way gti ae wa OO, 0,0
PactalMlOplwMies th) Mee a ee Oe Weta ee, 9,8 0,2
Distribution

See Figure 10.

Discussion

Hall (1961) also advocates the synonomy of Phasidus in Agelastes, basing
her argument largely on analysis of juvenile characters.

THE EVOLUTION OF GUINEA-FOWL 101

Genus Guttera
Guttera Wagler, 1832
Fig. 8(0G4-9)
Guttera plumifera plumifera (Cassin, 1857)
Figs 1C, 12A(OTU2), 13(0TU2)
Numida plumifera Cassin, 1857: 321.

Description

Larger than Agelastes spp.; crown surmounted by a crest of long, straight,
bristly feathers; small occipital fold of blue-black skin; no nape, throat or
cere adornments; well-developed, pointed, blue gape wattles; orbital and
throat skin blue-black; collar plumage spotted; body plumage spotted without
vermiculations; tarso-metatarsus without spurs, scales pentagonal, not in
rows; iris brown; outer margin of secondaries white; furcula blade-shaped;
caecum up to 150 mm; abdominal plumage spotted bluish-white without
vermiculations.

Statistics for quantitative characters (N = 26)
(See p. 65 for statistical comparisons with G. p. schubotzi.)

Character D4 S.D:

Bill length 24,5 mm 17
Wing length 225,6 mm 5.3:
Tarso-metatarsus length 81,7 mm at
Wattle basal width . 10,7 mm £2
Wattle length 11,6 mm 22
Dorsal spot size 10,7 units 1,0
Crest frontal length 24,6 mm 3.1
Crest rear length 46,9 mm 56
Crest central height 31,8 mm G7
Crest basal length . 30,0 mm 2,6
Anterior crest curliness 1,0 0,0
Posterior crest curliness 1,0 0,0
Dorsal black collar OO Y 0,0
Ventral black collar 0:0 7 0,0
Occipital fold ops evA 1D
Ear patch 0,4 mm 0,6
Throat red On, 0,0
Orbital red . C057, 0,0
Dorsal spot number Dies |
Total spot barbs 5,4 0,6
Total within spot barbs 35 0,6

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character
Spot barb blueness .
Chestnut blotch size
Chestnut blotch extent
Distribution

See Figure 14.

xX
0,8

0,0 units

L004

Guttera plumifera schubotzi Reichenow, 1912

Figs 1D, 12A(OTU1), 13(°0TU]1)
Guttera plumifera schubotzi Reichenow, 1912: 320.

Description

S.D.
0,6
0,0
0,0

As G. p. plumifera, except that the base of the nape and an area anterior

to the ear are covered by patches of orange-yellow skin.

Statistics for quantitative characters (N = 71)

(See p. 65 for statistical comparisons with G. p. plumifera)

Character

Bill length

Wing length
Tarso-metatarsus length
Wattle basal width .
Wattle length

Dorsal spot size

Crest frontal length
Crest rear length

Crest central height
Crest basal length .
Anterior crest curliness
Posterior crest curliness
Dorsal black collar
Ventral black collar
Occipital fold

Ear patch

Throat red

Orbital red .

Dorsal spot number
Total spot barbs

Total within-spot barbs
Spot barb blueness .
Chestnut blotch size
Chestnut blotch extent

xX
22,8 mm

227,2 mm

80,7 mm
9,6 mm
8,9 mm

‘10,0 units

24,3 mm
38,4 mm
32,4 mm
25,9 mm

1,0

1,0

COZ

0,0 %
24,0 %
Heysilsreotan

ONO

0:0 FY
2hg2

Dil

eT)

1,6

0,0 units

0,0 %

S.D.
1,4
8,2
3,4
C5
2,4
1,4
5,9
4,8
6,4
2,4
0,0
0,0
0,0
0,0
8,5
2,8
0,0
0,0
2,9
0,6
0,5
0,6
0,0
0,0

THE EVOLUTION OF GUINEA-FOWL 103

Distribution
See Figure 14.

Discussion

The one intermediate specimen between G. p. schubotzi and G. p. plumifera
is more schubotzi in appearance, but the patches of orange on the nape and ear
are much reduced in extent.

Guttera pucherani pucherani (Hartlaub, 1860)

Figs 1E, 12A(OTUS), 13(O0TUS), Tables 4-5 (OTUS)
Numida pucherani Hartlaub, 1860: 341.

Description

Larger than G. plumifera; crown surmounted by a long crest of relatively
curly, downy feathers; well-developed occipital fold of blue-black skin; no
nape and cere adornments; orbital and throat skin red; throat skin folded;
rudimentary, relatively short blue wattles at gape; collar plumage spotted; body
plumage spotted without vermiculations; tarso-metatarsus without spurs,
scales pentagonal, not in rows; iris red; outer margin of secondaries white;
furcula hollow; caecum length up to 150 mm; abdominal plumage spotted
bluish-white without vermiculations.

Statistics for quantitative characters (N = 64)
(See Tables 4-5 for statistical comparisons with other subspecies.)

Character xX S.D.
SUL IShG. 0 a 25,5 mm 1,8
Wing length ee eee ee ae 2 OleOr mn 14,8
Warse-metatarsus length . . . . 3. 6 el 90,2 mm 6,9
Miiasmcubasdiwidth. . . . . .« . . « a 4 9,2 mm 1S)
We SE So a en 2.7) ton 0,7
OS] SOCT.S 4 11,3 units [3
MSiecsminomialiometh . . . . ». « » ws «2 « 19,4 mm 3.5)
LPSS Sar le 1 rr rr a re 29,5 mm Bit
Mecemecntralheisht . . . 9 2 . os 6 let. % 26,4 mm 6,6
Mewemssaienemy 2 2. fe Ol 34,7 mm a
Pemctiom crest Curliness:. . . . . «=. »«» «© « « 1,0 Oo?
Ememem@merest CUrIIMESS . - . - 6 s « ss 20 0,2
Dorsal black collar Py CRIS poe Eee: RAT er end ae Deny A ES
Mimeraieplack-collar =. sw 7,4 % 10,4
Mesias es kl i 74,6 % 195
Ef O20Gf -.0 2 a a no earns 0,0 mm 0,0
“erceih rad. a. ne 9330774 A

SeGCISRCORP MR et et OR a) SS we 95,4 % 2,4

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character X S.D.
Dorsalspotmumber f° = ess Sh, 2 eee 26,0 4,3
Wotalispotibarbstas 23° 74. 2/e) os eee lee 6,3 13
Total. within=spot barbs, 454 3 a eee 4,3 0,7
Spot barb blueness ee Vee, Ones ere Cee a ae 9 0,9
Chestnutbloteh sizes... ee ee ee 0,0 units 0,0
Chestnut blotchrextenG@ ) 45 (29 eh =e 0.07 0,0
Distribution

See Figure 31.

Discussion

From breeding experiments with captive birds, Ghigi (1936) demonstrated
complete interfertility between individuals ascribed to G. edouardi and
G. pucherani. He also found that red throat and orbital skin, characters used
to distinguish G. pucherani, were invariably absent among FI hybrids. This
suggests that these character states may be recessive. Ghigi (1936) further
hypothesized that Guttera edouardi suahelica (Neumann 1908: 14) and
G. e. granti (Elliot 1871: 584), subspecies described from specimens collected in
southern Tanzania, are in fact intergrades between G. p. pucherani and
G. p. barbata. The results of this study support that hypothesis, since pucherani-
barbata intermediates in discriminant analysis (Fig. 13) almost invariably fit
descriptions of these taxa, and the distribution attributed to them falls in the
high total COV region between the suggested parental subspecies (Fig. 30B).

Guttera pucherani verreauxi (Elliot, 1870)

Figs 1F, 12A(OTU3), 13(0TU3), Table 4(0TU3)

Numida verreauxi Elliot, 1870: 300.

Guttera cristata sethsmithi Neumann, 1908: 13.
Guttera pallasi Stone, 1912: 208.

Guttera edouardi schoutedeni Chapin, 1923: 73.
Guttera edouardi chapini Frade, 1926: 139.
Guttera edouardi kathleenae White, 1943: 19.

Description
As G. p. pucherani, except crest longer; less well-developed occipital fold;

orbital skin colour blue; collar plumage black with no vermiculations; spotting
with stronger blue hue; iris brown.

Statistics for quantitative characters (N = 159)
(See Table 4 for statistical comparisons with other subspecies.)

Character xX S.D.

emotive tes. Seat iry gute As, Ce ew a 23,8 mm 6
Wing length OR ay ik ie, OE, 0a SR ee (Pe 11.2

THE EVOLUTION OF GUINEA-FOWL

Character X
Weno-metatarsusiengsth « . £ wok ke 89,2 mm
Piseeesoasdl Wit i a 9,2 mm
ES [SPC eh” ee a ee 2,7 mm
PieuseecmOU size ka 9,5 units
Macveummetarteneth. ke DA i wanes:
SS) Pdi? LE ee 37,4 mm
Meesmecminralheiont =. we 26,5 mm
MeemsIGMOOM ek a a 31,9 mm
Pemceiaisckest CUTIINeSS © . 9. ef lk ll it
Beistomi@r Chest CUTIMESS 2 5 3 6 kk wk ek 2,0
Mepssemisckecollar 3... kl a Dit
DeeMrmOaCKee@at! ge 8 a ea LES
DEC MEEOIC NG te 47,1 %
20 20S pe ee ee ee ne cee 0,0 mm
SOE SC. 1 eee re PT or
EE PSC co ee OO
MRORSESMOLMUMIDEL <6 24,1
DUEL SOC SST a os ane 6,0
formmmaumm-spot barbs . . . . . . . «. 39
POMMIMEHONOINCMESS. 2 . . we lk lk Des)
Micsmiimeiloren Size... kw 0,0 units
@isccmuboloteh extent . . 9 . 3. 2 8 we 8 O02,
Distribution

See Figure 31.

Guttera pucherani sclateri Reichenow, 1898
Figs 1G, 12A(OTU4), 13(0TU4), Table 4(0TU4)
Guttera sclateri Reichenow, 1898): 115.
Description
As G. p. verreauxi, except anterior crest much shorter.
Statistics for quantitative characters (N = 12)

(See Table 4 for statistical comparisons with other subspecies.)

Character x
IRE ey RP 25.2 7am
Der ee ee we ey ee ASO
fmse-metatarsus leneth . . . ke 89,9 mm
iemceaSMWwiGt 2 a we Ree 10,4 mm
Ee S ep” ae ee sar 2,3 mm
MIS IESPOUSZCr 8 OR 11,5 units

Macwainoniailenea: |. 6 kg we 5,8 mm

105

155

S.D.
DED.
10,4
4.5
2
0,5
ee
Zeal

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character X
Grestircar leneth ss 57, ce te eee Oe eee 40,8 mm
Crest centralshetomt.| 4. 2740 See ce Sa ee ee ee 18,5 mm
@rest basalalensth... “5 8. “sea ee 32,1) iam
AntehOr Crest CUMIMESS, 5 Sells, | Stun een ens el
Posterior Chest: cunlmess: 5 ea. aimee ne Den
Dorsal blackecollak =) xs a ee 18,0 %
Ventral blackecolllar 1%) 4) 2) 8G: Seen ee 23,5074
Occipitalfolde Wats Ga 4) ue eae en ae O2SigA
Barpateh. : foe. ee OO a oh lg eee 0,0 mm
WhEOAt Ted: =. BEE aT Oh Ri ne ne a 95,374
Orbitalted. 6) 222 6 ee ey ae ee 0:0, 7,
Dorsal spot numbery. 2.42 6 Ge Ge elles ee Boe ee 23
Lotal:spot barWs "j2k 5 Gaye awe, Sa oe 6,3
Rotalswithmespor barbs ss) aya es | ee eee 4,3
Spot-barbsbluenesss.. . Go. 4G eee See 1,8
Chestnut blotchisize “bk Roe Ses eee 0,0 units
Chesinutiblotchiextent 85) 25 eis, = eae oe 00%
Distribution

See Figure 31.

Guttera pucherani barbata Ghigi, 1905
Figs 1H, 12A(OTU6), 13(0TU6), Table 5(OTU6)
Guttera barbata Ghigi, 1905: 194.

Description

1/6

0,0

As G. p. pucherani, except crest longer; throat and orbital skin blue;
collar plumage black with no vermiculation; body plumage spots occasionally

interspersed with chestnut blotches.

Statistics for quantitative characters (N = 28)
(See Table 5 for statistical comparisons with other subspecies.)

Character X
BillSlemethy.. sae <x Wieu yn Merge i le i, se ees 25,3 mE
Wing lemsth Gf= = . an Oe a ae
iarso-metatarsus tempeh) ).c.w 64. a) 2 ee ee 90,2 mm
Wattle*basalewadthe.s "798-539". nw wh ee Gk 9,4 mm
NVieretlelemethah ey Pie. kw STM og gh we ae 2,8 mm
Dorsalspotisizere.. Aieel en hoo ce GA no Ro 10,9 units
Crest trontaliensthy Vy ee ye ho Ge ee 23,3 mm
Cresimicurlenetlin eG. te Ge “se. LN ne eM ee 32,2 mina
Grest.cemrateneiolt 3.0 a. 2k te 6 se bs ee BES 26,9 mm

Crest basalsienothy 0-5) wn Bee ee ae ee 34,6 mm

Sap),
1,3
11,0
4,5
1,3
0,8
12
a7
3,9
8,5
2,9

THE EVOLUTION OF GUINEA-FOWL 107

Character X SD:
PUMGMOL GREStCUGINESS 6 2. 2 1 . «6 se A 0,3
Memamcrest CURIINESS 2 9. 5 « sh ke 1,9 0,3
Mensigolackicollar 2. °. 3. «ot elu hl 21,4 % 4,5
Meemmeemlack collar. 3. 2 05 6 » 8 s + 20D/ 255)
EGMECMEROIG 2 ye 76,4 % feles
eer kk 0,0 mm 0,0
Sse each 2) Sa ee er a aaa 6,9 25,8
MOCERCOMI fe ES ee 0,3 39
Manairspotmumber 2. 2 « + + 8 + es 2959 4,9
MeveseO@tbarbS: 2... + se ee kk 539 0,7
Mormiavicnim-spot barbs . . . . . + . «= . 4,0 0,5
SaemiathonbleNesS. . 6°. 3 klk ke eS) 0,7
Mmesinmimoloreh Size . 2. wk tk ls 2,7 units ligil
Micsmuigolotch extent 2 2 2. 7 ek le AS 7, 2,6
Distribution

See Figure 31.

Discussion

Ghigi (1936) mentions G. p. barbata specimens with brown irides. This is
further evidence of gene flow between the brown-eyed western and red-eyed
eastern subspecies of G. pucherani.

Guttera pucherani edouardi (Hartlaub, 1867)

Figs 11, 12A(OTU7), 13(OTU7), Tables 4-5(OTU7)

Numida edouardi Hartlaub, 1867: 36.
Guttera lividicollis Ghigi, 1905: 195.
Guttera edouardi symonsi Roberts, 1917: 3.

Description
As G. p. barbata, except crest curlier; occipital fold is of whitish skin;
no throat fold; black collar plumage extensively covered with chestnut blotching.

Statistics for quantitative characters (N = 34)

Character X S.D.
[eli HSICGEID ~ SS earet a aee 25,6 mm 4,9
rier MMe ae he we ee st 25538 mM 7,4
Wearsoometatarsus length. . . . . . 2-2. 4 86,9 mm 4,8
Remletoasal width. . .« .« «7. 2 - is ww 9,9 mm 1,0
SeeeemeCMOtMbet ie oe sw we eS P 2:5. mm 0,6
MRS ESPOIMGIVCR I 0 ee Me a we 9,9 units 1,0
Ciaesemomtawiensth” . 2. 6s wk Oe 22,6 mm 4,7

TSS) PER era) ee en rn 38,0 mm 4,1

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character Xx S.D.
Crest central erates Wan 23.0 it cee we ee ce 25,9 mm 5
Crestibasall lemoehnas hz) 20 a sek ee 32,8 mm 2,4
AMtenlor CreSE.CURNMESS =) a en eee 5 OFF
POStEhIOk CRESL CURMIMCSS omar ye ye en 2 0,3
Dorsal black-collar™ 1 \: Aag)Ancde ke ee ee 214-9 4,8
Ventral black collar 22°...) 09? te Se ey aes 3258 VA 4,7
Occipital fold? a". iva Sh Ree? Se Dae D558) 4 96
Bar pate: -. GN Seok he ar Aare ae 0,0 mm 0,0
Pheo@at red: Byres tee jh iano. aa ee Be 0,0 % 0,0
Orbital Tee). Sey ee ae a a O10 0,0
Dorsab:spot mumber:: 45) Whee a ha pein ty aes ee 2354 4,2
Totalspot barbs.2 2. 2: 7 oe Se ee ee ee ee D52 0,7
otal within=spot,barbs “27 6) 2 a ee ae 35 0,6
Spot barb blMemess se .. Seen La lye Ge eae ae 0,8 0,5
@hestnut blotchisize. 9". 0 ee Ge ee 10,8 units 3,4
Chestnut blotch extent ei.) Ju uae Cott Vik arene eee 342-7 LST
Distribution

See Fisure 31.

Genus Acryllium
Acryllium Gray, 1840

Fig. 8(0G3)
Acryllium vulturinum (Hardwicke, 1834) °-
Fig. 1J
Numida vulturina Hardwicke, 1834: 52.

Description

The largest guinea-fowl species; no crown, nape, throat or cere adorn-
ments; occiput covered by short, dense, downy chestnut-coloured feathers;
rudimentary blue-grey wattles at gape; orbital and throat skin blue-grey;
well-developed collar hackle; body plumage spotted with vermiculations;
tarso-metatarsus usually with bump(s), scales pentagonal, not in rows; iris red;
outer margins of secondaries lavender; furcula blade-shaped; caecum longer
than 200 mm; abdominal plumage blue.

Statistics for quantitative characters (N = 43)

Character X S.D.
BUMeHeCne ee Ae 8 I) eM Ee ine oer ae 28,5 mm 2,0
Wing length Veen Ie, ae ee a eR 2S) 3 aE 10,1
Tarso-metatarsus leneth . oo. .° . . 2 « ©» WOG@aama 6,9

Wattle basakwidth' § 5-0. (Bey Mao! ee gaa & 7,2 mm EA

THE EVOLUTION OF GUINEA-FOWL 109

Character xX S.D:
RULE LEGS aye en 2,2 mm 0,6
Pee SeRUGLUTE MUINOCE fk Ok 1,8 1,4
Tarsal structure length een PRR eT Pn fcc la ee At 2,6 mm 1,9
Distribution

See Figure 10.

Genus Numida
Numida Linne, 1766
Fig. 8(0G10-14)
Numida meleagris meleagris (Linne, 1758)

Figs 1K, 12B(OTU4), 32(O0TU4), Table 6(OTU4)

Phasianus meleagris Linne, 1758: 158.

Numida ptilorhyncha var. major Hartlaub, 1884: 217.
Numida ptilorhyncha omoensis Neumann, 1904: 407.
Numida ptilorhyncha toruensis Neumann, 1904: 410.
Numida ptilorhyncha macroceras Erlanger, 1904: 97.
Numida ptilorhyncha neumanni Erlanger, 1904: 97.
Numida ptilorhyncha var. inermis DuBois, 1915: 18, 27.

Description

Larger than Guttera spp.; crown surmounted by a bony helmet; no occipital
or throat adornments; well-developed, rounded, blue gape wattles; nape
covered by short, downy feathers; orbital and throat skin blue; cere sur-
mounted by a tuft of cartilaginous bristles; collar black, finely barred with
white; body plumage spotted with vermiculations; tarso-metatarsus lacks
spurs, scales pentagonal, not in rows; iris brown; outer margins of secondaries
banded black and white with vermiculations; furcula blade-shaped; caecum
less than 150 mm; abdominal plumage spotted with faint vermiculations.

Statistics for quantitative characters (N = 311)
(See Table 6 for statistical comparisons with neighbouring subspecies.)

Character xX SD:
PRI eS ee em A 25,0 mm 7
See oe AS as 8 2622 om (ey
Masso-mictatatsus length 2. . . . 6 we a 84,3 mm 532
SeeteomoasolewiGth., 9-0 3. 6 we we 14,1 mm 7
Pe mermICI ah beh a 13,5 mm 3,0
Pemrerperecnt ole 2 2 kk lt ee wed Sal oy, Da
mcimicmMROMmAINCHOIA . 5. a 22,6 mm 10,2
Pemiirencanicneti: . 2 6k ee el 11,8 mm 15

memmemcenttalhereht .-.. 2). 8 oh ee 11,7 mm 8,0

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character xX S.D.
elmet basalvlemegin . 5995 e incu eee ee 1952 mim 3:0
Helinet thickness’ 42. a ee a 6,4 mm ei
Cerestructunce ieneth’ . % oa, bee eee 6,0 mm 3,6
Cere structure thickness: == <2 3) ea eee 1,1 mm 0,4
Collar plumage. . ee GIRS, eee, Wade ee ge vee Ds 0,6
Nape filoplume iederie sng ee OP ays ta: eee 14,9 mm ead
Nape filoplume anteroposterior coverage . . . . 3853 fA 17,4
Nape filoplume lateraltcoveracey =) 2) | ae 955657 5,7
Nape filoplume density . . . ir, ate See Ds 0,3
Secondary remex outer web Gemniculation eRe 558 0,7
Wine Covert barrin® | 9 7 2 ee ee eee 155 0,7
Dorsalvermiculation e+ 509) Seka 5 eee ee 3,0 0,7
Worsal spot Sizee "9s Pe eae ths) ee 18,8 units 7,
Distribution

See Figure 46.

Discussion

Two taxa, N. m. strasseni (Reichenow, 1911: 82) and N. m. blancoui
(Grote 1936: 158), have been described from the region in which N. m. galeata
and N. m. meleagris meet. Specimens attributed to these taxa are invariably
intermediate between galeata and meleagris in discriminant analysis, and their
collection sites fall in the region of high total COV between the parental forms
(Fig. 45A). They are treated as intergrades.

Numida meleagris sabyi Hartert, 1919
Figs 1L, 12B(OTU1), 32(;0TU1), Table 7/;0TU1)
Numida sabyi Hartert, 1919: 69.

Description

As N. m. meleagris, except gape wattles red; nape featheration long
filoplumes restricted to the mid-dorsal line; no cere adornment.
Statistics for quantitative characters (N = 4)

(See Table 7 for statistical comparisons with neighbouring subspecies.)

Character X S.D.
Bll enechifher «sy ts.-° ok team ek) Gee ce ne eee 24,0 mm 0,8
Wing lenothit “00 40 ave - t Ge 6 oe SO onoame 10,3
iarso-metatarsusiiensthy «5 “aes! Le an ee 88,3 mm all
Wiattle basaliwidtiie= Gx tan Wea “es eA woe, ee 15,5 mm 1,0
NVacleleneuhe ay ae A Pe a ae 17,8 mm a7

WiatelespemceminOlUGy e: “al Ps ee ee ge 0,0 % 0,0

THE EVOLUTION OF GUINEA-FOWL 11]

Character Xx S.D.
memctinontanieneth . . . .« . . « « s . 22,5 mm 3,0
Pemvencamiemetm . 2 i fo bd 8 Sl 15,0 mm G35
Eemicwcenttal hersht 9 16,3 mm 26
feeimeebusaliiiensth 8. - 2. 6k oe ke 20,0 mm 3,4
BMMCINEMIGKMICSS 2 5 kk a 5,3 mm 0,5
Mecermucimuneieneth 2... 6k kee 0,0 mm 0,0
Mrnoesmctune MMICKMESS 2. 6s ee 0,0 mm 0,0
Collar plumage. . OE Ok es Se ar aA e 25 0,6
Nape filoplume lencia . es ee fe he ot) Lee 34,0 mm Tel
Nape filoplume anteroposterior coverage . . . . 100,0 % 0,0
iINapemloplume lateral coverage . . . . . . . SOA 17,9
Nape filoplume density... Si Se Nails Ate 3,0 0,0
Secondary remex outer web mexmiculanon Se ie 3,8 eS
Dyameecovers barming . . . . 2. .« . ws ws 0,0 0,0
Dersaiayenmiculation . .. . . .« 2. 2 wo we 29 0,6
DOES) SOCE SZ en 7 Se. (Os5)
Distribution

See Figure 46.

Discussion

Although this taxon is represented by only four specimens, its validity
was upheld owing to its isolation and correspondence with criteria set for
subspecies.

Numida meleagris galeata Pallas, 1767

Figs 1M, 12B(OTU2), 32(0TU2), Tables 6-~7(OTU2)

Numida galeata Pallas, 1767: 13, 15.
Numida marchei Oustalet, 1882: 1.

Description
As N. m. sabyi, except smaller with collar plumage grey to blue-grey.

Statistics for quantitative characters (N = 137)
(See Tables 6-7 for statistical comparisons with neighbouring subspecies.)

Character xX S.D.
PCCM me ee) we ae) Ges 22,3 mm 1,8
eMPMCMPOM NTS ee ee a we S| 624 im 14,6
emsopmetalansusseneth 2... ee RR 81,8 mm 6,4
ReeilerbaSalewiGd . 6 ee 14,2 mm 1,9
SES ead 1 a ee re 15,5 mm 29

SremelespemcentONIG a Sony, 4,3

112 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character X S.D.
Felmet irontallenethy 4 ee ee ee 15,5 imam 5,8
iclmet reandenethh) cays. a. 15) wee ee re 7,0 mm 259
Helmet central heisht (4 5. e 5,8 mm 2,9
Helmet-basaljlengthe 4) 3) 20 2 ee ee 18,6 mm 353
Helmet thickmesse 0." SV ee eo a ee ae eee 4,9 mm He)
Cére Structumevlem atlas 1) iy pa ee ce ee 0,3 mm 0,9
Gére structure thickmessiaur.. a5 ees, ee ee eee 0,3 mm 1,6
Collar plumage. . A eae ts yy Re cg eae 0,1 0,3
Nape filoplume leva. LaO Stee in + eee 20,4 mm 4,3
Nape filoplume anteroposterior coverage . . . . DONA 16,0
Nape tloplume lateral coverage’ =). . 2 Seeee TOSS 5303
Nape filoplume density . . ee 2,8 0,4
Secondary remex outer web Tecmiculagan <i Cae 4,4 13
Wine covert baring.) 29 5000 7 ee ee ee 0,6 0,2
Dorsalbvermiculation ». 92). tao ee 3,6 0,6
OKSANSPOb SIZE hy ae et okra Sh 8, ee ene 12,2 units 3,8
Distribution

See Figure 46.

Numida meleagris somaliensis Neumann, 1899

Figs 1N, 12B(OTUS), 32(O0TUS), Tables 6, 8(OTUS)
Numida somaliensis Neumann, 1899: 25.

Description

As N. m. meleagris, except gape wattles blue with red tips; cere tufts
much longer and more numerous; and nape featheration long filoplumes
restricted to the middorsal line.

Statistics for quantitative characters (N = 44)

(See Tables 6, 8 for statistical comparisons with neighbouring subspecies.)

Character Xx S.D.
ler SO eee he cian he eile Rime Me 25,1 mm 1,4
Wine Tenet 3) Oe ee ment 11,0
arso=metatarusmenctiny-) 2 eo eh 2 fe ee 84,2 mm 4,2
Wattle basanites. « a GOW say Bees © cn) ie 13,6 mm 2,4
Wrattlewemetlag Bete. es-C iii yee ie Yen We ee 14,6 mm 353
Wattle per cemtiOlicaine <2 trian 26th ie ne 87,5 7% 14,9
elimet trontalliengtiay. 2 oS re a 20,0 mm 7,6
RSet reat eMm tite OG, ee ee ie Pale! an eee i 10,3 mm 6,0

elmer centralthieicht a ee ee 12,9 mm 12,3

THE EVOLUTION OF GUINEA-FOWL

Character

Helmet basal length

Helmet thickness

Cere structure length

Cere structure thickness

Collar plumage .

Nape filoplume ange
Nape filoplume anteroposterior coverage
Nape filoplume lateral coverage .

Nape filoplume density
Secondary remex outer web Tenmiculation :
Wing covert barring

Dorsal vermiculation

Dorsal spot size

Distribution
See Figure 46.

x
17,4 mm
6,6 mm
13,3 mm
1,0 mm
3,0
15,8 mm
Aled: 7%
53,01 74
2,8
a2
0,8
2,4

23,6 units

Numida meleagris marungensis Schalow, 1884

Figs 10, 12B(OTU6), 32(0TU6), Tables 6-7, 9(;0TU6)

Numida coronata marungensis Schalow, 1884: 105.
Numida marungensis maxima Neumann, 1898: 21.

Description

3

S.D.

39
153
156
0,1
0,7
35
20,7
34,0
0,5
2,0
0,6
0:9
6,5

The largest subspecies; helmet characteristically thicker and longer basally;
long gape wattles, blue with red tips; no cere adornment, and wattles pointed.

Statistics for quantitative characters (N = 97)

(See Tables 6-7, 9 for statistical comparisons with neighbouring subspecies.)

Character

Bill length

Wing length
Tarso-metatarsus length
Wattle basal width .
Wattle length

Wattle per cent blue
Helmet frontal length .
Helmet rear length .
Helmet central height .
Helmet basal length
Helmet thickness

Cere structure length
Cere structure thickness

xX
26,7 mm

283,5 mm

90,2 mm
11,8 mm
16,6 mm
(ERO YA
47,7 mm
13,4 mm
18,7 mm
39,2 mm
12,4 mm
0,3 mm
0,3 mm

S.D.
1,4
a
4,7
155
3,1
8,5
8,5
ay
3,8
337
256
0:7
0,9

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character X S.D.
Collar plumage. . Uae cay ter mayer gre Ps 2) 0,7
Nape filoplume ‘engi. Ae fl, wore Ai see a 26,3 mm 3;3
Nape filoplume anteroposterior suas Se eges oe T1533) Ya se)
iINape hlloplumelateralcoverase =) eee 2901 / 75S
Nape filoplume density . . ee eee D5) 0,4
Secondary remex outer web sepinioulle on Zncleny aera 0,9 1,0
Wine covert Daring. «2. ¢ 2. 2 . Re eee 0,0 0,0
Dorsal vermiculation <2 S45 32> 45 5. eee LS 0,8
IDOrsal SPOtsizZeug —o— as df “wees ee ee eee 23,5 units 4,5
Distribution

See Figure 46.

Discussion

N. m. frommi (Kothe 1911: 13) and N. m. rikwae (Reichenow 1900: 40)
have been described from the region between N. m. marungensis and N. m.
mitrata, and N. m. callewaerti (Chapin 19326: 1) from the region between
N. m. galeata and N. m. marungensis. Specimens attributed to these three forms
are invariably intermediate between two subspecies in discriminant analyses,
and their collection localities fall within regions of high total COV (Fig. 45A).
They are treated as intergrades.

Numida meleagris reichenowi Ogilvie-Grant, 1894
Figs IP, 12B(OTU8), 32(;0TU8), Tables 6, 8;OTU8)
Numida reichenowi Ogilvie-Grant, 1894: 536.

Description

Similar to, and nearly as large as NV. m. marungensis; except wattles rounded
and red, helmet taller and sabre-shaped, and nape featheration less dense.

Statistics for quantitative characters (N = 121)
(See Tables 6, 8 for statistical comparisons with neighbouring subspecies.)

Character x S.D.
DiMlMlenote A? WP vad captinae cahpsal le Seeeae Mere Semeye 24,7 mm 1,6
Wing length a ee eee ee Mee 252.2) ean 9,1
iharso-metatansus lenethe . a. oly Wen oe ee 90,7 mm 11,4
Wattle basalivviditii.,, “s se “aia, ae a, av aah ve os 11,9 mm 1,4
Wrattlelenetht yo" ac fe «Gallen. eas ahha <P eg, (ee 12,8 mm Vis)
Wattle percent ble. ois )aer ar at acer ae ae We) 7. 29
Fieinettromtallemeth 4. 4) sw sa a a 44,2 mm 10,9
elimetrearlenet se =) oO. = Sue = Suh 29,5 mm 9,8

Ficlimet centralhecicht. 2. = “ae = oS 4 eee 29,5 mm 9,0

THE EVOLUTION OF GUINEA-FOWL 115

Character Xx S.D.
Meumetbasaliensth =. . 2. . Ww . ww ee 24,9 mm 29
BrcmMeetMeKNeSSy 6 ee we 8,3 mm 17
Memecmuctineleneth 2 2. eee 0,7 mm 1,8
Meremsmucture thickness . . © . . 2 os es 0,4 mm 0,8
MOMMIES 218) 0,5
iNapemloplumelensth. . . . ee ee eA 20,5 mm 3,6
Nape filoplume anteroposterior coverage . . . . O94 49,2
iNepealoplume lateral coverage . . . . . «= . 30,0 % DAES
Nape filoplume density . . vee Sg ote 2 0,6
Secondary remex outer web o iniculanion eee ee 23) ee)
meseovernt barrime 2 6 6 kk ke O2 0,5
Moralvermeculation . . . «1. . .« .« « . 2,4 0,7
AGSESOINSIZE 5 ek 23,2 units 35
Distribution

See Figure 46.

Discussion

N. m. intermedia (Neumann 1898: 21), and N. m. ansorgei (Hartert 1899:
331) have been described from the transition area between N. m. meleagris
and N. m. reichenowi. N. m. uhehensis (Reichenow 1898a: 88) has been described
from the transition area between N. m. mitrata and N. m. reichenowi. For
reasons given in other such instances, these forms are treated as intergrades.

Numida meleagris mitrata Pallas, 1767

Figs 1Q, 12B(OTU7), 32(O0TU7), Tables 6, 8-10(OTU7)
Numida mitrata Pallas, 1767: 18.

Description

Smaller than the last two subspecies; phenotype as N. m. marungensis
except that helmet less well developed.
Statistics for quantitative characters (N = 293)

(See Tables 6, 8-10 for statistical comparisons with neighbouring sub-
species. )

Character 4 S.D.
ES Teng." i on re eer 25,7 mm KS
Ser MeMOUME I ele wt ae | 2A 8,2
Narno-metatarus length 2... ee 86,1 mm 4,2
Breulemoasa@lwidti. 2 6 ow ek 10,9 mm 163)
Pr micwlemetni ee ek 16,8 mm 3,0

Boil pemicentDMe Fo 2 wk te 64,6 % O83)

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character X S.D.
Helmet irental lemeth 2. 2) Go er. eee 38,3 mm SLs
Helmet neartenetine .. Wie os a ee 18,8 mm Syl
Felmet centralheight sede) ae ee ee 19,4 mm Je
Ricimetebasalmlengtha “awe i a ae Se 23,3) taiia 3,8
FISiMetthickMessiyos 7% AG eo cok ew Oe het ee 8,3 mm 2,0
Cere Structinenleneth 4s, 1 ee ee 0,2 mm 0,8
Gere structirethickmess. = 319 > 2955 Wale 0,2 mm 0,7
Collar plumasey, We 15. 20 Sy - a & Sl 0,6
Nape tiloplumetensth: © 2 <2 ih) «as =) sae ae 23,0 mm 38)
Nape filoplume anteroposterior coverage . . . . 59a oeT
Nape floplumedateralicoverase: 9 5 =) = “eee 23,0074 8,3
Nape filoplume density . . h5 cule DES 0,7
Secondary remex outer web senmieniagon £5) Sa eae iPS 3
Wineecovert Darume 2 oy 20a ue Ra) el eae eee 0,0 0,0
Dorsal vermiculaGion YO. Seis Le 0,7
Dorsal Spotsize: i+ TU FeO eee ee eee 19,4 units 4,5
Distribution

See Figure 46.

Numida meleagris coronata Gurney, 1868

Figs 1R, 12B(OTU10), 32(0TUI10), Table 10(0TU10)

Numida coronata Gurney, 1868: 253.
Numida transvaalensis Neumann, 1899: 26.
Numida papillosa limpopoensis Roberts, 1924: 77.

Description

As N. m. marungensis, except smaller overall size; decidedly thinner and
taller helmet, collar plumage more streaked than barred with white.
Statistics for quantitative characters (N = 136)

(See Table 10 for statistical comparisons with neighbouring subspecies.)

Character xX S.D.
BMPR Ot ey ek td oe ge eh 2531 mama 7
Wing length a RE ig a ae aS oo Wh Ge | a 8)
Parso-metatansus lemethy “5.5 ° =@ i& Get ee 85,6 mm 552
Wattle basalewidthiv. et! Ly Ge ae 2 els PR 11,0 mm SS
Wiattle tenet ey ac acy Uae Be a Le Sk et 18,7 mm 3,2
Wale percenivbluc. .. etl ar aw An aS ol Se 10,0
Melmetirontaldenethy {18 se 7 Tete Oe 54,8 mm 9,6
icimetteamiemetiane ie “wel eo ou oo Awl ew) el” wah oe 25,1 mm 5,8
lncimet central mersat’ .. Fie eo e e 24,7 mm 5,3

Pelmetbasailenoen io Ge en ee a ee 25,8 mm Zo

THE EVOLUTION OF GUINEA-FOWL 117

Character x S.D.
BACMMGEEMIGKMESS Ph we 8,8 mm Oa
MGeESeRMCIURe (EMC oo ae 0,4 mm 0,7
Mcrcesenuctune tMIGKMESS. 2 5. kk 0,3 mm 0,4
Collar plumage. . RT ee eM i Pree na 4,2 Ji
Nape filoplume lene. Bee eS sia! ED ag chen 18,5 mm 4,6
Nape filoplume anteroposterior coverage . . . . 40,0 % 22,8
iINapewmleplume lateralcoverage . . . . . . . be 304 4,9
Nape filoplume density . . Te ios ee 0,8
Secondary remex outer web Penniculanion Wee, a Aue Des 1,4
Puiemcovert DaTTIN® 5 6 2 ok ye oe ek 0,0 0,0
Monstevermiculation 2. 5. 5 eee a 2,6 0,6
(OS, SSCE SZ a ae 20,4 units 5,6
Distribution

See Figure 46.

Discussion

Numida meleagris papillosa (Reichenow 1894: 145) has been described from
the region south of Lake Ngami, i.e. the transition area between NV. m. damarensis
and NV. m. coronata. For reasons given in other instances this form is taken to
be an intergrade.

Numida meleagris damarensis Roberts, 1917

Figs 1S, 12B(OTU9), 32(0TU9), Tables 9-11(OTU9)
Numida papillosa damarensis Roberts, 1917: 2.
Description

As N. m. coronata, except well-developed papilli at cere; collar spotted;
helmet less well developed.

Statistics for quantitative characters (N = 102)
(See Tables 9-11 for statistical comparisons with neighbouring subspecies.)

Character x S.D.
allo es a Wa 24,6 mm 1,6
Wing length Me er, eae erie ee TR ait ae oh Se anita 16
iiaeso-metatarsus lensth 2. 2... C. 83,4 mm 3,9
Piavlesmasalowidtly; <0 ee ee 1 noma 188)
SvciMememeihe fo 6 kk a 18,9 mm DES
Deaulcapemeent blue. 2. we eG 64,4 % 8,5
memichimontalicneth 2. 9.3 ke Bw eed 45,2 mm 1
PicMeincammlCnebhye NY yy he te we 22,9 mm 4,6

Incimcecembralheieht =. 0. os 8. Ge ak 22.3- mm 4,3

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character x S.D.
Helmet basalilemeth@ 9% “4.5 eee 19,7 mm Syl
Flelmet UhickineSsisy © 0) 22) pen oe) ae ees ree 9,2 mm 1,3
Cereistructunelemethin.. Ua 4 eee oe 3,7 mm hs)
@eke Simuctunestinickmesss 2) ee ene 229 sia 0,3
Collar plumage. . git Htoy (ogee. cee 4,9 1,0
Nape filoplume lena - ee oe ee 12,3 mm 8,5
Nape filoplume anteroposterior coverage . . . . 24,7 % DD
Nape filoplumetlateralicoverace ene ON ay 8,0
Nape filoplume density. . ee 0,5 0,7
Secondary remex outer web ceaaioniauon es 358 1,4
Wins covert-barting ) 2 xe, ee 0,0 0,0
Dorsaltvermiculation’) 49.) 4). a ee 2,6 0,5
DorsaluSpOt:SiZe® ciens oo econ Se Se ee 22,5 units 8,5
Distribution

See Figure 46.

PHYLOGENY

PRIMITIVE AND DERIVED CHARACTER STATES

The following hypothetical primitive-derived sequences are postulated for
the characters listed in Appendix 1. The number in parentheses following each
character name is its number in Appendix 1.

Crown (1), occipital (2), nape (3), and throat (6) adornments

All francolins and guineafowl-phasianid hybrids have feathered crowns,
occiputs and napes. Very few francolins have crests, and then only rudimentary
ones. No francolin has a throat fold. Therefore, a naked crown, nape or occiput
is taken to be derived relative to the feathered condition. Among guinea-fowl
with feathered crowns, a well-developed crest of downy feathers is taken to be
derived relative to a short crest of downy feathers, and a well-developed crest
of bristly, erect filoplumes derived relative to a well-developed crest of downy
feathers. The latter assumption is based on the idea that a modification of a
downy, drooping crest to an erect bristly crest allows species or individual
recognition in social encounters. Also, among guinea-fowl which have these
regions unfeathered, any likely secondary elaboration of the unfeathered con-
dition (e.g. a helmet, long filoplumes, patches of skin of contrasting
colour, folds of skin) is taken to be derived, since these structures may be
adaptive in a social (individual and species identification) or physio-ecological
(thermoregulation, crypsis) context (Brown 1963; Reynolds 1977). Finally,
folded throat skin is taken to be derived relative to the unfolded condition.

THE EVOLUTION OF GUINEA-FOWL 119

Orbital (4), throat (5), and gape wattle (8) skin colour

In nearly all francolins, areas of naked skin on the head and throat are
red. Therefore, red is taken to be the primitive condition relative to any other
colour for the above characters.

Gape adornment (7)

No francolin has a gape wattle, and guineafowl-phasianid hybrids have at
most rudimentary wattles. Therefore, well-developed gape wattles are taken
to be derived.

Cere adornment (9)

No francolin or guineafowl-phasianid hybrid has an adornment at the
cere. Therefore, any cere adornment is taken to be derived.

Collar (10), body (11) and abdominal (18) plumage

The plumage of francolins and guineafowl-phasianid hybrids is usually
dark, often streaked, barred or vermiculated with lighter colours. Moreover,
none of the above possesses an elaborate collar-hackle. Therefore, among
guinea-fowl, white or blue plumage is taken to be derived relative to dark
plumage, and spotted plumage derived relative to barred, streaked or vermicu-
lated plumage. Among guinea-fowl with spotted plumage, spotted plumage
without peripheral vermiculation is taken to be derived relative to that with
vermiculation. Also, regardless of plumage pattern, an elaborate collar hackle
is taken to be derived relative to an undifferentiated collar.

Tarso-metatarsal scales (12) and adornment (13)

All francolins have their posterior tarso-metatarsus covered by small
imbricated scales in rows. In nearly all francolins, the male at least possesses
spurs. Therefore, among guinea-fowl any modification of the spurred condition
(e.g. naked tarsi or tarsal bumps), and any scalation pattern other than that
described above are taken to be derived conditions.

Tris colour (14)

Nearly all francolins have brown eyes. Therefore, among guinea-fowl, a
red iris is taken to be derived.

Outer margins of secondaries (15)

In most francolins the secondaries are brown to grey, sometimes faintly
streaked or vermiculated with white. In only a few francolins do the outer
margins of these feathers appear to be strikingly different from the general
body plumage. Therefore, among guinea-fowl, secondaries that contrast with
the general body plumage pattern, 1.e. character states 2-4, are taken to be
derived.

Furcula (16)

All francolins have blade-shaped furculas. Therefore, a hollow furcula is
taken to be derived.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Caecum length (17)

In francolin and guinea-fowl of the genera Numida and Guttera caecum
length is about 16 per cent that of the large and small intestines combined
(Beddard 1898). There is no information on caecum length for Agelastes spp.
In Acryllium yulturinum, the length of this organ is more than 23 per cent
that of the intestines (Beddard 1898). Therefore, among guinea-fowl a relatively
long caecum is taken to be derived.

PHYLETIC ANALYSIS

The most parsimonious guinea-fowl phylogeny based on primitive-derived
character sequences is given in Figure 47. In this figure, the four shared derived
character states comprising character suite | delineate the subfamily Numidinae
from francolin-like phasianids. These derived character states, common to all
guinea-fowl taxa (Ghigi 1936), and their primitive counterparts (in parentheses)
are:

1. large size (small size);

2. third and fourth sacral vertebrae with robust transverse processes (only
third vertebra with such a process);

3. second metacarpal lacks a backward process (process present);

4. largely naked head with at least rudimentary wattles (feathered head, and
no wattles).

Agelastes sp., with their preponderance of primitive character states,
probably more closely resemble the proto-guinea-fowl than does any other
member of the subfamily. These two species share only one derived character
state, white abdominal plumage. Adult A. niger (Fig.. 1B) shows primitive
character states for all characters investigated except for its naked occiput.
The white abdominal plumage linking it to A. meleagrides (Fig. 1A) is present
only in juvenile birds. It is also possible that the additional derived character
states attributable to Agelastes due to the naked head and neck of A. meleagrides
are the result of convergent evolution, and do not suggest any closer affinity
to other guinea-fowl genera.

Character suite 2 consists of three shared derived character states (spotted
body plumage, non-imbricated, pentagonal scalation of the posterior tarso-
metatarsi, unspurred tarso-metatarsi) which link the three remaining genera.
Character suite 3 consists of three shared derived character states (unfeathered
crown, blue orbital and throat skin) which link Acryllium (Fig. 1J) and Numida
(Fig. 1K-S). Character suite 4 consists of four shared derived character states
(folded occipital skin, white outer secondary margins, body plumage spotted
without peripheral vermiculation, hollow furcula) which link the two Guttera
species (Fig. 1C—I), and distinguish them from Acryllium and Numida. Character
suite 5 consists of four derived character states (helmeted crown, naked occiput,
well-developed wattles, cere with tufts or papilli) which distinguish Numida
Fig. 1K-S) from Acryllium (Fig. 1J). Character suite 6 consists of three

THE EVOLUTION OF GUINEA-FOWL 194

Agelastes Guttera Acryllium Numida

Fig. 47. An hypothetical phylogeny for the Numidinae. Each monophyletic lineage is charac-
terized by a suite of derived character states (Mf), the primitive counterparts of which (LJ)
are found in members of the co-ordinate sister-group.

derived character states (hackled collar, long caecum, red eye) which distinguish
Acryllium from Numida.

SPECIATION

ORIGIN AND DERIVATION OF THE NUMIDINAE

Several hypotheses have been offered concerning the origin and evolution
of the Numidinae. Ghigi (1936) states that the proto-guinea-fowl originated
in Africa, and probably was derived from a francolin ancestor. This hypothesis
is based on the fact that guinea-fowl are found only in Africa (Arabian and
Malagasy populations being probably the result of introductions by man),
and phenotypic similarities between the most primitive guinea-fowl (Agelastes
spp.) and Francolinus spp. Cracraft (1973), in support of his hypothesis of a
Gondwanaland origin for the Galliformes, suggests a North American origin
from New World quails. Olson (1974), suggests a possible Asiatic origin from a
pheasant-like bird, based on analysis of a single Eocene fossil femur from
Mongolia. This bone is intermediate in shape between femurs of extant
pheasants and Agelastes niger. However, it is only about 70 per cent the length
of an Agelastes femur, i.e. well within the size range of many extant francolins
(Mackworth-Praed & Grant 1952, 1962, 1970).

There is no compelling evidence favouring any of these three hypotheses.
When, and if, sufficient information comes to the fore, it will probably support

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

elements of all three. Accordingly, the working hypothesis taken herein is:
the Numidinae are derived from a francolin-like Asiatic phasianid; but evolution
and radiation of extant guinea-fowl has occurred solely in Africa.

EVOLUTION OF GENERA

The first opportunity for colonization of Africa by Asiatic faunal elements
arose with the mid-Miocene (c. 17-18 m.y.B.P.) union of the African and Asian
plates (Axelrod & Raven 1978). At this time, forest was much more extensive
in Africa than at present, possibly even exceeding limits depicted in Figure 5.
However, the colonization corridor connecting these two continents was
covered by relatively arid savanna vegetation in the mid-Miocene, and has
almost certainly not had more lush vegetation since then (Axelrod & Raven
1978). Thus, any Asiatic ancestral guinea-fowl was probably a bird which
lived in savanna habitat, and, upon its arrival in Africa, encountered vast
forest, and much less extensive savanna adaptive zones. Since Agelastes spp.
possess so few derived character states (see above), it is likely that radiation of
proto-Agelastes into the forest took place soon after colonization.

With the joining of the two continents, the mild climate that favoured
widespread forest vegetation throughout the late Cretaceous and early Tertiary
began to deteriorate. Africa became progressively more arid; and savannah and
desert-steppe biomes expanded, at the expense of forest, throughout the latter
Miocene and Pliocene (Axelrod & Raven 1978). Moreover, this period was
characterized by widespread uplifting, rifting and tectonic activity adding
considerable topographic diversity to the continent, and partitioning its
expanding and contracting biomes (Axelrod & Raven 1978). These conditions
would have favoured radiation in and into expanding forest-edge, savanna
and desert-steppe biotopes; and it is possible that proto-Guttera, Numida, and
Acryllium were the result of such Mio-Pliocene radiations.

EVOLUTION OF SPECIES AND SUBSPECIES

After the uniformly arid Pliocene, the relatively rapid wet—dry climatic
fluctuations and continuing rifting and mountain building during the Pleistocene
provided additional opportunities for radiation in Africa (Moreau 1966;
Hamilton 1974; Livingstone 1975; Axelrod & Raven 1978). Expanding
forest and wetlands in relatively moist phases would have divided savanna
biome into more or less isolated tracts, and restricted desert to relatively small
refugia in Somalia, northern and south-western Africa. If wet-phase forest
and wetlands were distributed as in Figure 5, Acryllium vulturinum (Fig. 10)
and sub-Saharan subspecies attributed to Numida meleagris herein (Fig. 46)
would have had the opportunity to diverge in isolation, since portions of their
present-day ranges are encompassed by isolated tracts of savanna and desert—
steppe refugia. Also, expanding forest may have favoured a second radiation
into relatively widespread lowland forest, culminating in Guttera plumifera.

During arid phases, forest-living guinea-fowl, and Moroccan N. meleagris,

THE EVOLUTION OF GUINEA-FOWL 123

would have been restricted to island-like refugia. If vegetation were distributed
as in Figure 6, N. m. sabyi (Fig. 46), Agelastes meleagrides (Fig. 10), A. niger
(Fig. 10), Guttera plumifera plumifera and G. p. schubotzi (Fig. 14) and, to a
lesser extent, forest-edge taxa (subspecies attributed to G. pucherani) had the
opportunity to diverge in allopatry, since portions of their present-day ranges
would have had access to refuges of suitable biotope. Thus, we need look no
further than the Pleistocene for biogeographic events which could have allowed
allopatric evolution of extant guinea-fowl species and subspecies.

BIOGEOGRAPHY
RESULTS

A map of hypothetical African avifaunal zones drawn from evolutionary
patterns found in guinea-fowl is given as Figure 48. Zonal boundary lines in
this map agree remarkably well with those in Chapin’s (1932a) faunal map
(Fig. 49) based on distribution patterns of “many species and races of birds’,
and with species and subspecies boundaries in distribution maps of selected
francolin species and subspecies (Figs 50-51). In the hypothetical avifaunal
map, subregion boundaries are those of the relatively phenotypically homo-
geneous (i.e. no subspecies) genera, Agelastes and Acryllium (Fig. 10). Provincial
boundaries coincide closely with those of species, and district boundaries with
those of subspecies (Figs 14, 31 and 46). Chapin neither lists the taxa whose
ranges form the basis of his map, nor specifies criteria used in distinguishing
subregions, provinces and districts. The hypothetical map differs markedly
from Chapin’s in that it:

1. restricts the forest subregion (commonly labelled @) in Figs 48-49) to an
area somewhat larger than Chapin’s province no. |;

2. divides his province no. 4 into eastern and western, rather than northern

and southern districts;

divides his district 1B into two districts (1A and 1B in Fig. 48).

reapportions territory within the commonly labelled province no. 6;

divides his district no. 6E into two provinces (7 and 8 in Fig. 48);

does not recognize montane provinces.

a

The only differences between the hypothetical map and the francolin
distribution maps are relatively minor shifts in boundary lines, and a still
finer subdivision of districts by francolins. Also, district boundaries, which
Separate guinea-fowl subspecies, often delimit francolin species.

DISCUSSION

It is impossible to resolve differences between the two avifaunal maps in
zonal boundaries and hierarchical assessments, since Chapin does not specify
the data base and methodology underlying his map. His decision to unite the
continuous block of lowland forest with the surrounding forest—savanna
mosaic to form subregion @) (Fig. 49) is probably due to the abundance, in

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

exes $Ubregions

provinces

lA ——-—- districts

Fig. 48. Hypothetical African avifaunal zones based on evolutionary patterns found in
guinea-fowl.

the latter, of relict patches of lowland forest and gallery forests, which provide
suitable habitats for forest birds. Indeed, when distributional data are lacking
or equivocal, Chapin (1932a) and other zoogeographers (e.g. Davis 1962;
Moreau 1966) seem to have relied on the distribution of vegetation as a predictor
of bird distributions. The division of Chapin’s province no. 4 and district 1B
into east—west districts in the hypothetical map is due to effects (on guinea-fowl
evolution) of probable past forest—wetland and savanna barriers that bisected
these zones during the Pleistocene (see Figs 5-6). The reapportionment of
territory to districts in the commonly labelled province no. 6 may reflect a lack
of clear-cut avian distributional patterns within that province, a possibility
already suggested by Benson & Irwin (1966). The necessity of partitioning
Chapin’s district no. 6E was anticipated by that author (Chapin 1932a: 89),
and has been done by other authors (Moreau 1952; Davis 1962). The lack of
montane avifaunal zones in the hypothetical map is certainly due to the sub-
montane altitudinal limitation of guinea-fowl.

THE EVOLUTION OF GUINEA-FOWL (25

@) == subregions

] —— provinces

1A ---7— districts

Fig. 49. African avifaunal zones based on analysis of bird species and subspecies distributions
(after Chapin 1932a).

CONCLUSIONS

The results of comparisons of the hypothetical avifaunal map with Chapin’s
map and francolin distribution maps suggest three tentative conclusions, which
can serve as hypotheses in future evolutionary and biogeographic studies.

1. Distribution patterns found in guinea-fowl can be used as models for
broad patterns exhibited by African bird species and subspecies other than those
dependent on montane habitats.

2. At least some francolin species and subspecies have evolved as a result
of factors that have been important in the evolution of guinea-fowl.

3. Physical and ecological barriers which have allowed only subspeciation
in guinea-fowl have been sufficient to bring about speciation in francolins.

The first hypothesis is being tested by a cluster analysis (Hagemier &
Stults 1964; Sneath & Sokal 1973) of 119 equally sized areas of Africa according
to 1 099 passerine species and well-marked subspecies in Hall & Moreau (1970)
(Crowe in prep.). The preliminary results of this study, summarized in Figure 52,
are consistent with that hypothesis. The second and third hypotheses can be
tested, if patterns of character variation in francolin taxa are analysed using

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

——Subspecies boundaries
— species boundaries

Y extralimital areas

=| zones of sympatry

Fig. 50. Distributions of selected non-forest francolin species and subspecies (after Hall

1963 and Mackworth-Praed & Grant 1952). A. Francolinus bicalcaratus ayesha. B. F. clapper-

toni. C. F. icterorhynchus. D. F. leucoscepus. E. F. afer cranchii, intercedens and harterti.
F. F. hildebrandti. G. F. hartlaubi. H. F. adspersus. 1. F. natalensis. J. F. capensis.

methodology outlined herein, and the results are compared to those for
guinea-fowl.

SYNTHESIS

Taxonomy, phylogeny, speciation and biogeography are intimately related
aspects of guinea-fowl evolution. Their necessary separation under different
headings in the present study has been somewhat detrimental to the under-
standing of each. Accordingly, this section attempts to synthesize the author’s
conception of evolution in guinea-fowl (summarized in Fig. 53).

Guinea-fowl are characteristically African birds. Although the likely
ancestral guinea-fowl was an Asiatic francolin-like phasianid which could live
in arid savanna habitat, the evolution that has led to extant guinea-fowl taxa
occurred solely in Africa. Moreover, biogeographic patterns derived from

THE EVOLUTION OF GUINEA-FOWL 127

a ——

Ee Re,
<r Gee eae
——— Sy ay

= Francolinus ahantensis

ESTE squamatus squamatus

[J F.s. schuetti

Fig. 51. Distributions of selected forest-living francolin species and subspecies (after Hall 1963).

guinea-fowl species and subspecies boundaries closely parallel broad patterns
found in African birds as a whole. The Asiatic ancestral guinea-fowl probably
traversed the arid-savanna corridor linking Asia and Africa soon after the
mid-Miocene union of the two continents. This savanna-living bird encountered
an African continent dominated by forest, possibly unoccupied by potential
competitors. Such conditions favoured radiation into the forest, and it is
likely that Agelastes, the most primitive (i.e. most francolin-like) guinea-fowl
genus is a result of an early radiation into forest.

Relatively soon (on a geological time scale) after this successful invasion
of forest, the climate of Africa became more arid. Throughout the latter Miocene
and Pliocene savanna and desert biomes expanded at the expense of forest.
Such a situation favoured radiation in non-forest biomes, and into the expanding
forest-edge biotope, and it is possible that proto-Numida, Acryllium and

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

@) ee subregions

1 —— provinces

1A ----— districts

Fig. 52. Hypothetical African avifaunal zones based on a cluster analysis of 119 equally
sized blocks of Africa according to 1 099 species and subspecies in Hall & Moreau (1970).

Guttera were the result of such radiations. The relatively uniformly arid con-
ditions of the Pliocene subjected these four lineages to strongly divergent
selective pressures, and it is probable that the genera recognized herein were
already well defined at the beginning of the Pleistocene.

The arid climate of the Pliocene was replaced by a fluctuating wet—dry
climate in the Pleistocene. These climatic fluctuations had profound effects on
the distribution of African biomes. During moist phases, the forest biome
expanded considerably beyond its present extent, partitioning non-forested
biomes into more or less isolated tracts. Desert biome was confined to relatively
small areas, and savanna biome bridged the western Sahara, allowing dispersal
of N. meleagris into north Africa. Sub-saharan subspecies of N. meleagris are
the result of divergence in these wet-phase isolated tracts. Also, expanded forest
during mesic phases could have allowed a second radiation into lowland forest,
culminating in Guttera plumifera. During arid phases, the forest contracted
into island-like refugia, and N. m. sabyi was isolated, and presumably diverged
from sub-Saharan populations. The species Agelastes meleagrides and A. niger,

129

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THE EVOLUTION OF GUINEA-FOWL

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130 ANNALS OF THE SOUTH AFRICAN MUSEUM

and subspecies in the genus Guttera, are also a result of divergence in these
refugia. The fact that Guttera subspecies are much more well marked than are
those of N. meleagris suggests that isolation in forest refugia has been more
effective than in tracts of savanna partitioned during wet phases.

ACKNOWLEDGEMENTS

I am grateful to Mr P. Clancey of the Durban Museum and Art Gallery,
Dr F. Gill of the Philadelphia Academy of Natural Sciences, Mr M. Irwin of
the National Museum of Rhodesia (Bulawayo), Dr A. Kemp of the Transvaal
Museum, Dr M. Louette of the Musee Royal de lAfrique Centrale,
Dr R. Paynter of the Museum of Comparative Zoology, Dr L. Short of the
American Museum of Natural History, Dr D. Snow of the British Museum
(Natural History), and Major M. Traylor of the Field Museum of Natural
History for allowing me access to the guinea-fowl in their institutions’ collec-
tions. Mrs J. Adams painted the guinea-fowl shown in Figure 1. Mr S. Piper
helped with contour maps. Dr L. Throckmorton provided initial encouragement
to begin this analysis. Prof. W. R. Siegfried provided the necessary stimulus
to finish the analysis. Mr R. Brooke commented critically on several drafts of
the paper. My wife, Anna, helped in all phases of the study, particularly with
illustrations and data processing. The De Beers and University of Cape Town
Computer Centres gave valuable advice on data processing and computer time.
Generous grants from De Beers Consolidated Mines Ltd and the University
of Cape Town were essential to the project. The inclusion of Figure | was
made possible by a grant from the University of Cape Town Editorial Board.
I should like to thank the South African Museum for publishing this paper.

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132 ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE EVOLUTION OF GUINEA-FOWL 133

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APPENDIX |

Qualitative characters and character states analysed in this study.

Character name States

Crown adornment . : . 1 -—short crest of feathers
2 — long crest of feathers
3 — none, only naked skin
4 — bony helmet

Occipital adornment . . . 1 —short, dense, chestnut-coloured downy feathers
2 — long filoplumes confined to a mid-dorsal line
3 — fold of blue to black skin
4 — fold of whitish skin
5 — none, only naked skin

Nape adornment : , . 1 -—short downy feathers
2 — long filoplumes confined to a mid-dorsal line
3 — patch of orange-yellow skin at base, similar

patch anterior to ear

4 — none, only naked skin

Orbital skin colour. b . 1 — pink to red

2 — light blue to black
Throat skin colour. , . 1 — pink to red

2 — blue to black
Throat adornment. : . 1—no fold of skin

2 — fold of skin
Gape adornment : ; : 1 — rudimentary wattles

2 — well-developed and pointed wattles

3 — well-developed and rounded wattles
Gape wattle colour . d . 1-— pink to red

2 — blue to blue-grey

3 — blue with red tips

Cere adornment . : : eo none
2 — cartilaginous tufts or papilli
Collar plumage . : : . 1 — black with faint vermiculations

2 — black with no vermiculations
3 — black, finely barred with white
4 — grey to blue-grey

5 — spotted
6 — well-developed hackle
7 — white
Body plumage . : : . 1 — black with vermiculations

2 — spotted with vermiculations
3 — spotted without vermiculations
4 — as 3, with chestnut blotching between spots

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

No. Character name States
12 Tarso-metatarsal scales . . 1 —imbricated and in a row
2 — pentagonal, not in rows
13 Tarso-metatarsal adornment . 1 W— spurs or bumps
2 — none
14 Iriscolour . : : : . 1 — brown
2 — red
15 Outer margins of secondaries . 1 -— brown to black with faint vermiculation
2 — white

3 — lavender
4 —alternating bands of black and white with
varying degrees of black and white vermicula-
tions
16 + Furcula A ee bs: . . 1 — blade-shaped
2 — hollow, cup-shaped; found in guineafowl with
long crests (Chapin 1932a)
17 Caecum length . 4 5 ‘ 1 — up to 150 mm
2 — greater than 200 mm; in all specimens con-
forming to the description of Acryllium
vulturinum (Beddard 1898)
18 Abdomen plumage . : . 1 — white
2 — blue
3 — white spots with faint vermiculations
4 — bluish-white spots without vermiculations

APPENDIX 2

Quantitative characters analysed in this study. See Figures 1-4 for a pictorial representation
of many of the characters.

No. Name Units Description
1 Billlength . : ; : mm the chord measured from the base of the
cere to the tip of the maxilla
2 Wing length ; ' : mm the chord of the unflattened folded wing

from the farthest anterior tip of the wrist
joint to the tip of the longest primary

3 Tarso-metatarsus length . mm the diagonal chord from the posterior point
of articulation of the tarsometatarsus with
the tibia to the most distal undivided tarsal
scute on the dorsal surface of the middle toe

4 Wattle basal width .. mm the chord from the most anterior to posterior
points of juncture of the wattle with the
cheek

ae Wattle lencth 995 ; mm the chord from the juncture line of the wattle
to the most distal point along the wattle
margin

6 Wattle percent blue... Wh a subjective estimate of the surface area of
the wattle covered by blue pigment. It is
possible to assess the amount of this colour
in preserved material since, although natural
colour is lost soon after death, the demar-
cation between red and blue can be deter-
mined because red areas revert to a yellow
or translucent amber state, and blue areas
to an opaque blue-grey

7 Helmet frontal length : mm the curvilinear distance, as measured with a
flexible tape, along the anterior margin of
the bony helmet from the point of juncture
with the skull to the apex

THE EVOLUTION OF GUINEA-FOWL

Name

Helmet rear length

Helmet central height

Helmet basal length
Helmet thickness

Cere structure length .
Cere structure thickness

Collar plumage

Nape filoplume length

Nape ffiloplume antero-
posterior coverage

Nape filoplume lateral
coverage

Nape filoplume density

Secondary remex outer web
vermiculation

Wing covert barring

Dorsal vermiculation .

Dorsal spot size .

Crest frontal length

Crest rear length

Units

mm

0-4

135

Description

the curvilinear distance along the posterior
margin of the helmet from the point of
juncture with the skull to the apex

the chord perpendicular to the line of
juncture with the skull to the highest point
along the margin of the helmet

the chord from the anterior to posterior
juncture points of the helmet with the skul]
the maximum lateral width of the helmet at
its base

the chord from the base of the cere to the
distal tip of the longest tuft or papilla

the maximum thickness of the thickest tuft
or papilla

0 = grey; 1 = blue/violet; 2 = black, barred
white with a blue wash; 3 = black, barred
white; 4 = black, barred white with faint
longitudinal streaking; 5 = black, barred
white with some spotting; 6 = spotted

the chord from the base to the tip of the
longest straightened nape filoplume

the percentage of the nape, at the mid-dorsal
line, from the occiput to the upper-most
collar feathers, covered by filoplumes

the percentage of the nape covered laterally
by nape filoplumes at the mid-point of the
anteroposterior coverage

a subjective estimate, based on a comparison
with reference specimens encompassing the
range of variation, of density of the nape
filoplumes on a scale of increasing density.
0 = filoplumes not present, 4 = covered by
a mat of filoplumes

0 = absent; 1 = extends +
white bands; 2 = extends
white bands; 3 = extends
white bands; 4 = extends = way along the
white bands; 5 = extends to full length of
white bands; 6 = outer edge of white bands
obliterated by vermiculation

O= not present; 1 — present but faint:
Di hesene

a subjective estimate, as with character no.
18, O = faint, graded subjectively to 4 =
dense

the maximum width, as measured with a
dissecting microscope fitted with an ocular
grid, of a particular spot on a randomly
selected feather from the mid-dorsal region
the chord from the base to the tip of the
longest straightened crest feather within
5 mm of the cere

the chord from the base to the tip of the
longest straightened crest feather within
5 mm of the most posterior extent of the crest

way along the
way along the
way along the

Cal enleo expo

136

34

35

36
37

38

39

40

4l
42

43
44

ANNALS OF THE SOUTH AFRICAN MUSEUM

Name
Crest central height

Crest basal length

Anterior crest curliness

Posterior crest curliness
Dorsal black collar

Ventral black collar
Occipital fold
Ear patch

Throat red

Orbital red

Dorsal spot number

Total spot barbs .
Total within spot barbs

Spot barb blueness

Chestnut blotch size

Chestnut blotch extent

Tarsal structure number

Tarsal structure length

White collar
Facial filoplumes

Units
mm

%o

count

count
mm

/o
0-1

Description

the chord perpendicular to the line of crest
juncture with the skull from the base of the
crest feathers to the highest point along the
margin of the unstraightened crest

the chord from the anterior to posterior
juncture points of the crest with the skull

a subjective estimate as with character no. 18.
1 = straight; 2 = moderately curly; 3 = very
curly

as with character no. 27

the percentage of black plumage extending
from the most anterior dorsal aspect of the
collar to the base of the tail

as with character no. 29 but the ventral
surface

a subjective estimate of the lateral extent of
the occiput covered by a fleshy fold

the maximum width of any patch of non-
blue/black skin anterior to the ear

a subjective estimate of the anteroposterior
extent of red pigmented skin between the
throat and the ventral base of the neck. As
with character no. 6, this character may be
assessed since the red colour reverts to a
yellow state after preservation.

a subjective assessment of the amount of
red pigmented skin around the eye

the maximum number of spots falling within
a circle of 1 cm radius superimposed over
the dorsal feather discussed in character
non22

the number of non-black barbs associated
with the spot measured in character no. 22
the number of barbs encompassed by the
spot measured in character no. 22

a subjective assessment of the amount of
blue in the spot. 0 = white; 1 = faint blue;
2 = medium blue; 3 = darkest blue

the maximum width of any chestnut blotch,
as measured with a dissecting microscope
fitted with an ocular grid, found on the
feather examined for character no. 22

the extent of the spotted area of the feather
examined for character no. 22 covered by
chestnut blotching

the number of spurs or bumps on the tarsus
the chord measured from the juncture line
of the longest tarsal bump or spur with the
tarso-metatarsus to the apex of the structure
the percentage of white plumage in the collar
a subjective estimate of the extent of the
head, other than the occiput, nape and
crown, covered by filoplumes

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

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Name of new genus or species is not to be included in the title: it should be included in the
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Biological Abstracts.

T. M. CROWE

THE EVOLUTION OF GUINEA-FOWL
(GALLIFORMES, PHASIANIDAE, NUMIDINAEB)
TAXONOMY, PHYLOGENY, SPECIATION AND

BIOGEOGRAPHY

VOLUME 76 PART 3 AUGUST 1978 : ISSN 0303-2515

| 507.68

~ ANNALS

OF THE SOUTH AFRICAN
MUSEUM

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

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

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.
Ann, Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. 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.

(continued inside back cover)

=i

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

Volume 76 Band
August 1978 Augustus
Parte "3 > Deel

SOUTHERN AFRICAN CUMACEA
PART 3

FAMILIES LAMPROPIDAE
AND CERATOCUMATIDAE

By

JENNIFER DAY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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SOUTHERN AFRICAN CUMACEA
PART 3
FAMILIES LAMPROPIDAE AND CERATOCUMATIDAE
By
JENNIFER DAY
Zoology Department, University of Cape Town

(With 16 figures and | table)
[MS. accepted 20 June 1978]

ABSTRACT

The Lampropidae in southern Africa are represented by eleven species in five genera.
Seven species are new: Platysympus camelus, P. depressus, P. compressus, P. phylloides, Para-
lamprops margidens, Hemilamprops glabrus and Hemilamprops sp. Hemilamprops pellucidus
and Bathylamprops calmani are redescribed and new figures are given. Adult males of Para-
lamprops (formerly Platytyphlops) peringueyi and Stenotyphlops spinulosus are described and
figured for the first time. The generic diagnosis of Paralamprops is altered to accommodate
information obtained from adult males of P. peringueyi, while the genus Platytyphlops is
invalidated.

Keys are given to the genera of the Lampropidae, the southern African members of the
family, the world species of Paralamprops, Platysympus, Bathylamprops, and Ceratocuma, and
to the species of Hemilamprops from the southern hemisphere.

The general distribution of lampropids is discussed and a more detailed account given
of the southern African representatives. It is concluded that lampropids are bipolar in distri-
bution, preferring deep and/or cold waters and avoiding the tropics. No member of the family
is found at depths of less than 188 m in these waters.

The only southern African member of the Ceratocumatidae, Ceratocuma horridum,
is redescribed and refigured and is considered to belong to a local subspecies, C. horridum
australe, which is polymorphic. The ceratocumatids are too poorly known to generalize
effectively about their distribution, but they all appear to be deep-water, essentially Atlantic
forms. None has been found at depths of less than 196 m or further from the Atlantic than
the south-east coast of South Africa and Kerguelen.

CONTENTS
PAGE
Introduction ; ; : . d ; : : = E38
Material and station data : , : : : : ‘ 5 dl sks
Methods . : 140
Key to the southern African Lampropidae and Ceratocumatidae 140
Family Lampropidae : : : ; : : ey At
Key to the genera of Lampropidae : , : ; : : 142
Stenotyphlops . ; : : : i ; a) 43
Paralamprops . : : 4 ; : : : : . 146
Platysympus 5 : : ; : : ' : 2 5 > ed
Bathylamprops . : : : y ‘ 5 : : oF AGS
Hemilamprops . : : : : ; : 2 £68
Distribution of the Lampropidae : : , ‘ seek he
Distribution of the southern African Lampropidae : ; Bern ies
Family Ceratocumatidae . : 5 : : : , 3 : 180
Ceratocuma : : ‘ A ‘ : a ng! Bok
Distribution of the Ceratocumatidae sige Uinae ‘ ‘ Le ST,
PNCKIOWICUBeIMeNtshine! oy Vs ie OY Sh a br ie ge GAS et NS
References) Po cee eet : ‘ : : : ao disks:
(37

Ann. S. Afr. Mus. 76 (3), 1978: 137-189, 16 figs, 1 table.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

This is the third in a series of papers on the Cumacea (Crustacea) of
southern Africa. The first two deal with the family Bodotriidae (subfamily
Vaunthompsoniinae (Day 1975), subfamily Bodotriinae (Day 1978)). The
reader is referred to the first of these for a discussion of the morphology and
terminology of the group as a whole.

Since the Lampropidae are essentially cold-loving forms, the only species
occurring in these waters are found at depths greater than 188 m where tempera-
tures are uniform and generally low (less than 12°C in these latitudes). Only
four species have previously been described from the southern African region:
Hemilamprops pellucidus Zimmer, 1908, Platytyphlops peringueyi Stebbing, 1912,
and Stenotyphlops spinulosus Stebbing, 1912, from southern Africa and Bathy-
lamprops natalensis Jones, 1969, from the South-western Indian Ocean. The
other species previously known from the African continent is Bathylamprops
calmani Zimmer, 1908, from deep waters off equatorial east Africa. A further
Six Species are described here, bringing the total number of named species for
southern Africa to ten. There is a further species (probably of Hemilamprops),
but all the individuals are too badly damaged to allow adequate descrip-
tion.

The Ceratocumatidae, a family known until recently (Jones 1969) from a
single species, appear to occur only in waters deeper than 196 m. One of the
two findings of the type species, Ceratocuma horridum Calman, 1904, was
recorded by Stebbing (1912) from Natal. Further individuals are now available
but they are morphologically variable and until more material is forthcoming
it will not be possible to say with certainty whether all individuals belong to
Calman’s species.

MATERIAL AND STATION DATA

The vast bulk of the material available to the author was provided by
the South African Museum (SAM). Part of it was obtained by the S.S. Pieter
Faure in 1898-1907 from deep waters round the coast of South Africa, and the
remainder was collected aboard the R.V. Meiring Naude in 1976-1977 during
a survey conducted by the Museum in deep waters off the east coast of South
Africa. A few of the samples come from the deepest stations of transects con-
ducted by the Zoology Department of the University of Cape Town (UCT)
aboard the University’s Research Vessel, the R.V. Thomas B. Davie, off Still Bay
and Lambert’s Bay.

Depth records for some of the Pieter Faure stations are approximate and
have been estimated from charts. Newly available information on depths off
the Cape Peninsula shows that the depth for SAM-—A10602 is about 800 m,
rather than 400 m as previously estimated.

Figure 1 shows the positions at which lampropids and ceratocumatids
were found. The code letters used are as follows:

SOUTHERN AFRICAN CUMACEA: PART 3 139

South African Museum SAM: Pieter Faure samples
SM: Meiring Naude samples
Zoology Department,
University of Cape Town LBT: transect at Lambert’s Bay, 200 km north
of Cape Town
SST: transect at Still Bay, 270 km east of
Cape Town
WCD: benthic survey off the western Cape
Province

* SAMA 10601

SAM-A 10602 a
SAM-A 594 * ,SAM-A.10606
SAMA 10607 *

Fig. 1. Coastline of the Cape Province showing positions of stations at which lampropids and
ceratocumatids were collected. Inset: coastline of Natal.

Dotted line indicates 200 m depth contour. See text for explanation of code letters.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

METHODS

Collections: a variety of gear was used for sampling: dredges in the Pieter
Faure and Meiring Naude programmes, and Van Veen grabs and Cape Town
dredges in the Thomas B. Davie programmes.

Length measurements were made from the anterior tip of the carapace to
the posterior tip of the telson. Exhalant siphons and uropods were excluded in
every case.

KEY TO THE SOUTHERN AFRICAN LAMPROPIDAE AND
CERATOCUMATIDAE

It should be noted that this key is designed to assist in the identification
even of damaged animals and those of varying ages in which the sex may be
difficult to determine. For this reason it should always be used in conjunction
with the generic keys for final identification.

1 Telson small, semicircular, lacking apical spines (may be deflected over anal valves)
(Fig. 15B); carapace sculptured into numerous rounded (Fig. 15A) or digitiform (Fig. 16A)
PEOCESSES” se ee se ale ha ere eco ene Ceratocuma horridum australe (Figs 15 & 16)

— Telson large, elongate, with at least three apical spines (Fig. 21); sculpturing variable but
NOt AS ADOVE 266 cece cee cess cd Kowa 's sue aiewia de alsd 4-0 elle] och eet 2

2 Carapace rounded, more or less circular in cross-section, totally devoid of marginal or
lateral carinae or longitudinal ridges on posterior half at least (Fig. 11A).............. 3

— Carapace dorsoventrally flattened anteriorly at least, marginal or lateral carinae or longi-
tudinal ridges present on most or all of carapace (Figs. 2A, 4A)............0.2000 cece 6

3 Carapace with irregular transverse rows of minute denticles; pseudorostrum almost
one-fiith totallength of carapace. 22... cee eee ee Bathylamprops calmani (Fig. 10)

— Carapace without transverse rows of denticles; pseudorostrum distinctly less than one-fifth
total ‘feneth of carapace (Fig. 11A) 22.22.00 0.005 3 oe cen os bee ee eee 4

4 Pseudorostrum truncate anteriorly with short, poorly defined ventrolateral carinae; telson
with five spines apically and none laterally.............. Hemilamprops glabrus (Fig. 13)

— Pseudorostrum pointed anteriorly, without lateral carinae (Fig. 11A); telson with three
apical and at least five pairs of lateral spmes .........::..-:.--22 5 eee 5

5 Anterolateral corner of carapace with several long, slender spines; telson less than half
lengthvof peduncle Of mropodsse. Sie Sncrd oso cere See Hemilamprops sp. (Fig. 14)

— Anterolateral corner of carapace smooth or minutely denticulate; telson at least two-thirds
length of peduncle of uropod........6.. 5400626 Hemilamprops pellucidus (Figs 11 & 12)

6 Carapace extraordinarily flat and leaf-like, almost as wide as abdomen is long..........

Platysympus phylloides (Fig. 6)

— Carapace rounded or flattened but not leaf-like, not nearly as wide as abdomen is long
(BIg. SA) ooo ee ee bec eile div ob ca che Sub ecen Oe 6 6 Rae ye Sr 7

7 Carapace and body with a number of longitudinal ridges formed by rows of small denticles;
carapace almost rectangular in dorsal outline; fifth pereiopod reduced to two segments

Stenotyphlops spinulosus (Figs 2 & 3)

— Carapace with a single sharp marginal carina; square or oval in dorsal outline; fifth
pereiopod consisting of at: least four seements. .. 6.004 5.5. 50.4.4. 5 see 8

8 Abdomen twice as long as cephalothorax; carapace almost square in dorsal view........

Paralamprops peringueyi (Fig. 4)

— Abdomen subequal in length to cephalothorax; carapace oval in dorsal view (Fig. 7B)... .9

9 Marginal carina of carapace strongly dentate; exopod present on pereiopod 2 of female

Paralamprops margidens (Fig. 5)

— Marginal carina of carapace not dentate (Fig. 6A, C); exopod absent from pereiopod 2 :

female. cece ei eae ec oe eae Reg be sl tlee bes De teh O8 ee eee ane

SOUTHERN AFRICAN CUMACEA: PART 3 141

10 Carapace smoothly oval or with a few small, rounded projections; not laterally compressed
dorsal to marginal carina; middorsal carina hardly evident.....................-e00.
Platysympus depressus (Fig. 7)

— Carapace smooth, laterally compressed dorsal to marginal carina (Fig. 8A); middorsal
EE TURD GIT (rier y foil B55 SOUS kant eae teat oe a ROR a mete AMEE eee Nga Ni cate alt eraee i

11 Dorsal edge of carapace smoothly arched; pseudorostrum pointed anteriorly in lateral view
Platysympus compressus (Fig. 8)

— Dorsal edge of carapace sinusoidal; pseudorostrum dorsoventrally truncate anteriorly in
Wee ARC WVER free ie iB es Sec nahh ala austere snalic idl wc Rypiors 4» e a">-aiaud le Platysympus camelus (Fig. 9)

Family Lampropidae Sars, 1878
Diagnosis
Antenna | with flagellum well developed. Antenna 2 of male with short
segments, of female with at least three segments. Mandibles of normal (boat)
shape. Palp of maxilla 1 absent or bearing one or two filaments. Exopods
"present on maxilliped 3 and pereiopod 1 in both sexes and on pereiopods 2 to
4 in male. Exopods present on pereiopod 2 and rudimentary on pereiopods 3
and 4, or absent from all three, in female. Pleopods in male 0 to 3 pairs, with
an outer process to the inner ramus. Telson moderate to large, well developed
post-anally, with three to five apical spines.

Type genus
Lamprops Sars, 1863.

Remarks

The presence of a well-developed telson with at least three apical spines
together with the well-developed first antenna is characteristic of the family.

Earlier workers tended to distinguish a greater number of families than
are now accepted. The families Chalarostylidae, Paralampropidae, Platy-
sympodidae, Pseudodiastylidae and Lampropidae of Stebbing (1913) are now
all included in the larger family Lampropidae.

The family is well defined and consists at present of ten genera, five of
which are represented in the present collection. One of these (Stenotyphlops
Stebbing, 1912) is known only from southern Africa.

A problematic feature of the taxonomy of the family is the fact that, being
deep-water forms for the most part, relatively few species are known and many
of these are represented by only one sex. Since the major distinction between
some genera (notably Lamprops, Mesolamprops and Hemilamprops) is the
number of pairs of pleopods present in adult males (zero, two and three pairs
respectively), females and juveniles cannot always be placed in a genus with
any certainty. This in turn makes it difficult to construct a useful key to the
genera. In the key below, an attempt has been made to use characters other than
those found only in adult males, but these are not very clear-cut. The geographic
distribution of the species of Lamprops, however, shows that it is essentially a
shallow-water genus confined to high latitudes of the Northern hemisphere. The
distribution of Hemilamprops, on the other hand, is much wider and its species

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

tend to occur in deeper waters. Thus there is little doubt that the species from
deep water in the Southern hemisphere for which only females are known—
H. lotusae Bacescu, 1969, H. ultimaespei Zimmer, 1921, H. glabra sp. nov. and
Hemilamprops sp.—are, indeed, members of the genus Hemilamprops. The single
specimen of Lamprops? comata Zimmer, 1907, from Tierra del Fuego is a
fragmentary female, so that its systematic position must remain indeterminate
for the present.

The presence in the collection of adult males of both Stenotyphlops and
Platytyphlops with three pairs of pleopods throws some light on the relation-
ship between these genera and closely allied ones. Details of the findings are
presented in the discussion of Paralamprops on page 147.

The genera of the family as a whole are morphologically unremarkable for
the most part, but for the fact that in several cases the carapace is very strongly
flattened dorsoventrally, with a single, sharp lateral carina encircling the entire
carapace apart from the posterior edge (and here called a marginal carina to
distinguish it from the more usual lateral carinae found widely in several
families). The reason for this adaptation is not clear, but since all the species
exhibiting this character are deep-water forms, it may have evolved as a means
of increasing the surface area to prevent sinking into the oozy mud of the
seafloor.

A singular genus, described by Bacescu (1972), is Archaeocuma. It is mono-
typic and known only from the Peruvian Trench. It is distinguished by the
presence in both sexes of a single pair of pleopods. In all other characters it is
typically lampropid.

KEY TO THE GENERA OF LAMPROPIDAE

All males have three pairs of pleopods unless otherwise stated.

1 Antenna 1 with third segment no longer than second and accessory flagellum minute;
telson and peduncle of uropods more than four times length of telsonic somite ........
Pseudodiastylis Calman, 1905

— Antenna | with third segment no longer than second and accessory flagellum well developed
or with third segment elongate and accessory flagellum very small; telson and peduncle of
uropod no more than four times leneth of telsonic somite. ... 2.2.0.2 eee p2

2 Telson small, subequal in length to telsonic somite and a third length of peduncle of uropod
(etiaa learns OW) eclec bose eaten otro eatin ae eieeae cao cupaie une Chalarostylis Norman, 1879

— Telson distinctly longer than telsonic and at least half length of peduncle of uropod...... 3
3 Pereiopod 5 reduced to a minute, 2-segmented projection; a single filament on palp of
PAA KATA s,s ors ease tae Pavsiel Skee ait Me onaeekas<ancreey A) cal Sim ae ea auc Stenotyphlops Stebbing, 1912
Pereiopod 5 normal, or if reduced, at least 4-segmented; palp of maxilla 1 absent or with
two filaments: (25.2. Pew eastern Be Dae teh os WO 4 ele a ote wie 0 pr OR 4
4 Basis of pereiopod 4 longer than entire length of pereiopod 5...............0 cece ne eeee 5
— Basis of periopod 4 subequal to, or shorter than, entire length of perelopod 5............ 6
5 Basis of maxilliped 3 much shorter than remaining segments together; male and female
Oth! Withone pair Ol MlEODOdS. 04.0 ack eke eae eee Archaeocuma Bacescu, 1972

— Basis of maxilliped 3 not shorter than remaining segments together; male with three pairs
On pleopods and! female with NOUS... 5 acm .2.os 40 ee ee Paralamprops Sars, 1887

6 Maxilla 1 lacking palp; exopods absent from pereiopods 2-4 in female; carapace with
Sirons marginal Carinae iin ees okton, ele eee attuned he Platysympus Stebbing, 1912

SOUTHERN AFRICAN CUMACEA: PART 3 143

— Palp of maxilla 1 with two filaments; exopods normal on pereiopod 2 and rudimentary on
pereiopods 3 and 4 in female; number of pairs of pleopods in male 0, 2 or 3; carapace
SNM LSTA OI CALI Devas cin cise cei < cel aia es: specie oe waters gic oie ee ware le wie Sn ReRSee ORL TepeR dhe ois s a

7 Pseudorostrum at least a fifth of total length of carapace; third segment of antenna 1 longer
and considerably more slender than second..............%. Bathylamprops Zimmer, 1908

— Pseudorostrum distinctly less than a fifth total length of carapace; third segment of antenna |
Rename longer or much more slender than second ........0....0 6.00 es es sade cew ees 8

8 Male without pleopods; antennal notch small but distinct; eye present; basis of pereiopod 1
fperexinately equal in length to rest of limb.................-..25: Lamprops Sars, 1863

— Male with two pairs of pleopods; antennal notch present or absent; eye present or absent;
basis/or pereiopod | shorter than rest of limb................ Mesolamprops Given, 1964

— Male with three pairs of pleopods; antennal notch usually absent; eye present or absent;
basis of pereiopod 1 distinctly shorter than rest of limb........ Hemilamprops Sars, 1883

Stenotyphlops Stebbing, 1912
Generic diagnosis
Carapace not strongly flattened. Eye absent. Both flagella of antenna |
well developed. Palp of maxilla 1 with one filament. Exopods of pereiopods 3
and 4 of female rudimentary. Pereiopod 5 reduced to a minute, 2-segmented
projection. Male with three pairs of pleopods.

Type species
S. spinulosus Stebbing, 1912 (by monotypy).

Remarks

The genus is monotypic and the single species is known only from southern
Africa. The combined presence of a single filament on the palp of maxilla 1
and the greatly reduced fifth pereiopod is diagnostic. The adult male described
below is the first known for the genus. The presence of three pairs of pleopods
confirms Stenotyphlops as typically lampropid, while the extreme reduction of
the fifth pereiopod and the nature of maxilla 1 clearly separate it from the other
genera in the family. In general morphology it is otherwise very close to
Paralamprops.

Distribution of Stenotyphlops
Deep water off southern Africa.

Stenotyphlops spinulosus Stebbing, 1912
Figs 2-3
S. spinulosus Stebbing, 1912: 162-163, pl. 60.

Records

SAM-A10602 (PF 17440) 34°25’S 17°45’E 800m 1 ¢: 10,5 mm; 3 92: 11,2 mm and
damaged; 4 juvs

SAM-A10607 (PF 16982) 34°40’S 17°50’E 1200m 1 adult ¢: 13,8 mm; 1 damaged

adult 2
SM 60 27°09’S 32°58’E 800 m_ 1 damaged 3
SM 103 2832'S 32:34 5 680 m_ 1 damaged 3; 1 ovig. 2: 12,5 mm
SM 123 30°33’S 30°48’E 690 m 2 22: 7,0 mm and damaged

SM 129 30538 30328 = 850an 1.02 956 mm 19: 99am

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

Previous records

Holotype only.

Holotype

Adult female, deposited by Stebbing in the British Museum (Natural
History). Type locality: approximately 370-550 m, off the Cape Peninsula
G4°25'S 17°S0'E).

Description

Ovigerous female, length 12,5 mm (SM 103). Integument slightly trans-
lucent, armed with very small denticles. Carapace (Fig. 2A) slightly flattened
anteriorly and inflated posteriorly with three longitudinal rows of denticles on
either side and two on the anterior sinus. Middorsal carina evident anteriorly

Fig. 2. Stenotyphlops spinulosus

Ovigerous female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3.
E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Pereiopod 5. I. Telson and peduncle of
uropod.

Young female. J. Uropod.

Scale line = 4 mm for A-B; 2 mm for C-G; 1 mm for I-J; 0,5 mm for H.

SOUTHERN AFRICAN CUMACEA: PART 3 145

(Fig. 2B) behind eyelobe, minutely denticulate. Eyelobe eyeless. Carapace
inflated posterodorsally on either side of middorsal depression. First three
thoracic somites flanged laterally. Abdominal somites cylindrical. Cephalo-
thorax and abdomen subequal in length.

Antenna | (Fig. 2C) of moderate size; first segment subequal in length to
next two together. Both flagella well developed.

Palp of maxilla 1 with a single filament.

Maxilliped 3 (Fig. 2D) fairly stout, basis subequal in length to rest of
limb. Carpus long and parallel-sided, propodus and dactyl slender.

Pereiopod | (Fig. 2E) not elongate, basis subequal in length to next four
segments together. Ischium small, merus and carpus elongate. Propodus and
dactyl cylindrical.

Pereiopod 2 (Fig. 2F) elongate. Basis subcylindrical, carpus long and stout,
armed with a row of spines on inner edge. Exopod small and slender.

Pereiopods 3 (Fig. 2G) and 4 similar. Basis subequal in length to rest of
limb, merus longest of remaining segments. Dactyl minute. Exopod very small,
2-segmented.

Pereiopod 5 (Fig. 2H) reduced to a minute, 2-segmented stump.

Telson (Fig. 21) distinctly wider proximally than distally, about three-
quarters of length of peduncle or uropod, distally armed with five to six pairs
of spines laterally and three single spines terminally. Rami of uropod damaged
in adult female. Peduncle of uropod of young female (Fig. 2J) subequal in
length to endopod. First segment of exopod much shorter than second. First
segment of endopod nearly twice length of second and third together. Endopod
longer than exopod by one segment.

Adult male, length 13,8 mm (SAM-—A10607). As female, except as follows:
carapace (Fig. 3A) less flattened anteriorly and less inflated posteriorly. Ridges
on carapace more distinct, not always denticulate. Lateral flanges of thoracic
somites scalloped (Fig. 3B).

First segment of flagellum of antenna | (Fig. 3C) bearing numerous short
aesthetascs. Antenna 2 reaching about half way along length of body. Palp of
maxilla | (Fig. 3D) illustrated. Segments distal to basis of maxilliped 3 and
pereiopod 1 missing. Exopod of pereiopod 2 larger. Pereiopod 3 (Fig. 3E)
stouter, dactyl minute and apparently continuous with small terminal spine.
Pereiopod 5 (Fig. 3G) longer, but still 2-segmented. Three pairs of normal
pleopods present.

Telson (Fig. 3F) shorter, about half length of peduncle of uropod. Armature
of peduncle and proximal part of endopod of uropod more extensive. Distal
portions missing.

Remarks
No adult males have previously been described. The differences between the

female described here and Stebbing’s holotype are slight, and the differences
between adult male and adult female are within the limits expected between

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Stenotyphlops spinulosus

Adult male. A. Lateral view. B. Dorsal view of cephalothorax. C. Antenna 1. D. Maxilla 1.
E. Pereiopod 3. F. Uropod and telson. G. Pereiopod 5.

Scale line = 4 mm for A-B; 2 mm for C, E-F; 1 mm for D, G.

sexes. Stebbing figures his female with a slightly narrower carapace and a larger
first segment of antenna 1. The flagellum of his female has four segments and
the present one five. The greatest difference is in the telson: Stebbing figures it as
being not very much longer than the telsonic somite and little more than half
the length of the peduncle of the uropod. The telsons of both male and female
figured here are twice the length of the telsonic somite and in the female is
nearly as long as the peduncle of the uropod. In the male the peduncle is much
longer, nearly twice as long as the telson. However these differences may
simply be due to individual variation since the lengths of the peduncle and
telson vary somewhat among the individuals available at present. Apart from
this, they agree well with Stebbing’s figures and there is little doubt that they
belong to the same species.

Distribution
From Cape Point (about 500 to 1 200 m) to Natal (680 to 850 m).

Paralamprops Sars, 1887
Generic diagnosis

Carapace slightly or strongly depressed dorsoventrally with a marginal
carina or at least one pair of lateral carinae. Antenna | with both flagella well
developed. Palp of maxilla 1 absent or with two filaments. Exopods of
pereiopods 3 and 4 of female rudimentary or absent. Pereiopod 5 small to

SOUTHERN AFRICAN CUMACEA: PART 3 147

rudimentary, no longer than basis of pereiopod 4. Male with three pairs of
pleopods. Telson well developed.

Type species
Paralamprops serratocostata Sars, 1887.

Remarks

The genus has consisted up to now of seven species: P. serratocostata Sars,
1887, P. orbicularis Calman, 1904, P. aspera Zimmer, 1907, P. semiornata Fage,
1928, P. grimaldi Fage, 1928, P. arafurensis Jones, 1969, and P. rossi Jones,
1971. The genus Platytyphlops was established by Stebbing (1912) on the basis
of several specimens including a fragmentary ovigerous female and at least one
young male. He decided that the male was probably mature and therefore
characterized the genus as having no pleopods in the male and a greatly reduced
fifth pereiopod in both sexes. One fully adult and several subadult males of
the same species are now available, however, and possess three pairs of normal
pleopods. They should thus be placed in Paralamprops. It also turns out that
the reduction of the fifth perelopod is less evident in the adult male than in
immature males or adult females in P. peringueyi and that, in fact, the limb is
reduced to some extent in all species of Paralamprops. But the degree of reduc-
tion varies considerably, reaching its limit in P. peringueyi. Thus the generic
diagnosis of Paralamprops has been slightly altered accordingly.

The genus is morphologically rather variable. The first maxilla in P. serra-
tocosta and P. margidens sp. nov. lacks a palp while in P. orbicularis, P. semi-
ornata, P. grimaldi and P. peringueyi there is a normal palp with two filaments.
In these last four species and in P. rossi the carapace is very strongly flattened
with a single, sharp marginal carina forming a wide, flat, flange encircling the
entire carapace apart from the extreme posterior edge. The first maxilla is not
described for P. rossi, P. aspera or P. arafurensis, all of which are known from
single specimens which would have been badly damaged by dissection of the
anterior mouthparts. In P. serratocostata, which lacks a maxillary palp, the
carapace is not strongly flattened dorsoventrally but, in common with P. ara-
furensis and P. aspera, does possess a number of longitudinal ridges. P. margi-
dens also lacks a maxillary palp, but the carapace is somewhat flattened and
bears a single dentate marginal carina, thus being intermediate between the two
types described above.

Calman (1912) was of the opinion that the absence of a palp on maxilla 1
was ‘so important and unexpected that it might justify the creation of a new
genus’. The present author agrees that this character is of considerable signifi-
cance and suggests that in the future Paralamprops may well be split into two
genera on the combined characters of the first maxilla and the carapace. How-
ever, as Jones (pers. comm.) has pointed out, the practical difficulty of examining
the maxilla in rare species and the lack of information on the adults of many
of the species under discussion, precludes the splitting of the genus at present.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stenotyphlops, distinguished by a single filament on the palp of maxilla 1,
a greatly reduced fifth pereiopod and several longitudinal ridges on the cara-
pace, is very close to Paralamprops, as is Archaeocuma, which is distinguished by
a single pair of pleopods. The most closely allied genus of all is Hemilamprops.
In fact, it is difficult to find a really satisfactory set of characters to distinguish
Hemilamprops from some members of Paralamprops, other than the roundness
of the carapace and shortness of the abdomen in Hemilamprops, and the fact
that there is little or no reduction of the fifth pereiopod in this genus. However,
these characters are easily distinguishable and seem to be uniform. In order to
avoid making Hemilamprops unwieldy and even more diverse than it is at
present, the two genera must be kept apart for convenience’ sake.

Hemilamprops mawsoni Hale, 1937, which was not a satisfactory member of
that genus, can, however, now be placed in Paralamprops, where it is very
similar to P. rossi. This brings the total number of species of Paralamprops to ten.
Distribution of Paralamprops

Known at depths from 232 to 3 789 m in the Atlantic, Antarctic and
East Indies.

KEY TO THE SPECIES OF PARALAMPROPS

1 Carapace with a single, shatp marsinal cara. .2..........). ose eee D
— Carapace with at least three pairs of lateral and/or dorsolateral carinae .............. 8
2 Catrapace no more elevated posteriorly than anteriorly... ... 4.0 3
— Carapace more elevated posteriorly tham anteriorly. /.).:...... 2 eee 4
3 Telson little longer than telsonic somite; second and third segments of antenna 1 subequal

in length; ischium of pereiopod 1 about as wide as long Br
P. orbicularis Calman, 1904—N. Atlantic
— Telson more than twice length of telsonic somite; third segment of antenna 1 half length
of second; ischium of pereiopod 1 much wider than long
P. semiornata Fage, 1928—W. Portugal

4 Telson nearly equal in length to last two somites together. |...) ee eee 5
—' Telson about half length of last two somites together... ...2.. .. a2 snes ih
>) Marcinalicanna strongly dentate. 2.2.5 nae oe ee ee P. margidens sp. nov.
= Marginal carina smooth ....... 0.666 als ec ts we oe ooo wel ce eiese oust SRS ee nen ee ee 6
6 Minute exopods on pereiopods 3 and 4 of female; carapace not transversely ridged in mid-

dorsal gutter; pseudorostrum pointed anteriorly in dorsal view
P. mawsoni (Hale, 1937)— Antarctic
— No exopods on pereiopods 3 and 4 of female; carapace lightly ridged transversely in mid-
dorsal gutter; pseudorostrum rounded anteriorly in dorsal view
P. rossi Jones, 1971—Ross Sea
7 Fifth pereiopod 5-segmented (female) or 6-segmented (male), much less than half length
of pereiopod 4; basis of pereioped 2 shorter than’ rest of limb. 2 ee eae eee
P. peringueyi (Stebbing, 1912)—South Africa
— Fifth pereiopod 7-segmented, as long as basis of pereiopod 4; basis of pereiopod 2 longer
than srest soft limba): Data een eeees J eels Lee ee P. grimaldi Fage, 1928— Azores
8 Middorsal carina not serrate; telson little narrower posteriorly than anteriorly with five
SPINES Trans VenselyaGkOssta exe ene ancien acne P. arafurensis Jones, 1969—East Indies
— Middorsal carina serrate; telson distinctly narrower posteriorly than anteriorly with three
apical’spines’ <i .8 6e ee eee bE held s wis Saeed Oe Cee Si Ok eee 9
9 Telson no more than twice length, of telsonic somite..-..5-.2 52s eeeeeeee
P. serratocostata Sars, 1887—Kerguelen
Telson three times length of telsonic somite.......... P. aspera Zimmer, 1907—Antarctic

SOUTHERN AFRICAN CUMACEA: PART 3 149

Paralamprops peringueyi (Stebbing, 1912)
Fig. 4
Platytyphlops peringueyi Stebbing, 1912: 159-161, pls 58-59.

Records

SAM-A596 (PF 17585) 34°48’S 18°03’E 369-554m 3 29:9,3 mm, 9,9 mm, damaged
(paratypes)

SAM-A10602 (PF 17440) 34°25’S 17°45’E 800 m__1 adult gd: 14,7 mm; 1 damaged

subadult 3; 3 dd: 7,7-10,6 mm;
3 292: 8,0-16,0 mm; 2 damaged
juvs

SAM-A10606 (PF 16769) 34°37’S 17°50’E 1394m 1 ovig. 2: 14,7 mm

Previous records

Type locality only.

Holotype

Not designated: syntypes include ovigerous female and young males from
two samples (PF 17585 and PF 17643), deposited in the British Museum
(Natural History). Type locality: between 370 and 550 m, off Cape Point
(34°48’S 18°03’E).

Description

Adult male, length 14,7 mm (SAM-A10602). Integument with minute
triangular denticles. Sides of carapace strongly depressed (Fig. 4A), marginal
carina very evident; median part of carapace compressed laterally, slightly
more elevated posteriorly than anteriorly. Middorsal line defined anteriorly by
extremely well-developed carina and posteriorly by a narrow gutter flanked by
a pair of flattened dorsal elevations, denticulate on their anterior edges and
bent outwards slightly (Fig. 4B). Eyelobe small, eyeless, flanked by flattened,
upturned, lateral extensions of pseudorostral lobes. Carapace slightly longer
than wide.

First pedigerous somite exposed dorsally only; second to fourth slightly
flanged laterally, fifth cylindrical. Cephalothorax more than three-quarters
length of cylindrical abdomen.

Antenna | (Fig. 4C) fairly large; first segment longer than next two together.
Flagellum elongate, 4-segmented. Accessory flagellum 5-segmented with
numerous short aesthetascs on basal segment.

Palp of maxilla 1 with two filaments.

Maxilliped 3 (Fig. 4D) short, leg-like. Merus slightly expanded, carpus
and propodus cylindrical, subequal in length.

Basis of pereiopod 1 (Fig. 4E) stout, ischium slightly expanded. Carpus
slightly longer than ischium and merus together. Part of propodus and dactyl
missing. Exopod large and stout, basal segment almost circular.

Basis of pereiopod 2 (Fig. 4F) fairly short, carpus longer than ischium and

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Paralamprops peringueyi
Adult male. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Maxilliped 3.
E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Pereiopod 5.
Adult female. I. Lateral view. J. Dorsal view of carapace. K. Pereiopod 3. L. Pereiopod 5.
M. Uropod and telson.

Scale line = 4 mm for A-B, I-J; 2 mm for C-H, K, M; 0,5 mm for L.

merus together, armed with a row of sharp spines. Dactyl incomplete. Exopod
large.

Pereiopods 3 (Fig. 4G) and 4 similar, basis very large in comparison with
rest of limb, of which merus is longest.

Pereiopod 5 (Fig. 4H) very small, 6-segmented. All segments distal to
basis subequal in length. Entire limb less than half length of basis of pereiopod 4.

SOUTHERN AFRICAN CUMACEA: PART 3 151

Three pairs of pleopods present.

Uropods and telson as in female (Fig. 4M) except that telson has three pairs
of lateral spines, not two.

Adult female, length 16,0 mm (SAM-A10602). As male, except as follows:
posterodorsal elevations of carapace narrower and curling over laterally
(Fig. 41). Middorsal carina much less well developed, minutely denticulate.
Carapace almost as wide as long (Fig. 4J). First pedigerous somite visible
laterally as well as dorsally.

Antenna | with third segment slightly longer, first segment of flagellum
without aesthetascs. Second antenna 4-segmented. Exopod of pereiopod 2
smaller. Basis of pereiopods 3 (Fig. 4K) and 4 relatively smaller, distal segments
larger and stouter; exopods minute. Pereiopods 5 (Fig. 4L) minute, 5-segmented,
with a stout terminal spine.

Telsonic somite (Fig. 4M) nearly twice as long as broad, two-thirds length
of telson. Telson with two pairs of lateral spines and five terminally. Peduncle
of uropod slightly longer than telson with numerous small spines on inner edge.
Rami of uropods incomplete.

Remarks

The adult male has not previously been described but corresponds well
with the female in most respects. The female differs slightly from that described
by Stebbing (1912), mainly because his was considerably smaller and less mature.
In particular the carapace is square, not rounded, in dorsal view and the carpus
and propodus of maxilliped 3 are smaller in the present specimens. The exopods
of the thoracic limbs of Stebbing’s male are smaller, again because it is immature.
Nevertheless these characters are of little specific significance and there is no
doubt that all individuals belong to the same species. It should be noted that
there is some individual variation in the degree to which the median part of the
carapace is elevated, particularly in some of the younger individuals.

P. peringueyi is characterized by the very great reduction of the fifth
pereiopod, particularly in the female. It is most similar to P. grimaldi, from
which it may be distinguished by this character.

Distribution

Only known off the Cape Peninsula from about 369 to 1 394 m.

Paralamprops margidens sp. nov.

Fig. 5
Records
SAM-A595 (PF 15785) 34°39'S 18°10’E 500m _ 1 9:6,1 mm

SAM-A10602 (PF 17440) 34°25'S 17°45’'E 800m _ 2 99: 6,1 mm (holotype + 1); 3 dd:
5,8-6,1 mm

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

Young female, in the South African Museum, SAM-A15721, collected by
the S.S. Pieter Faure in about 1900. Type locality: 800 m, off the Cape Peninsula
(34°25'S 17°45’E).

Description

Young female, holotype, length 6,1 mm. Integument lightly calcified,
reticulate on body and very slightly denticulate on some limbs. Carapace
(Fig. 5A) slightly wider than deep, nearly twice as long as deep, somewhat
depressed immediately behind eyelobe. Marginal carina strongly dentate. Mid-
dorsal carina shallow, serrate anteriorly. Branchial regions somewhat inflated.
No anterolateral angle (Fig. 5B). Pseudorostral lobes short in dorsal view
(Fig. 5C). Eyelobe triangular, eyeless.

First three pedigerous somites denticulate laterally. Abdominal somites
subcylindrical. Cephalothorax subequal in length to abdomen.

Antenna | as in male (Fig. 5N), but third segment slightly longer and
flagellum 4-segmented. Accessory flagellum damaged in all females.

Antenna 2 (Fig. 5D) of moderate size, 4-segmented, with the last two
segments relatively long.

Maxilla | (Fig. 5E) with no sign of palp.

Maxilliped 3 (Fig. 5F) leg-like, basis subequal in length to rest of limb.
Ischium small, merus slightly expanded on outer edge with two large terminal
spines. Carpus inserting subterminally on merus, subequal in length to last two
segments together. Exopod well developed.

Pereiopod | (Fig. 5G) very long, basis half length of rest of limb. Merus
and carpus subequal in length, propodus almost as long as merus and carpus
together, dactyl slightly shorter with several terminal spines. Exopod well
developed.

Pereiopod 2 (Fig. 5H) long and slender. Basis subequal in length to next
four segments together. Carpus stout with six large spines on lower edge. Last
two segments slender. Only basal segment of exopod present.

Pereiopods 3 (Fig. 51) and 4 similar, basis of pereiopod 4 relatively shorter.
Basis of pereiopod 3 very slender, more than twice length of rest of limb.
Exopod very small, 2-segmented.

Pereiopod 5 (Fig. 5J) hardly more than half length of basis of pereiopod 4.

Telsonic somite (Fig. 5K) wider than long, little more than a third length
of telson. Telson tapering evenly from base, armed with five pairs of short
spines laterally and three longer ones terminally. Peduncle of uropod slightly
longer than telson. Exopod reaching end of second segment of endopod.
Endopod 3-segmented, first armed with fine setae on inner edge.

Subadult male, paratype, length 6,1 mm. As female, except as follows:
carapace (Fig. 5L) somewhat shallower, denticles slightly larger. No depression
behind eyelobe. Middorsal serrations (Fig. 5M) larger and extending further
back. Third and fourth pedigerous somites denticulate dorsally.

SOUTHERN AFRICAN CUMACEA: PART 3 153

Fig. 5. Paralamprops margidens sp. nov.

Adult female, holotype. A. Lateral view. B. Detail of anterior end of carapace. C. Dorsal view
of carapace. D. Antenna 2. E. Maxilla 1. F. Maxilliped 3. G. Pereiopod 1. H. Pereiopod 2.
I. Pereiopod 3. J. Pereiopod 5. K. Uropod and telson.

Subadult male, paratype. L. Lateral view. M. Detail of anterior end of carapate. N. Antenna 1.
O. Basis of pereiopod 1. P. Pereiopod 3.

Scale line = 2 mm for A, C, L; 1 mm for B, F--K., M, O-P; 0,5 mm for D-E, N.

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Each segment of antenna | (Fig. 5N) slightly longer than succeeding one;
flagellum 5-segmented, accessory flagellum 3-segmented. Basis of pereiopod 1
(Fig. 5O) strongly dentate. Distal segments of exopods missing from pereiopods
1 and 2. Bases of pereiopods 3 (Fig. 5P) and 4 shorter and stouter, exopods
3-segmented. Basis of pereiopod 5 slightly longer. Armature of uropods reduced
(but possibly lost due to age).

Remarks

Lacking a palp on maxilla 1, P. margidens falls within the serratocostata—
aspera—arafurensis group of Paralamprops. It is easily distinguished from these
and from the South African P. peringueyi by the denticulate marginal carina.

Distribution
Known only from about 500 to 800 m off the Cape Peninsula.

Platysympus Stebbing, 1912
Platyaspis Sars, 1869: 158 (preoccupied name).

Generic diagnosis

Carapace strongly flattened dorsoventrally with strong marginal carina.
Both flagella of antenna | well developed. Maxilla 1 without palp. Pereiopods 2
to 4 of female without exopods. Male with three pairs of pleopods. Telson well
developed.

Type species
Platyaspis typicus Sars, 1869.

Remarks

The genus is well-defined and easily recognizable due to the very charac-
teristic flattened carapace with a strong marginal carina. The absence of exopods
on pereiopods 2 to 4 in the female is unique in the family although in other
genera they may occasienally be absent from pereiopods 3 and 4. The presence
of four new species in southern African waters brings the total number for the
genus to seven.

Distribution of Platysympus
Europe from 226 to 1 100 m; North Atlantic from 219 to 957 m; Antarctic
at 385 m; South Africa from 188 to 1 200 m.

KEY TO THE SPECIES OF PLATYSYMPUS

1 Carapace with three longitudinal ridges dorsal to marginal carina.........:.....5.-++-+-
P. tricarinatus Hansen, 1920—N. Atlantic
Carapace with middorsal and marginal carinae forming only major longitudinal ridges... .2
Pereiopod 5 half length of pereiopod 4; female with rudimentary exopods on pereiopod 2
P. brachyurus* Zimmer, 1907— Antarctic
— Pereiopod 5 more than half length of pereiopod 4; pereiopod 2 of female without exopod. .3

N

SOUTHERN AFRICAN CUMACEA: PART 3 155

3 Carapace remarkably flattened and leaf-like: almost circular in dorsal view and wider than

© SVL SIRRVETT, TSMLGUD Se Gai Sheth OR anny er ee P. phylloides, sp. nov.
— Carapace flattened but not leaf-like: longer than wide in dorsal view and considerably
narrower than abdomen is long............. 4
4 Dorsal outline of carapace undulating sinusoidally, forming two evenly-spaced elevations;
bases of pereiopods 1 and 2 with flattened, scale-like edges........... P. camelus sp. nov.
— Dorsal outline of carapace smooth or with several low, unevenly-distributed tumidities;
bases of pereiopods 1 and 2 without flattened, scale-like edges...............0 00 e cues 5

5 Dorsal third of carapace strongly compressed laterally, forming a very deep middorsal
carina; basis of pereiopod 5 less than half length of rest of limb... .P. compressus sp. nov.

— Dorsal third of carapace not strongly compressed laterally, middorsal carina negligible or
incorporating much less than a third of its depth; basis of pereiopod 5 more than half
“2 £121 SL OSE OUNTAOL soeh ie ee aa ae me RHO SL Oh Se Alle eee er ems poet 6

6 Middorsal carina distinct over whole length of carapace; carapace of female smooth; merus,
carpus and propodus of pereiopod | wide and flattened... .P. typicus (Sars, 1869)— Europe

— Middorsal carina of carapace only evident posteriorly; carapace of female with several
tumidities and depressions; merus, carpus and propodus of pereiopod 1 not wide or flattened
P. depressus sp. nov.

*P. brachyurus is known only from one incomplete female individual. Some characteristics
suggest that when further material is available the species will be found to fit better in
Paralamprops.

Platysympus phylloides sp. nov.

Fig. 6
Records
SAM-A10607 (PF 16982) 34°40’S 17°50’E 1200m _ 1 adult 2: 7,2 mm (holotype);
1 damaged 2
SM 129 30°54’S 30°51’E 850m _ 2 ovig. 22: 7,4 and 7,7 mm (both
damaged)
Holotype

Adult female, in the South African Museum, SAM-—A15682, collected by
the Pieter Faure in about 1900. Type locality: approximately 1 200 m, off the
Cape Peninsula (34°40’S 17°50’E).

Description

Adult female, holotype, length 7,2 mm. Integument very delicate and trans-
lucent. Carapace (Fig. 6A) remarkably flat and leaf-like, almost circular in
dorsal view (Fig. 6B) with middorsal carina faintly evident anteriorly. Eyelobe
small, rounded and eyeless. Carapace flattened and paper-thin at edges forming
an almost transparent wide flange extending round entire edge except where
attached to abdomen. Posterolaterally, extensions of this carina form flaps
overlapping third pedigerous somite on each side and leaving a gap on either
side of second somite. All cephalothoracic appendages except fifth peretopod
entirely covered by carapace (Fig. 6C); bases of maxilliped 3 and pereiopod 1
pointing forward and attached underneath carapace by a thin membrane,
being thus quite immobilized.

All five pedigerous somites free, last four wider than deep. Carapace
wider than abdomen is long. Cephalothorax about half as long again as cylin-
drical abdomen.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Platysympus phylloides sp. nov.
Adult female, holotype. A. Lateral view. B. Dorsal view. C. Ventral view of cephalothorax.
D. Antenna 1. E. Antenna 2. F. Maxilla 1. G. Maxilliped 2. H. Maxilliped 3. I. Pereiopod 1.
J. Pereiopod 2. K. Pereiopod 3.

Scale line = 4 mm for C; 2 mm for A-B; | mm for G-K; O,5 mm for D-F.

Antenna | (Fig. 6D) small, first segment subequal in length to next two
together. Both flagella short but well developed, main flagellum 3- and accessory
flagellum 2-segmented.

Antenna 2 (Fig. 6E) reasonably large, 3-segmented.

Maxilla 1 (Fig. 6F) without palp.

Maxilliped 2 (Fig. 6G—from ovigerous female, not holotype) with distal
segments short. Oostegal setae long and numerous.

Basis of maxilliped 3 (Fig. 6H) very flexible but immobile, being attached

SOUTHERN AFRICAN CUMACEA: PART 3 1S)

to the ventral surface of the carapace. Exopod normal but setae very small
and poorly setulose. Ischium small; merus slightly expanded; carpus large
and stout; propodus and dactyl slender.

Basis and exopod of pereiopod 1 (Fig. 61) as in maxilliped 3. Ischium
small. Carpus elongate, subequal in length to propodus and dactyl together.

Pereiopod 2 (Fig. 6J) lacking exopod. Basis slender, subequal in length
to rest of limb. Carpus long and well armed.

Pereiopods 3 (Fig. 6K) to 5 all similar, lacking exopods. Limbs slender,
basis subequal in length to rest of limb; merus, carpus and propodus subequal
in length.

Telson (Fig. 6B) twice length of telsonic somite, armed with only three
small apical spines. Peduncle of uropod subequal in length to last somite and
telson together, unarmed. Exopod unarmed, about two-thirds length of endo-
pod. Each segment of endopod armed with a single small spine distally on
inner edge.

Males have not been found.

Remarks

In the absence of male individuals it is not possible to state with certainty
that this species belongs to Platysympus. It is certainly quite distinctive in the
nature of the carapace, but apart from this, the appendages are very similar
to other members of the genus. It may be that this is merely the ultimate con-
dition in a genus in which the carapace is always flattened, and that the peculiari-
ties of its morphology are necessary to overcome problems associated with a
greatly flattened carapace. For example, the exopods, on those limbs which
have them, are reduced and it is difficult to see that they would be of any use
if they were present. Pleopods would seem to be unimportant as a means of
locomotion in such a flattened animal and it may prove that none are present
even in the adult male. If this should be so, then the species will have to be placed
in a separate genus.

The peculiar attachment of the third maxilliped and first pereiopod to
the floor of the carapace is presumably also an adaptation to the overhanging
carapace, as is the presence of the inhalent aperture on either side of the second
pedigerous somite.

Distribution
Cape Point at 1 200 m and Natal at 850 m.

Platysympus depressus sp. nov.
Fig. 7
Records
SAM-—A10602 (PF 17440) 34°25’S 17°45’E 800m 1 adult ¢: 5,8 mm (holotype);
2 subadult gd: 5,4 mm; 2 young

36d: 4,5 mm, 4,8 mm; 3 ovig. 29:
5,8-6,4 mm; 6 2°: 5,1-6,4 mm

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM 86 2H -59'S'32-40'E ( 550m 1 subadult 3: 5,4 mm

SM 103 28°31’S 32°34’E_ =—- 680 m 1 adult 3: 7,0 mm; 1 damaged 9
SM 129 30°54’S 30°31’E =850m 1 young 3, 2 29, all damaged
LBT 38J 32°07'S 16°31’E 440m 1 3: 4,8 mm

Holotype

Adult male, in the South African Museum, SAM-—A15683, collected by
the Pieter Faure in about 1900. Type locality: 800 m, off the Cape Peninsula
(34°25’S 17°45’E).

Description

Adult male, holotype, length 5,8 mm. Integument fairly thin and translucent
without obvious denticles or reticulations. Carapace (Fig. 7A) dorsoventrally
depressed with a single marginal carina around entire periphery except pos-
teriorly. Dorsal outline low and smoothly arched. Middorsal carina faintly
evident posteriorly only (Fig. 7B). Eyelobe small, eyeless. Pseudorostral lobes
short and expanded laterally.

All five pedigerous somites visible, first four slightly flanged laterally.
Abdominal somites cylindrical, together slightly shorter than cephalothorax.

Antenna | (Fig. 7C) fairly large. Flagellum 4-segmented, first bearing
numerous short aesthetascs. Accessory flagellum 3-segmented.

Antenna 2 (Fig. 7D) fairly short, reaching beyond posterior edge of cara-
pace. Segments short and poorly setose.

Maxilla 1 without palp.

Maxilliped 3 (Fig. 7E) fairly short. Basis unexpanded distally, subequal
in length to rest of limb. Merus slightly expanded, carpus large. Propodus and
dactyl slender.

Basis of pereiopod 1 (Fig. 7F) longer than rest of limb. Ischium as wide as
long, next three segments subequal in length.

Pereiopod 2 (Fig. 7G) slender, carpus longer than propodus and dactyl
together, armed with four stout spines.

Pereiopods 3 (Fig. 7H) and 4 similar. Basis slightly shorter than rest of
limb in pereiopod 3 and slightly longer in pereiopod 4. Exopods present.
Distal segments of pereiopod 5 missing.

Three pairs of normal pleopods present.

Uropods and telson as in female: parts of uropods missing from holotype.

Adult female, paratype, \ength 6.4 mm. As male, except as follows: cara-
pace (Fig. 7J) with a number of slight swellings and depressions, slightly more
elevated dorsally; middorsal carina better defined. No wider anteriorly than
posteriorly in dorsal view. First pedigerous somite wider and third narrower.

Distal segments of flagella of antenna 1 missing. Antenna 2 (Fig. 7K)
4-segmented. Distal segments of pereiopod | flatter and wider. Pereiopods 2 to 4
without exopods. Pereiopod 2 much more slender, carpus longer; bases of
pereiopods 3 and 4 longer and slightly thinner. Pereiopod 5 (Fig. 71) very

SOUTHERN AFRICAN CUMACEA: PART 3 159

Fig. 7. Platysympus depressus sp. nov.

Adult male, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Antenna 2.
E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Pereiopod 5.
Adult female, paratype. J. Lateral view. K. Antenna 2. L. Uropod and telson.

Scale line = 2 mm for A-B, J; 1 mm for C-I, K-L.

slender, basis equal in length to next three segments together; merus and carpus
subequal in length.

Telson (Fig. 7L) nearly twice length of preceding somite, wider proximally
than distally and serrated on edges; armed only with three small spines apically.
Peduncle of uropod longer than telson, subequal in length to endopod, serrated

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

on inner edge. Second segment of exopod with three slender distal spines. First
segment of endopod longer than next two together.

Remarks

P. depressus 1s most similar to P. typicus (Sars, 1869) and P. compressus
sp. nov. It may be distinguished from both in that it lacks a middorsal carina
on the carapace and from P. compressus by the latter being laterally rather
than dorsoventrally compressed, lacking low protuberances on the carapace
of the female and having a larger first antenna in the male. P. depressus differs
from P. typicus mainly in the shape of the carapace: the dorsal and marginal
carinae are better defined and the female lacks low protuberances on the cara-
pace in P. typicus. Also in this species the first antennae are smaller, the distal
segments of pereiopod | are flattened and the second pereiopod is more slender
with a longer dactyl.

Distribution
From Lambert’s Bay to northern Natal at depths from 440 to 850 m.

Platysympus compressus sp. nov.

Fig. 8
Records
SAM-A10601 (PF 12605) 30°33’S 30°58’E 805m 1 subadult 3: 4,8 mm
SM 60 27°09’S 32°58’E 800m 1 adult ¢: 5,5 mm (holotype):
3 dd: 5,8-6,1 mm; 3 ovig. 92:
5,8-6,1 mm
Holotype =

Adult male, in the South African Museum, SAM-A15681, collected by the
Meiring Naude, 19 May 1976. Type locality: 800 m, off northern Natal
(27°09'S 32°50’E).

Description

Adult male, holotype, length 5,5 mm. Integument smooth without reticula-
tions or denticles. Carapace (Fig. 8A) smooth with marginal carina as in
P. depressus, dorsoventrally depressed for the most part but the dorsal third
strongly compressed laterally, forming a very distinct, narrow middorsal carina.
Pseudorostral lobes narrowed anteriorly in lateral view, rounded and fairly
short in dorsal view (Fig. 8B). Eyelobe small, rounded and eyeless.

All pedigerous somites exposed, all of similar length. Abdominal somites
subcylindrical, together shorter than cephalothorax.

Antenna | (Fig. 8C) large, first two segments slightly serrated. Second
segment much wider distally than proximally, wider than long. Third segment
short and subconical. Accessory flagellum missing. First segment of flagellum
with numerous short aesthetascs, last three short and subequal in length.

Maxilla 1 without palp. Maxilliped 3 and pereiopods | and 2 represented
by bases only.

SOUTHERN AFRICAN CUMACEA: PART 3 161

Fig. 8. Platysympus compressus sp. nov.

Adult male, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Pereio-
pod 3. E. Pereiopod 5. F. Pleopod 2. G. Uropod and telson.
Ovigerous female, paratype. H. Lateral view. I. Pereiopod 2. J. Antenna 1.

Scale line = 2 mm for A-B, H; 1 mm for D-E, G, I-J; 0,5 mm for C, F.

Pereiopods 3 (Fig. 8D) and 4 similar, last two segments of each missing.
Basis stout, ischium short, merus and carpus subequal in length. Exopods
moderately large.

Pereiopod 5 (Fig. 8E) short and slender.

Three pairs of pleopods present. (Fig. 8F)

Last abdominal somite as wide as long, less than half length of preceding
one. Telson (Fig. 8G) slightly more than half length of peduncle of uropod,
serrated distally on lateral edges and with three small sharp spines apically.
Peduncle of uropod fairly stout, subequal in length to endopod with numerous
small sharp spines distally on inner edge. Exopod slightly longer than first

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

two segments of endopod, unarmed. First segment of endopod longer than next
two together with several small spines on inner edge. Second segment longer
than third.

Ovigerous female, paratype, length 6,1 mm. As male, except as follows:
carapace (Fig. 8H) slightly deeper, pseudorostral lobes less narrowed in lateral
vew. Pedigerous somites shorter, abdominal somites together subequal in
length to cephalothorax. Marsupium small.

Antenna | small, segments not expanded. Flagella both 2-segmented.
Pereiopod 2 slender, without exopod; carpus longer than last two segments
together. Segments distal to basis missing from maxilliped 3 and pereiopods 1, 3
and 4. Pereiopods 3 and 4 without exopods. Telson relatively longer—two-
thirds as long as peduncle of uropod—but otherwise as in male.

Remarks

Although the appendages of none of the individuals are complete, the
carapace is distinctively different from that of the other species in the genus.
The lateral compression of the dorsal part of the carapace is characteristic, as
is the large, wide first antenna in the male.

Distribution

Known only from depths between 800 and 810 m off northern Natal.

Platysympus camelus sp. nov.

Fig. 9

Records

SM 86 27°359'S'32°40'R 550'im 1 adult gd: 6,8 mm (holotype);
3 ovig. 22: 6,1-6,8 mm

WCD 450D 34°18 1870S°E «188 m 1 subadult 3: 3:93mmemt 2:
4,8 mm; 1 damaged ovig. 2

SST 1K 35° 22'S 22°31 E> 2005m 1 2: 3,9 mm

SST 17N 35,2279) 22,51 EE -2001m 1 damaged adult g; 1 subadult 3:
4,1 mm; 4 92: 4,1 mm

Holotype

Adult male, in the South African Museum, SAM-—A15684, collected by the
Meiring Naude, 22 May 1976. Type locality: 550 m, off northern Natal
(27°59'Si32 40 BE):

Description

Adult male, holotype, length 6,8 mm. Integument smooth, slightly trans-
lucent. Carapace (Fig. 9A) strongly dorsoventrally depressed at edges; slightly
compressed laterally in midline forming distinct dorsal carina undulating in
lateral view due to two rounded elevations, one middorsally and one postero-
dorsally. Marginal carina strong, forming a flattened flange around entire
carapace except posteriorly. Eyelobe (Fig. 9B) small, rounded and eyeless. All

SOUTHERN AFRICAN CUMACEA: PART 3 163

Fig. 9. Platysympus camelus sp. nov.

Adult male, holotype. A. Lateral view. B Dorsal view of carapace C. Antenna 1. D. Maxilli-
ped 3. E. Pereiopod 1. F. Pereiopod 2. G. Pereiopod 3. H. Uropod and telson.
Ovigerous female, paratype. I. Lateral view. J. Antenna 1. K. Uropod and telson.
Juvenile. L. Lateral view of carapace.

Scale line = 2 mm for A-B, I, L; 1 mm for C-H, J-K.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

pedigerous somites visible, of approximately equal length. Abdominal somites
fairly large, rounded, together subequal in length to cephalothorax.

Antenna | (Fig. 9C) fairly large and stout; third segment wider than long.
Both flagella 4-segmented, first segment of main flagellum surrounded by
numerous short aesthetascs.

Maxilliped 3 (Fig. 9D) short and fairly stout. Basis longer than rest of
limb, ischium wider than long. Carpus longer than ischium and merus together.

Bases of pereiopods | to 4 flattened, edges expanded by means of flat,
transparent scales fusing to form flanges. Basis of pereiopod | (Fig. 9E) slightly
longer than rest of limb, both inner and outer edges flanged distally. Ischium
small, merus and carpus subequal in length, both flanged on inner edge; pro-
podus slightly longer, dactyl short and cylindrical.

Basis of perelopod 2 (Fig. 9F) flanged distally on both edges. Ischium
half length of merus, together about half length of carpus. Propodus and dactyl
short and cylindrical.

Pereiopods 3 (Fig. 9G) and 4 similar, inner edges flanged. Basis distinctly
longer and stouter than rest of limb, of which merus, carpus and propodus are
subequal in length and cylindrical.

Basis of pereilopod 5 about half length of basis of pereiopod 4, distal
segments the same.

Three pairs of pleopods present.

Telsonic somite as wide as long, less than half length of telson. Telson
(Fig. 9H) tapering evenly from base, little more than half length of peduncle
of uropod, serrated distally on both edges and with three small apical spines.
Peduncle of uropod with several small sharp spines distally on inner edge.
First segment of exopod unarmed, second slightly serrated on both edges with
three small terminal spines. First segment of endopod slightly longer than
subequal second and third segments together.

Ovigerous female, paratype, length 6,8 mm. As male, except as follows:
carapace (Fig. 9I) relatively larger, undulations of dorsal surface more marked.
Pedigerous somites shorter, the first deeper. Carapace almost round in dorsal
view. First segment of antenna | (Fig. 9J) larger, third narrower. Both flagella
with three segments. Merus of maxilliped 3 longer and thinner, carpus stouter.
Distal segments of pereiopod 1 less flattened, edges not flanged, together
slightly longer relative to basis. Pereiopods 2 to 4 without exopods. Basis of
pereiopod 2 more slender, remaining segments longer and thinner. Bases of
pereiopods 3 and 4 more slender. Uropod slightly shorter and stouter, inner
edges of peduncle and rami serrated and with fewer spines.

Remarks

The peculiar flanged edges of the bases of pereiopods 1 to 4 and the undu-
lating dorsal edge of the carapace clearly distinguish this species from the others
in the genus. It should be noted that this undulation is most marked in juveniles

,

SOUTHERN AFRICAN CUMACEA: PART 3 165

(Fig. 9L) and least evident in adult males, although these are none the less easy
to distinguish.

The flanges on the pereiopods are occasionally found in members of other
families of Cumacea (e.g. Ceratocuma horridum, Fig. 16D). The functional
significance of this feature is uncertain. It may be to increase the surface area
(for digging?) with the smallest possible increase in weight.

Distribution
Northern Natal to the Cape Peninsula at depths from 188 to 550 m.

Bathylamprops Zimmer, 1908
Generic diagnosis

Carapace not flattened. Pseudorostral lobes large and acutely produced.
Eye absent. First and third segments of antenna | elongate, accessory flagellum
minute. Palp of maxilla | with two filaments. Pereiopods 3 and 4 of female with
small exopods. Male with three pairs of pleopods. Telson large and well
developed.

Type species
Bathylamprops calmani, Zimmer, 1908.

Remarks

The genus consists of three closely allied deep-water species, B. calmani
Zimmer, 1908, and B. natalensis Jones, 1969, both of which are known only
from the east coast of Africa, and B. motasi Bacescu & Muradian, 1976, found
off Florida. The genus is clearly recognized by the greater development of the
pseudorostrum than is usual in the family, and the large first antenna with the
minute accessory flagellum. It undoubtedly has close links with Hemilamprops,
which it resembles in general morphology.

Distribution of Bathylamprops

Deep waters between | 300 and 3 800 m off the east coasts of Africa and
the United States.

KEY TO THE SPECIES OF BATH YLAMPROPS

1 Telson more than three times length of telsonic somite; carapace with numerous low,

denticulate transverse ridges...... B. calmani Zimmer, 1908—east and south-east Africa
— Telson no more than two and a half times length of telsonic somite; carapace minutely
BoMMeUIALeEOL Smooth: bUL WIthOUt HIMES. oc. .c ec 0 Le ove ayers wis Perso ike oe aeras wont so ale leceus 2

2 Basis of maxilliped 3 twice length of remaining segments together, carpus no wider than
merus; exopod of uropod shorter than telson; uropodal rami subequal in length........
B. natalensis Jones, 1969 —Natal

— Basis of maxilliped 3 little longer than remaining segments together, carpus much wider
than merus; exopod of uropod longer than telson and longer than endopod............
B. motasi Bacescu & Muradian, 1976—Florida

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bathylamprops calmani Zimmer, 1908
Fig. 10
B. calmani Zimmer, 1908: 173-175, figs 60-70.

Records
SM 109 28°41’S 32°36’E 1300 m 1 adult 2: 15,4 mm

Previous records

Off Dar-es-Salaam, 2 959 m (Zimmer 1908); off Durban, 2 720-3 530 m
(Jones 1969).

Holotype

Damaged adult female, deposited by Zimmer in the Berlin Zoologisches
Museum. Type locality: 2 959 m, off Dar-es-Salaam (6°12’S 41°17’E).

Fig. 10. Bathylamprops calmani

Adult female. A. Lateral view. B. Detail of anterior end of carapace. C. Dorsal view of
carapace. D. Antenna 2. E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Telson and
uropod. I. Pereiopod 4.

Scale line = 4 mm for A—B; 2 mm for C-I.

SOUTHERN AFRICAN CUMACEA: PART 3 167

Description

Adult female, length 15,4 mm. Carapace (Fig. 10A) large, strongly vaulted
posteriorly and pointed anteriorly. Integument of carapace minutely denticulate
(Fig. 10B) forming numerous transverse ridges, particularly anteriorly. Pseudo-
rostral lobes elongate, denticulate immediately in front of eyelobe; denticles
also forming an indistinct and very short lateral carina behind the small and
indistinct anterolateral angle. Carapace narrow in dorsal view (Fig. 10C),
slightly depressed between a pair of large posterolateral expansions. Middorsal
carina present for a short distance behind the eyelobe. Eyelobe very small and
eyeless. Pseudorostrum about one-fifth of total length of carapace.

All five pedigerous somites short, visible. Cephalothorax slightly longer
than abdomen.

Antenna | (Fig. 10B) elongate, first segment more than twice length of
second, third slightly longer than second. Accessory flagellum minute,
2-segmented. Flagellum elongate, 3-segmented.

Antenna 2 (Fig. 10D) short, 4-segmented. First segment stout, second and
third small, fourth elongate (distal tip missing).

Maxilliped 3 (Fig. 1OE) very stout. Basis slightly longer than rest of limb;
ischium much wider than long; merus short and slightly expanded; carpus very
large and expanded; propodus and dactyl small and cylindrical.

Pereiopod | (Fig. 10F) elongate, basis longer than rest of limb. Ischium
small, merus slightly longer. Carpus, propodus and dactyl cylindrical and
elongate.

Pereiopod 2 (Fig. 10G) stout, basis shorter than rest of limb. Ischium
small; carpus long with several strong spines; propodus and dactyl small,
narrow.

Pereiopods 3 (Fig. 101) and 4 similar, basis and exopod of pereiopod 4
shorter. Exopod very well developed for a female, 2-segmented.

Pereiopod 5 shorter and more slender than pereiopod 4.

Telsonic somite (Fig. 10H) slightly widerthan long. Telson well developed,
more than three times as long as telsonic somite; pre-anal part short, distinctly
wider proximally; eight pairs of sharp spines distally on lateral edges and three
short ones terminally. Peduncle of uropod slightly longer than telson with
several small spines on inner edge. Exopod missing. First two segments of
endopod present, first unarmed, second very slightly serrated on both edges.

Adult males are unknown.

Remarks

From the shape of maxilliped 3 and the sculpturing of the carapace it is
clear that the present specimen belongs to the same species as Zimmer’s. How-
ever he figures the pseudorostrum of his unique, damaged specimen as being
somewhat shorter than that figured here. It is difficult to say if his was dis-
torted due to mutilation or whether the length is variable.

B. calmani is very similar to both B. natalensis and B. motasi. However,

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

in both of the latter the carapace lacks transverse rows of denticles, the exopods
of pereiopods 3 and 4 are much smaller and the telson is shorter. In B. natalensis,
too, the carpus of maxilliped 3 is not expanded and the fifth pereiopod is larger.

Distribution
Confined to deep waters off the east coast of Africa from northern Natal
to Dar-es-Salaam at depths from | 300 to 3 530 m.

Hemilamprops Sars, 1883
Generic diagnosis
Carapace not strongly dorsoventrally flattened. Eye present or absent.
Pseudorostrum short. Flagella of antenna | well developed. Palp of maxilla 1
with two filaments. Exopods on pereiopods 3 and 4 of female rudimentary.
Male with three pairs of pleopods. Telson well developed.

Type species
Not designated: Sars included H. rosea (Norman, 1863), H. cristata (Sars,

1870), H. uniplicata (Sars, 1872) and H. assimilis Sars, 1883, in his first descrip-
tion of the genus in 1883.

Remarks

Hemilamprops Sars, 1883, Mesolamprops, Given, 1964, and Lamprops Sars,
1863, are very closely-related genera, differing mainly in the number of pairs of
pleopods in the adult male: none in Lamprops, two pairs in Mesolamprops
and three pairs in Hemilamprops. This is the only character which invariably
separates the species of the three genera but there are some other differences
which are usually reliable in distinguishing them. For example, there is a
small but usually distinct antennal notch in Lamprops; it may be present or
absent in Mesolamprops and is usually absent in Hemilamprops. An eye is
present in Lamprops, variable in Mesolamprops and usually absent from Hemi-
lamprops. The proportions of the basis of pereiopod | to the rest of the limb are
perhaps most reliable. In Lamprops the basis is approximately equal in length
to the rest; in Mesolamprops it is slightly shorter and in Hemilamprops it is
distinctly shorter. Although this character appears to be constant, it is not
always of practical value since the distal segments of the pereiopods are fre-
quently lost or damaged, particularly in deep-water forms. However the com-
bination of the characters mentioned above should allow most individuals to be
placed in the correct genus. But it should be stressed that only the number of
pleopods in the male is genuinely diagnostic.

Distribution

The genus consists of 22 species, widespread in the Arctic, Antarctic,
Pacific, Atlantic and Southern Indian Oceans. The depth distribution is also
wide) trom! to 2) 725) 1m.

The species occur in three distinct groups geographically: one group of
eight species is found in the North Pacific, another of five species in the North

SOUTHERN AFRICAN CUMACEA: PART 3 169

Atlantic and Arctic and the remaining group of eight species in southern
oceans. Three species (two of which are new) in the last group are found in
southern African waters.

There are no records for tropical or subtropical waters and no species is
found both north and south of the tropics.

Rey tO LFHE SPECIES OF HEMILAMPROPS FROM THE
SOUTHERN HEMISPHERE

Since the tropics form a very distinct boundary between Northern and
Southern hemisphere species, a key is given only to those species occurring
south of 30°S.

1 Carapace somewhat carinate ventrolaterally, more than one and a half times as broad as
SSET yy Le SER OR coe eee H. lata Hale, 1946—Australia

— Carapace not carinate ventrolaterally, less than one and a half times as broad as deep... .2
2 Telson longer than peduncle of uropod; carapace robust, subtruncate anteriorly; pseudo-

rostral lobes not meeting anterior to eyelobe............ H. diversa Hale, 1946—Australia
— Telson shorter than peduncle of uropod; carapace delicate and elongate; pseudorostral
Sa wmiceuine Tor some distance anterior to eyelobe. . 6. i. 000s eee cc vce we ech ee sess 3

3 Telson with ten to twelve pairs of lateral spines; middorsal carina of female finely serrate;
carpus of pereiopod 2 long and slender, distinctly more than half length of basis; pseudo-
rostrum pointed anteriorly in lateral view... .H. pellucidus Zimmer, 1908 —Sovihern Ocean

— Telson with no more than six pairs of lateral spines; middorsal carina and pseudorostrum
padi. carpus of perciopod.2 less than half length of basis.............0...2.00--50 4

4 Anterior tip of pseudorostrum slightly uptilted; anterolateral edge and middorsal carina
bearing long, slender spines; telson less than half length of peduncle of uropod with five
pairs of small spines laterally and three long ones terminally........ H. sp.—South Africa

— Anterior tip of pseudorostrum not uptilted; carapace without spines middorsally or laterally
(may be denticulate); telson as long or almost as long as peduncle or uropod with zero to
Sees ou spines laterally and three to five terminally..............2..0ccecenseeeets 5

5 Telson with five long spines terminally and none laterally; carapace fairly broad and truncate
anteriorly in dorsal view; basis of maxilliped 3 widely expanded, carpus very large and
aap Ne: cs 3 0k as Sree aes dase eins Sera ible s way eee ee sl H. glabrus sp. nov.

— Telson with three long spines terminally and one to six pairs laterally; carapace fairly
slender and pointed anteriorly in dorsal view; neither basis nor carpus of maxilliped 3 much

fe ee I Os, oe dh a5 8.5, 3 cies hs Gre SMOSH gerade 6% US EOS Sees oe 6
6 Peduncle of uropod subequal in length to endopod....H. lotusae Bacescu, 1969— Argentine
— Peduncle of uropod subequal in length to exopod.................4. 7

7 Telson with six pairs of lateral spines; carapace finely denticulate middorsally............
H. serrulata Ledoyer, 1977—Kerguelen

— Telson with three pairs of lateral spines; carapace smooth middorsally................05
H. ultimaspei Zimmer, 1921—Tierra del Fuego

Hemilamprops pellucidus Zimmer, 1908
Figs 11-12

H. pellucidus Zimmer, 1908: 172-173, figs 53-59; 1913: 456-457. Stebbing 1912: 144-145,
pl. 52. Jones 1963: 52-53, figs 192-201; Jones 1969: 119.

Records

SAM-—A594 (PF 17386) S427 Sli Bi 849m 1 subadult ¢: 7,4 mm; 1 2:
8,0 mm

SAM-A595 (PF 15785) 34°39’S 18°10’E 500m 4 subadult gd: 5,8-8,0 mm;
4 ovig. 99: 8,3-8,6 mm; 9 99:
6,1-9,0 mm

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-A10601 (PF 12605) 30°33’S 30°58’E 805m ___1 subadult 3: 6,8 mm

SAM-A10602 (PF 17440) 34°25’S 17°45’E 800m 2subadult Jd: 7,0-8,0 mm; 1 2:
6,9 mm; 2 ovig. 99: 8,6-9,9 mm;
1 juv. 4,6 mm

SM 60 27°09’S 32°58’E 800m 1 adult gd: 7,8 mm; 2 subadult
3d: 5,8-7,0 mm; 3 99: 6,4-9,6
mm

SM 69 DHEAGESS BVLNS 18 660m _ 1 damaged g

SM 86 27°59’S 32°40’E 550m _ (6 adult $d: 7,4-8,6 mm; 6 sub-

adult 3d: 6,1-7,7 mm; 5 ge:
5,8-6,4 mm; 5 ovig. 92: 9,0-11,2
mm; 5 99: 5,4-8,3 mm; 5 juvs:

3,8-5,4 mm

SM 109 28°41’S 32°367E_ —«-:11300m_ =1 = subadult ¢: 7,4 mm: 1 2°:

' 7,4 mm

SM 129 30°53’S 30°31’E 850m _ I damaged ¢; 2 ovig. 29: 9,0-9,3
mm

SST 17P 35522 S122 3 1B 200m _ 1 2: 6,4 mm

LBT 67A 32°04’S 17°12’E 200m _ 1 subadult 3: 6,7 mm; 3 99:
5,8-7,0 mm

LBT 70C S2707 SAE 330m: 1 juves 531mm

LBT 72D 32°07’S 17°31’E 400m 3 subadult 3d: 5,8-7,7 mm

Previous records

Agulhas Bank, 126 to 596 m (Zimmer 1908, 1921); Chatham Rise, 238 to
535 m (Jones 1963); Antarctic, 2725 m (Zimmer 1913); Great Australian
Bight, 1 320 to 1340 m (Jones 1969); off Recife, Brazil, >1000 m (Jones
pers. comm.).

Holotype

Not designated: young male and female syntypes deposited in the Berlin
Zoologisches Museum. Type locality: 564 m, on the Agulhas Bank
(35°09’S 18°32’E).

Description

Adult male, length 7,4 mm (SM 86). Integument thin, pellucid, finely
reticulate. Gut-contents black. Carapace (Fig. 11A) elongate, twice as long
as deep. Pseudorostral lobes short, roundly pointed anteriorly. Antennal notch
a slight excavation below pseudorostrum. Eyelobe (Fig. 11B) small, pointed,
with several small denticles in midline. Carapace swollen posteriorly on either
side of shallow middorsal depression. Middorsal carina evident anteriorly,
denticulate only on eyelobe. Carapace less than half length of rest of body.
Pedigerous somites all visible, not flanged laterally. Abdominal somites
cylindrical.

First segment of antenna 1 (Fig. 11C) longer than next two together.
Flagellum 4-segmented with several small aesthetascs at base; accessory flagellum
3-segmented.

Flagellum of antenna 2 (Fig. 11D) reaching almost to end of body.

Palp of maxilla 1 with two filaments.

SOUTHERN AFRICAN CUMACEA: PART 3 171

Fig. 11. Hemilamprops pellucidus

Adult male. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Antenna 2.
E. Maxilliped 3. F. Pereiopod 2. G. Pereiopod 3. H. Pereiopod 5. I. Pleopod 2. J. Uropod and
telson.

Scale line = 4 mm for A-B; 2 mm for C-J.

Basis of maxilliped 3 (Fig. 11E) slightly longer than rest of limb. Carpus
stout and slightly expanded. Propodus about twice length of dactyl.

Distal segments of pereiopod 1 missing from all specimens. Exopod large
and very well developed.

Basis of pereiopod 2 (Fig. 11F) little more than half length of rest of limb.

Carpus elongate with several fine spines. Exopod large and almost circular in
outline.

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pereiopods 3 (Fig. 11G) and 4 similar. Basis of pereiopod 3 very large and
stout, of pereiopod 4 less so.

Pereiopod 5 (Fig. 11H) shorter, basis subequal in length to rest of limb.

Three pairs of typical pleopods present (Fig. 11I).

Telsonic somite (Fig. 11J) less than half length of preceding somite, about
two-thirds length of peduncle of uropod. Pre-anal part of telson less than half
of total length, serrated laterally. Post-anal part narrower with ten to twelve
pairs of small lateral spines and three terminally.

Peduncle of uropod slender with many fine spines on inner edge. Distal
portions of both rami missing from all males.

Adult female, length 11,2 mm (SM 86). As male, except as follows: eyelobe
(Fig. 12A) without serrations; middorsal carina very finely serrate. Carapace
(Fig. 12B) considerably wider posteriorly than anteriorly.

Antenna | (Fig. 12C) smaller, without aesthetascs. Antenna 2 (Fig. 12D)
3-segmented. Maxilliped 3 (Fig. 12E) stouter, carpus relatively longer. Exopods
of pereiopods much smaller. Basis of pereilopod | (Fig. 12F) strongly serrate,
subequal in length to next four segments together; propodus and dactyl long,
slender and subequal in length. Basis of pereiopod 2 (Fig. 12G) shorter, carpus

Fig. 12. Hemilamprops pellucidus

Adult female. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1. D. Antenna 2.
E. Maxilliped 3. F. Pereiopod 1. G. Pereiopod 2. H. Pereiopod 3. I. Uropod and telson.

Scale line = 4 mm for A-B; 2 mm for F-I; | mm for C-E.

SOUTHERN AFRICAN CUMACEA: PART 3 173

longer. Bases of pereiopods 3 (Fig. 12H) and 4 shorter and more slender;
exopods 2-segmented.
Telson (Fig. 121) slightly longer, peduncle of uropod with fewer spines.

Remarks

This species is readily identifiable by the pellucid nature of the integument,
nearly always resulting in the gut being clearly visible: in the present specimens
it is black or dark brown. It closely resembles a number of other species in
which the carapace is of similar shape, but may be distinguished from them as
follows: H. cristata (Sars, 1869), H. glabrus sp. nov., H. ultimaespei Zimmer,
1921, H. lotusae Bacescu, 1969, H. serrulata Ledoyer, 1977, and Hemilamprops
sp. all have no more than five pairs of lateral spines on the telson. H. normani
Bonnier, 1896, has a shorter telson and uropodal peduncle and the middorsal
carina is slightly serrated in the male. H. pellucidus most closely resembles
H. tanseiana Gamo, 1967, differing from it in the greater number of lateral
spines on the telson, the reduced serrations on the middorsal carina, the longer
uropod and telson and the larger and more expanded merus and carpus of
maxilliped 3 in H. pellucidus.

The individuals figured by Stebbing (1912) and Zimmer (1908) differ in
several respects from each other and also from those figured here. The shape
of the carapace in subadult males differs slightly, as does the length of the
telson and its number of lateral spines. Adult males also have a longer telson
and more, but smaller, spines on the peduncle of the uropod in the present
specimens. Other differences can be attributed to varying age and sex. The
adult female differs from that figured by Zimmer in the length of the propodus
and dactyl of pereiopod | and from Stebbing’s as well as from Zimmer’s in the
slightly longer and stouter telson. However, it seems that the specimens from
SAM-A594 are the selfsame ones described and figured by Stebbing, and in fact
these fit well within the range of variation of the other individuals available
for examination. Thus it is clear that the present specimens can be referred to
Zimmer’s H. pellucidus without any great doubt.

Distribution
This is one of the few species of Cumacea which occurs in southern African

waters without being endemic. It appears to be widespread throughout the
southern oceans at depths from 126 to 2 725 m.

Hemilamprops glabrus sp. nov.

eels
Records

SM 109 23-AlS 32-368 1300m" 1 ove: 2: 7.0 mm holotype):
2 92: 6,4-7,2 mm; 2 mancas:
2,9-3,2 mm

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

Ovigerous female, in the South African Museum, SAM-—A15680, collected
by the Meiring Naude, 25 May 1976. Type locality: 1 300 m, off northern Natal
(28°41'S 32°36’E).

Description

Ovigerous female, holotype, length 7,0 mm. Integument rather thin and
translucent. Carapace (Fig. 13A) fairly short, swollen posteriorly on either side
of shallow middorsal depression. Pseudorostral lobes short and truncate
anteriorly with an indistinct anterolateral carina running posteriorly for a short
distance, bearing a few denticles above. Carapace in dorsal view (Fig. 13B)
slightly flattened, distinctly broader than deep. Eyelobe small, rounded and

Fig. 13. Hemilamprops glabrus sp. nov.

Ovigerous female, holotype. A. Lateral view. B. Dorsal view of carapace. C. Antenna 1.
D. Maxilliped 3. E. Pereiopod 2. F. Pereiopod 3. G. Pereiopod 4. H. Pereiopod 5. I. Uropod
and telson.

Scale line = 2 mm for A-B; | mm for C-I.

SOUTHERN AFRICAN CUMACEA: PART 3 175

eyeless. Pedigerous somites rather deep, all visible. Abdominal somites cylin-
drical, together subequal in length to cephalothorax. Marsupium small.

Antenna | (Fig. 13C) fairly short, both flagella 3-segmented, third segment
of accessory flagellum very short.

Palp of maxilla 1 with two filaments.

Basis of maxilliped 3 (Fig. 13D) very stout, slightly shorter than rest
of limb: no more than three times as long as wide and particularly expanded
distally. Merus wide, carpus large and flattened.

Segments distal to basis missing from pereiopod | in all specimens.

Pereiopod 2 (Fig. 13E) slender, basis subequal in length to rest of limb;
carpus relatively short, subequal in length to propodus and dactyl together.

Pereiopods 3 (Fig. 13F) and 4 (Fig. 13G) similar. Exopods very small,
2-segmented. .

Pereiopod 5 (Fig. 13H) short, basis shorter than rest of limb.

Telsonic somite (Fig. 131) almost square in dorsal outline, more than half
length of telson. Telson slightly shorter than peduncle of uropod, wider proxi-
mally; lateral edges entirely smooth; apex with five long stout complex spines.
Peduncle of uropod with several small sharp spines on inner edge. Rami missing
from all specimens.

Males are not available.

Remarks

It has already been remarked that without mature males it is not possible
to place species conclusively in Lamprops, Mesolamprops or Hemilamprops.
In this case the first pereiopod cannot be used either since the distal segments
are missing in all specimens. However, the general appearance of the specimens,
together with their locality, strongly suggests that they should be placed in
Hemilamprops. In some respects, they do bear a slight resemblance to Lamprops ?
comata Zimmer, 1907 from deep water off Tierra del Fuego, but since this
species is known only from a single fragmentary female, no reasonable com-
parison is possible.

Within Hemilamprops, H. glabrus is easily distinguished by the short, stout
basis of maxilliped 3 and the shape of the carapace. From those species which it
most closely resembles it may be distinguished as follows: from H. pellucidus
Zimmer, 1908, H. cristata (Sars, 1869), H. tanseiana Gam6é, 1967, H. normani
Bonnier, 1896, H. serrulata Ledoyer, 1977, and Hemilamprops sp. by the
absence in H. glabrus of lateral spines on the telson and denticles on the mid-
dorsal carina; from H. ultimaespei Zimmer, 1921, and H. lotusae Bacescu, 1969,
in that it has five terminal spines on the telson and none at all laterally and that
its carapace is much broader and truncate anteriorly.

Distribution

Known only from the type locality: northern Natal at 1 300 m.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hemilamprops sp.

Fig. 14
Records
SAM-A10607 (PF 16982) 1200m 34°40’S 17°50’E—__ 2: ovig. 22: 9,6-10,2 mm; 1 9:
9,1 mm; 1 damaged

Remarks

Although four individuals of this species are available, all of them are too
badly damaged to allow an adequate description. Figure 14A is a composite
drawing of the undamaged parts of both ovigerous females.

None the less the carapace and telson are quite distinct from those of any
other known species. In particular, the denticles anteriorly along the mid-
dorsal carina and on the anterolateral margin of the carapace (Figs 14A and
14B) are longer and sharper than those of any known species, while the telson
(Fig. 14C) is characteristically short, with the three terminal spines particularly
long and slender. In these characters it is unique in the genus, so that it should
not prove difficult to identify further specimens as belonging to the same
species.

It is tentatively placed in Hemilamprops for the same reasons as those
given above for H. glabrus. However, the same cautionary note must be sounded
until adult males are available to confirm its generic position.

Distribution

Known only from a single sample from a depth of 1 200 m off the Cape
Peninsula.

Fig. 14. Hemilamprops sp.
Adult female. A. Lateral view. B. Dorsal view of carapace. C. Uropod and telson.
Scale line = 4 mm for B; 2 mm for A; 1 mm for C.

SOUTHERN AFRICAN CUMACEA: PART 3 ‘77

DISTRIBUTION OF LAMPROPIDAE

With only fifty-eight species, the family is rather small, constituting less
than 10 per cent of known species of Cumacea. Table 1 details the world-wide
distribution of lampropids according to latitude and depth. It can be seen that
the family is generally confined to deep and/or cold waters and has a bipolar
distribution. 58 per cent of the species-records are from latitudes north of 20°N,
6 per cent between 20°N and 20°S and 36 per cent south of 20°S. Thus very few
species are known from the tropics: one of these is recorded from 390 m and the
rest from depths greater than 800 m, where temperatures are considerably lower
than on the surface. This pattern of distribution shows a direct contrast to
that of the Bodotriidae where 38 per cent of the species are found in the tropics
(between 20°N and 20°S) (Day 1978).

TABLE 1. Distribution of Lampropidae according to depth and latitude (data mainly from

Jones 1969). Species may be entered more than once if they have been recorded from widely

different depths or localities. The entry marked * is also entered under ‘5-200 m’ and is,
therefore, excluded from the total count of species.

Shore—5 m 5-200 m 200-2000m >2000m Total

no. Yi, NOY, Yh, 50X)5 Ye © 500): Yue 10. Us
INEoEdOON -. ... 1 ae Il ae 0 0 0 2 3
BOSTOGN . . 1 bee 18 9 13 0 0 Dp 32
2052 int 16 3 5 2 3 16 23
ZON=20°S . . ltt 0 0 3 5 1 1+ 4 6
PO=S0S . «4 . 3 5 12 18 1 1+ 16 24
SS D 3 0 0 0 0 2 3
SorvOs of 5. 1 1+ 2 3 3 5 6 9
Total no. of records 2 3 30 44 29 43+ 7 10+ 68 100
Total no. of species ic 1+ 26 44 OR) 42 8 13 59 100

Assuming that the distribution of the lampropids is indeed limited by
water temperature, then those species living in shallow water (less than 200 m)
would be expected to occur at higher, cooler latitudes. This is shown by the
fact that there are no records for shallow waters between 20°N and 20°S, which
is not merely a reflection of collecting effort, since 36 per cent of bodotriids
are found in the same situation.

Similarly it can be shown that the lampropids preponderate in northern
waters, since there are nearly twice as many species (58 %) recorded north of
20°N as there are south of 20°S (36 %). Percentages for the bodotriids are
again in inverse proportion to this: 24 per cent occur north of 20°N and 37 per
cent south of 20°S.

Fully 55 per cent of lampropids are recorded from depths greater than
200 m (less than 8 % for bodotriids) and 12 per cent below 2 000 m (less than
1 % for bodotriids).

Thus the lampropids form a bipolar group, preferring deep, cold water and
avoiding the tropics. Because the collecting effort has been minimal in deep
waters, it is suggested that the number of species of lampropid is artificially

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

low. It is predicted that the number of new species in this family will increase
rapidly if more collecting is done in deep waters. This prediction is supported
by the fact that in the present paper, of the ten species described, six are new.

Of the eleven genera, four (Mesolamprops Given, 1964, from California,
Archaeocuma Bacescu, 1972, from Peru, Chalarostylis Norman, 1879, from the
North Atlantic and Stenotyphlops Stebbing, 1912, from South Africa) are
monotypic and their distribution need concern us no further.

Bathylamprops Zimmer, 1908, consists of three species from deep waters,
two off the east coast of Africa and one off Florida, and Pseudodiastylis Calman,
1905, of two species from deep tropical waters. The remaining genera together
consist of fifty species.

Lamprops Sars, 1863, contains twelve species, all from shallow waters at
depths less than 200 m and all from the northern hemisphere. Three of these
occur mainly between 20° and 50°N and the rest are found north of 50°N,
particularly in the region of the Bering Strait. The possible affinities of Lam-
props? comata are discussed above. Lamprops fasciata 1s the only species in the
family to have been found intertidally.

Hemilamprops Sars, 1883, consisting of twenty-one species, is very wide-
spread, representatives being found from the Arctic to the Antarctic, from the
Pacific, Atlantic, Indian and southern oceans at depths from 8 to more than
2 000 m. The species fall into three groups: 8 species occur in the North Pacific
(Japan to California), 5 in the North Atlantic and Arctic and 8 around South
America, South Africa and Australasia. None occurs between 30°N and 30°S.
The genus is bipolar, and follows the distribution pattern shown by the family
as a whole. =

Platysympus Stebbing, 1912, now consists of 7 species, all from waters
deeper than 200 m. 2 species are known from the North Atlantic and Norway,
and 5 from the Southern hemisphere— 1 from the Antarctic and the 4 new species
from South Africa.

Paralamprops Sars, 1887, consists of 10 deep-water species. 3 occur in the
North Atlantic from 600-3 789 m, 6 in the southern ocean from South Africa
to the Antarctic at 232-3 423 m, and | in the East Indies at 390 m.

Many of the species are known only from a single record and since so little
collecting has been done in deep water where many species normally occur, it is
predictable that the actual distribution of species will prove to be much wider
than it appears at present.

DISTRIBUTION OF THE SOUTHERN AFRICAN LAMPROPIDAE

Eleven species of lampropid are now known from these waters. Since they
are found only at depths greater than 200 m (with a single exception at 188 m),
and very little collecting has been done in deep water off southern Africa, it is
difficult and perhaps misleading to distinguish any clear distribution patterns.
The frequency of occurrence is also uncertain, because earlier records are

SOUTHERN AFRICAN CUMACEA: PART 3 179

incomplete and Cumacea are not caught in all samples, sometimes because the
substratum is unsuitable (rock or very coarse sand) or simply because cumaceans
are scanty for some other unknown reason. There is also the simple fact that
no sampling has been done off the west coast north of Lambert’s Bay or off
Mozambique at depths greater than 200 m. However, of the 12 grabs taken
by UCT at these depths around the coast, 6 contained lampropids, as did
7 of the ten SM samples which contained any Cumacea at all. The incomplete-
ness of the early Pieter Faure records is such that it is not possible to determine
the exact type of collecting gear nor the number of stations. But from the
previous two sets of figures it seems that lampropids are not at all uncommon
deep-water forms, although they are entirely absent from waters shallower
than 188 m.

In this way the South African lampropids differ from the Northern hemi-
sphere species, of which more than 60 per cent are shallow-water forms. This
is no doubt due to the fact that shallow waters in southern Africa are relatively
warm.

The southern African species do not fall into any obvious groups: tempera-
ture conditions in waters deeper than 200 m do not vary much round the coast
and there are many stretches of coast which have been sampled poorly or not
at all. In fact, there are only five rather small regions in which sampling has
been at all comprehensive. These are off Lambert’s Bay, the Cape Peninsula,
Still Bay, Durban, and in a fairly wide area off northern Natal.

Two species occur throughout the range: Platysympus depressus (21 speci-
mens) and Hemilamprops pellucidus (79 specimens), which are also the two
most common species in most samples. (This indicates, incidentally, that breaks
in the ranges of most species are due to a paucity of numbers rather than the
realistic limits of very confined ranges.) Two further species occur from the
Cape Peninsula to northern Natal: Stenotyphlops spinulosus (17 specimens) and
Platysympus camelus (14 specimens). The other six species, Paralamprops
peringueyi (14 specimens), Paralamprops margidens (6 specimens), Platysympus
phylloides (4 specimens), Platysympus compressus (8 specimens), Bathylamprops
calmani (1 specimen), Hemilamprops glabrus (5 specimens) and Hemilamprops sp.
(4 specimens) occur in only one or two regions and are therefore of little value in
Zoogeographic terms. The only one for which there is some little evidence for
a really restricted range is Paralamprops peringueyi which is not uncommon off
the Cape Peninsula (14 specimens in 3 samples) but has not yet been found
anywhere else.

Only two of the ten species are known outside South African waters: the
type specimen of Bathylamprops calmani was found off Dar-es-Salaam, and
Jones (1969) has since recorded three specimens off Durban. The type locality is
at a depth of 2 959 m and the other depth records are 2 720 and 3 530 m, sug-
gesting that the present specimen from | 300 m was at about the upper depth
limit for the species. H. pellucidus is one of the most widespread of southern
African Cumacea, occurring in southern oceans from the Chatham Rise, the

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Antarctic, Brazil and the Great Australian Bight as well as being by far the most
common lampropid in local waters, constituting half of the individuals in the
present collection.

Little can be deduced about depth distributions. In some cases there is
a slight tendency for individuals to occur in shallower waters off the cold
west coast (P. depressus at 440 m off Lambert’s Bay, and 550 to 680 m off
northen Natal; P. camelus at 188 m off the Cape Peninsula, and 550 m off
northern Natal; H. pellucidus at 200 to 400 m off Lambert’s Bay, and 550 to
1 300 m off northern Natal). However, collecting on the west coast has generally
been in shallower waters than on the east coast so the differences may be more
apparant than real.

There is a slight indication of a change in the fauna at very roughly 800
and 1 400 m. Four species (Platysympus depressus. P. compressus, P. camelus
and Paralamprops margidens) occur only at depths less than 850 m; three species
(S. spinulosus, P. peringueyi and P. phylloides) are found between 300 and
1 400 m and three (B. calmani, H. glabrus and Hemilamprops sp.) at 1 200 m or
more. Here again valid conclusions are limited by the fact that the greatest
depth at which sampling occurred was | 400 m.

The species-diversity of the family is fairly high. In comparison with the
Bodotriidae (which are by far the most abundant locally in terms of numbers,
both of individuals and of species (Day 1978)) the figures are as follows: the
Bodotriidae, with 4 582 specimens, 42 species and 649 records have a ratio of
7,1 individuals per record and a specimen : species ratio of 109. The Lampropi-
dae, with 169 specimens, 11 species and 39 records have a ratio of 4,3 indi-
viduals per record and a specimen: species ratio of 15,4: Thus lampropids
occur in fewer samples and are far less abundant where they do occur, but they
are far more diverse than are the bodotriids.

Family Ceratocumatidae Calman, 1904

Diagnosis

Five free pedigerous somites. Mandibles narrow (boat-shaped) at base.
Maxillipeds 2 and 3 elongate, 7-segmented, basis not produced distally. Exopods
present at least on maxilliped 3 and pereiopod 1 in both sexes. Propodus of
pereiopod 1 with two lobular setose processes. Male with four to five pairs of
pleopods. Telson small, unarmed and flap-like, hinged to cover anal valves.
Endopod of uropod 1-segmented, exopod 2-segmented with first segment very
short.

Type genus
Ceratocuma Calman, 1904.

Remarks

This is the smallest and most recently erected of the seven cumacean
families. The combination of four or five pairs of pleopods in the male, together

SOUTHERN AFRICAN CUMACEA: PART 3 181

with a small telson and the setose lobes on the propodus of the first pereiopod, is
quite characteristic. However, the telson is very small and is often held flapped
over the anal valves so that it is sometimes difficult to detect.

Until 1969, the family was known from four specimens of a single species
in two records, one from the north-western coast of Ireland at 705 m and one
from about 800 m off Durban. Examination of material from deep waters in the
Atlantic has increased numbers dramatically in the last few years so that the
family is now known from hundreds of specimens in seven species and two
genera. It is expected that further exploration of deep and abyssal waters will
continue to provide useful information on this family.

Ceratocuma Calman, 1904
Generic diagnosis

Body elongate, carapace dorso-ventrally flattened with protuberances.
Exopods present on pereiopod | in both sexes and on pereiopod 2 in the male.
Pereiopods 2 to 4 very long and slender, pereiopod 5 absent. Uropods slender
and elongate.

Type species
Ceratocuma horridum Calman, 1904.

Remarks

Ceratocuma is easily distinguished from Cimmerius, the only other genus in
the family, the generic diagnosis of which reads as follows: Body not elongate.
Carapace rounded, reminiscent of Campylaspis. Exopods present on pereiopods
1 to 4 in both sexes, reduced on pereiopod 4 in female. Pereiopod 5 present.
Pereiopods and uropods not particularly elongate or slender.

In general appearance the two genera are quite different, although they
both possess the familial characteristics.

Distribution of Ceratocuma

The four species are all known from the Atlantic and southern Indian
oceans: one from Ireland and Natal, one from the Puerto Rico Trench, one
from the Azores, and one from Panama, but all at depths of 680 m or more.

KEY TO THE SPECIES OF CERATOCUMA

1 About eighteen pairs of long, sharp, slender spines on carapace, others on posterior somites;
carapace oval in dorsal view and rounded anteriorly (ignoring spines); telsonic somite less
pianenalk lensth of peduncle of uropod..........2.....++- C. reyssi Jones, 1973 — Azores

— Protrusions of carapace (about ten pairs or fewer) and posterior somites confined to blunt
spines or rounded tubercles; carapace more or less rectangular in dorsal view; telsonic
Mmiteinalbicnstnor peduncle of uropod OF MOL; cs. i.e see ewe ee eee cave sete’ z

2 Carapace without paired dorsolateral tubercles or blunt spines: at most with three pairs of
Home OnaDUt extensive. FOUNGEE «PEOLUSIONS © 4 o0/6.5 ay.0s ass ate ocsce BOR UK Sours ayes ho crete ls Sele

C. amoena Jones, 1969— Puerto Rico Trench, off Surinam, Bay of Biscay

— Carapace with several pairs of dorsolateral protrusions, tubercles or blunt spines......... 3

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

3 Carpus of pereiopods 3 and 4 twice length of propodus; ischium and merus of pereiopod 1
together about half Jength of carpus..2....5..60s00-c000+ 4: 03) 7 eae
C. panamensis Bacescu & Muradian, 1974—off Panama

— Carpus and propodus of pereiopods 3 and 4 subequal in length; ischium and merus of
pereiopod 1 together subequal! im length to carpus... .44..2 4.) a. 4s eee 4

4 All tubercles on carapace, including anterior ones, sharply hooked... <3... 2.) eee
C. horridum horridum Calman, 1904—North Atlantic

— Tubercles on anterior half of carapace low and bluntly rounded, on posterior half blunt or
Sharply drooKed Heide ce ius hs feo Cea ae Creer aoe C. horridum australe subsp. nov.

Ceratocuma horridum Calman, 1904

C. horrida Calman, 1904: 37-40, pl. 55 (figs 57-75). Stebbing, 1913: 51-52.

Holotype

Not designated: three male syntypes, two of them mature, deposited by
Calman in the British Museum (Natural History). Type locality: 705 m, off
Northern Ireland.

Ceratocuma horridum australe subsp. nov.
Figs 15-16
C. horridus: Stebbing, 1912: 142-143.

Records

SM 103 28°31’S 32°34’E 680 m 1 adult g: 3,5 mm (holotype); 2 subadult gd:
2,9 mm; 1 ovig. 2: 3,7 mm; 2 juvs: 2,8 mm

SM 129 30°53’S 30°31’E 850 m 1 adult ¢: 4,5 mm; 1 damaged ovig. 2

SM 151 30°14’S 30°27’E 900 m 1 ovig. 9: 5,1 mm; 2234 none

Previous records
Natal, 805 m (Stebbing, 1912).

Holotype

Adult male, in the South African Museum, SAM-—A15722, collected by the
R.V. Meiring Naude, 24 May 1976. Type locality: 680 m, off northern Natal
(238° 31S)32 34):

Description

Adult male, holotype, length 3,5 mm (SM 103). Integument pale and thin,
brittle, faintly reticulate. Carapace (Fig. 15A) large, slightly flattened dorso-
ventrally, somewhat wider than deep and nearly twice as long as deep; irregu-
larly sculptured to form a number of low, rounded projections. In lateral view,
nine pairs visible: one large pair posterodorsally, four closely-adjacent pairs
posterolaterally, one pair anterolaterally, one pair immediately above the
anterolateral angle, one pair middorsal and one pair midlateral. The same
protuberances visible dorsally (Fig. 15B), the four outermost pairs forming the
lateral edges of the carapace and causing the lateral outline to be broken;

SOUTHERN AFRICAN CUMACEA: PART 3 183

Fig. 15. Ceratocuma herridum australe subsp. nov. (SM 103)

Adult male. A. Lateral view. B. Dorsal view. C. Antenna 1. D. Antenna 2. E. Maxilliped 1.
F. Maxilliped 3. G. Pereiopod 1. H. Distal tip of pereiopod 1. I. Pereiopod 2. J. Pleopod 3.
Ovigerous female. K. Lateral view. L. Antenna 2. M. Disial tip of pereiopod 1.

Scale line = 1 mm for A—B, D, F-G, I, K; 0,5 mm for C, E, J, L—-M; 0,3 mm for H.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

slightly medial to these are two pairs and the last three pairs are arranged in
longitudinal rows just lateral to midline. Middorsal carina absent. Eyelobe
small and eyeless. Carapace slightly less than twice length of pedigerous somites
together.

All five pedigerous somites visible, first slightly wider than deep without
lateral expansions; second, third and fourth expanded ventrolaterally forming
wide, pointed lateral extensions and dorsolaterally forming smaller, low pro-
tuberances. Last pedigerous somite without appendage, small and similar to
abdominal somites. First four abdominal somites produced to slight points
dorsolaterally, fifth elongate, sixth small and wider than long or deep. Cephalo-
thorax longer than abdomen.

Antenna | (Fig. 15C) short, first segment longer than next two together.
Accessory flagellum very small, l-segmented. Flagellum short, 3-segmented
with two aesthetascs.

Antenna 2 (Fig. 15D) shorter than carapace with short, poorly-setose
segments.

Palp of maxilla 1 with two filaments.

Maxilliped 1 (Fig. 1SE) with a single branchial leaflet; carpus very widely
expanded.

Maxilliped 3 (Fig. 15F) normal, merus and carpus very slightly expanded.

Basis of pereiopod 1 (Fig. 15G) subequal in length to next three segments
together. Ischium longer than merus, together subequal in length to carpus.
Carpus elongate with laminar expansion on inner edge. Propodus (Fig. 15H)
with two setose lobes, one at midlength and one subterminally. Dactyl narrow,
shorter than propodus, with a number of small spines.

Pereiopod 2 (Fig. 151) elongate with small exopod. Basis shorter than
rest of limb; ischium very short, merus little longer. Carpus elongate, half
length of propodus and dactyl together, unarmed. Terminal spine on dactyl
with flattened tip.

Pereiopods 3 and 4 elongate, without exopod. Pereiopod 5 lacking
(suppressed).

Telson very small, usually folded over anal valves; rounded when extended.

Peduncle of uropod much shorter than rami, serrated on inner edge. Both
rami serrated on inner edge, otherwise poorly armed: both appear broken at tip.

Ovigerous female, paratype, length 3,7 mm (SM 103). As male, except as
follows: carapace (Fig. 15K) with protuberances better developed, also nine pairs:
one large pair posterolaterally; two sets of three pairs arranged transversely, the
most dorsal pair in each case also being most anterior; two pairs laterally
immediately behind anterolateral corner. First two pedigerous somites wider
and deeper, all with much reduced lateral expansions, and dorsal expansions
hardly evident. Marsupium large and well developed.

Antenna 2 (Fig. 15L) small and apparently 2-segmented. Second and third
segments of antenna | slightly smaller. Propodus of pereiopod 1 (Fig. 15M)
smaller, one of the setose lobes at distal tip; dactyl inserting subterminally

SOUTHERN AFRICAN CUMACEA: PART 3 185

about a third of total length from distal tip. Pereiopod 2 without exopod.
Telson slightly rounder and broader. Rami similar but distal tips missing.

Adult male, length 4,5 mm (SM 129). At first glance the two individuals
from SM 129 and the ovigerous female from SM 151 look very different from
the rest. However, these differences appear to be confined to the sculpturing of
the carapace, and the appendages (except for the distal segments of pereiopod 1)
are identical in all important respects. The sizes of the individuals also vary
quite considerably. But it does not seem possible to pinpoint any really signifi-
cant differences which would allow specific differentiation and it would there-
fore seem that we are dealing with a single polymorphic species.

The adult male from SM 129 (Fig. 16A) differs from that figured in Fig. 15A
as follows: the five posterior pairs of protuberances on the carapace are drawn
out to form distinct points and the anterior pairs are either very much reduced
or absent. The tips of these points are very delicate, almost transparent and
very easily broken off, so that the apparent degree of development of these
may not be of great significance. The expansions of the second and third pedi-
gerous somites and the first four abdominal somites are very much better
developed: these too are easily damaged and one is broken off. In dorsal view
(Fig. 16B) the second lateral pair of protuberances from the anterior end is

Fig. 16. Ceratocuma horridum australe subsp. nov. (SM 129)
Adult male. A. Lateral view. B. Dorsal view of carapace. C. Pereiopod 1. D. Distal tip of

pereiopod 1.
Scale line = 2 mm for B; 1 mm for A, C; 0,5 mm for D.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

much smaller, the third and fourth better defined and more pointed, as are the
two posterodorsal pairs. The ovigerous female is the same in all respects as the
male.

Pereiopod | of the male (Fig. 16C) differs slightly in that the inner edge
of the ischium, merus and carpus have laminar expansions, the carpus is nearly
one and a half times the combined length of the ischium and merus and the
propodus is shorter (Fig. 16D). The peduncle of the uropod is also slightly
longer relative to the fifth abdominal somite.

Remarks

The systematic position of these individuals is not indisputable. The
appendages of all four species known so far are very similar, while the sculp-
turing of the carapace in particular and the rest of the body in general varies
quite considerably. C. reyssi Jones, 1973, from the Azores, can easily be dis-
tinguished by the numerous long, slender spines, not only on the carapace but
also on the first, fourth and fifth pedigerous and all abdominal somites except
the last. The pseudorostrum, too, is pointed anteriorly without a broad, flanking
projection on either side. There are other minor differences in the proportions
of the limbs as well, so that this species is clearly distinct from the others.

C. amoenum was described by Jones (1969) on the basis of the cephalo-
thorax of a single male from the Puerto Rico Trench. He has since (pers. comm.)
found undamaged specimens of the same species from Surinam and the Bay of
Biscay in which the sculpturing of the carapace is the same as that figured
and which species he confirms to be easily distinguishable from C. horridum
in all cases. It is further distinguished by ‘the uropod peduncles being relatively
long (compared with C. horridum) in proportion to the rami’, and some fully
adult males have only four pairs of pleopods. C. amoenum, then, is also clearly
distinct.

C. panamensis Bacescu & Muradian, 1974, from north-east of Panama,
differs from C. horridum in the smaller number of protuberances on the carapace
and thorax and the proportions of the distal segments of the pereiopods. As the
authors point out, it is very close to C. horridum; certainty about the validity
of the species will have to await the collection of adult males.

C. horridum was described by Calman (1904) on the basis of three males,
two of which were mature. Jones (pers. comm.) has since found large numbers
in the North Atlantic and off Surinam, all of which clearly belong to Calman’s
species. Stebbing (1912) identified (but did not figure) a single specimen from
Durban as C. horridum, saying that it differed from Calman’s description and
figures only ‘in a small bulbous expansion of the base of [the] peduncle (of the
uropod)’. The several other specimens now available from South Africa are
problematical. Some approach C. amoenum in having low protuberances on
the carapace, while others are very similar to Calman’s description of C. horri-
dum, having long, slender spine-like processes. However, close comparison of
the structure and positioning of the protuberances in the present specimens

SOUTHERN AFRICAN CUMACEA: PART 3 187

shows them to be very similar to each other, although the magnitude of the
sculpturing differs considerably. The sculpturing—in the form of sharp digiti-
form processes in some cases and merely raised bumps in others—is very similar
in arrangement to that figured by Calman for C. horridum. The structure of
the limbs is almost identical in both South African forms and in the description
of C. horridum, confirming that they are very similar. It therefore seems appro-
priate to consider these specimens as belonging to C. horridum.

Even in those specimens in which the carapace most closely approaches
that figured by Calman, the sculpturing is greatly reduced anteriorly and the
dactyl of pereiopod | inserts terminally on the propodus. In those specimens
with all the sculpturing reduced, the dactyl inserts subterminally, as it does in
Calman’s figure. Since the central and northern Atlantic form of the species
seems to be quite uniform, it is proposed to distinguish the local specimens
subspecifically as C. horridum australe. Due to the considerable variation
within as well as between samples, even the subspecies must be considered at
least to be highly polymorphic. Whether this polymorphism will prove sufficient
for erecting two species or subspecies in the place of C. horridum australe will
have to await the collection of a much larger number and wider range of indi-
viduals. At such time, it should also be possible to determine the relationship
between the local subspecies and C. horridum sensu Calman. -

Distribution
Natal from 680 to 900 m.

DISTRIBUTION OF CERATOCUMATIDAE

With the family consisting of only seven species in two genera, it is not
possible to draw many inferences from its distribution. It is worth noting,
however, that all seven species appear to be confined to the Atlantic and south-
western Indian Ocean (including Kerguelen). They are cold- and deep-water
forms, the depth at which they are found being reciprocal to the latitude,
except in records from the deepest waters.

Anatomically, Ceratocuma appears to form a cline of species differing
mainly in the sculpturing of the carapace but its distribution in the Atlantic is
rather haphazard and available evidence shows no relationship between mor-
phology and geographical distribution.

ACKNOWLEDGEMENTS

I wish to thank the South African Museum and Dr Brian Kensley for
providing me with the Pieter Faure material, and for offering me the oppor-
tunity to examine and describe the material collected by the Museum aboard
the R.V. Meiring Naude. I should also like to thank Professor John Day for

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

allowing me free access to all the material collected by the University of Cape
Town under his direction. I am particularly grateful to Dr N. S. Jones for his
very helpful criticism of this paper and other papers and for providing valuable
unpublished distribution data.

REFERENCES

BAcescu, M. 1969. Deux Cumacés nouveaux: Diastyloides carpinei n. sp. dans la Méditer-
ranée et Hemilamprops lotusae n. sp. dans l’ Atlantique Argentin. Rev. Roum. Biol. [Bucha-
rest] 14: 163-171.

BAcescu, M. 1972. Archaeocuma and Schizocuma, new genera of Cumacea from the American
tropical waters. Rev. Roum. Biol. [Bucharest] 17: 241-250.

B&cescu, M. & MurRADIAN, Z. 1974. New Cumacea from the north-western Atlantic: Cerato-
cuma panamensis ni. sp., Cimmerius costlowi n. sp. and some comments upon Petalosarsia
declivis (G. O. Sars). Rev. Roum. Biol. [Bucharest] 19: 217-227.

BAcescu, M. & MURADIAN, Z. 1976. Bathylamprops motasi sp. n. from the west Atlantic and
some considerations of the genus. Comun. Muz. Stiint. nat. Bdcau. 1976: 15-19.

BonNIER, J. 1896. Résultat Scientifique de la Campagne du ‘“‘Caudan’’ dans le Golfe de Gas-
cogne. III. Edriophthalmes. Annls. Univ. Lyon. 26: 529-562.

CALMAN, W. T. 1904. The marine fauna of the west coast of Ireland. Part IV. Cumacea.

Scient. Invest. Fish Brch. Ire. 1: 1-52.

CALMAN, W. T. 1905. The Cumacea of the Siboga Expedition. Siboga Exped. Monograph 36:
1-23.

CALMAN, W. T. 1912. The Crustacea of the order Cumacea in the Collection of the U.S.
National Museum. Proc. U.S. natn. Mus. 41: 603-676.

Day, J. 1975. South African Cumacea. Part 1. Family Bodotriidae, subfamily Vaun-
thompsoniinae. Ann. S. Afr. Mus. 66: 177-220.

Day, J. 1978. Southern African Cumacea. Part 2. Family Bodotriidae, subfamily Bodo-
triinae. Ann. S. Afr. Mus. 75: 159-290.

Face, L. 1928. Cumacés et Leptostracés provenant des campagnes du Prince Albert 1°* de
Monaco. Résult. Camp. scient. Prince Albert 1. 77: 1-55.

Gamo, S. 1967. Studies on the Cumacea (Crustacea, Malacostraca) of Japan. Part 2. Publs.
Seto mar. biol. Lab. 15: 245-274.

GIVEN, R. R. 1964. The cumacean fauna of the southern Californian continental shelf. No. 2.
The new family Mesolampropidae. Crustaceana 7: 284-292.

Hae, H. M. 1937. Cumacea and Nebaliacea. Rep. B.A.N.Z. Antarctic Res. Exped. 4 (2):
38-56.

Hate, H. M. 1946. Australian Cumacea. No. 13. The family Lampropidae. Trans. R. Soc. S.
Aust. 70: 178-188.

HANSEN, H. J. 1920. Crustacea Malacostraca. IV. VI. The Order Cumacea. Dan. Ingolf
Exped. 3B: 1-86.

Jones, N. S. 1963. The marine fauna of New Zealand: Crustacea of the Order Cumacea.
Bull. N.Z. Dep. scient. ind. Res. 152: 8-80.

Jones, N. S. 1969. The systematics and distribution of Cumacea from depths exceeding
200 metres. Galathea Rep. 10: 99-180.

Jones, N. S. 1971. The Fauna of the Ross Sea. Part 8. Cumacea. Bull. N.Z. Dep. scient. ind.
Res. 206: 33-41.

JonEs, N. S. 1973. Some new Cumacea from deep water in the Atlantic. Crustaceana 25:
297-319.

LEDOYER, M. 1977. Cumacés (Crustacea) des Iles Kerguelen recueillis par le N.O. “La Japo-
naise”’ en 1972 et 1974 et par le M.S. “‘Marion- Dufresne” en 1974. C.N.F.R.A. 42: 193-213.

NorMan, A. M. 1879. Crustacea Cumacea of the ‘‘Lightning’’, ‘“Porcupine” and ‘“Valorous”
Expeditions. Ann. Mag. nat. Hist. (5) 3: 54-73.

Sars, G. O. 1863. Beretning om en i Sommeren 1862 foretagen zoologisk Reise i Christianias
og Throndhjems Sifter. Nyt Mag. Naturvid. 12: 193-252.

Sars, G. O. 1869. Nye Dybvandscrustaceer fra Lofoten. Forh. VidenskSelsk. Krist. 1869:
147-174.

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Sars, G. O. 1878. Middelhavets Cumaceer. Part 1. Arch. Math. Natur. 3: 461-512.

Sars, G. O. 1883. Oversigt af Norges Crustaceer med forelgbige Bemaerkninger over de nye
eller mindre bekjende Arter. 1. Forh. VidenskSelsk. Krist. 1882: 1-124.

Sars, G. O. 1887. Report on the Cumacea collected by H.M.S. Challenger during the years
1873-1876. Rep. scient. Results Voy. Challenger. Zoology. 13 (37): 1-73.

STEBBING, T. R. R. 1912. South African Crustacea. Part 6. The Sympoda. Ann. S. Afr. Mus. 10:
129-176.

STEBBING, T. R. R. 1913. Cumacea. Tierreich 39: 1-210.

ZIMMER, C. 1907. Neue Cumaceen von der Deutschen und der Swedischen SiidpolarExpedi-
tion aus der Familien der Cumiden, Vaunthomsoniiden, Nannastaciden und Lampropiden.
Zool. Anz. 31: 367-374.

ZIMMER, C. 1908. Die Cumaceen der ,,Deutschen TiefseeExpedition’”. Wiss. Ergebn. dt.
Tiefsee-Exped. ‘Valdivia’ 8: 155-196.

ZIMMER, C. 1913. Die Cumaceen der Deutschen SiidpolarExpedition. Dt. Sudpol.-Exped.
1901-1903. 14, Zool. 6: 437-491.

ZIMMER, C. 1921. Einige neue und wenige bekannte Cumaceen des Swedischen Reichsmuseums.
Ark. Zool. Stockholm 13 (21): 1-9.

an Toe

2 hee

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Palaiype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

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

,

ee. 2. . 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 but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

JENNIFER DAY

SOUTHERN AFRICAN CUMACEA
PART 3

FAMILIES LAMPROPIDAE
AND CERATOCUMATIDAE

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ve LUME 76 PART 4 AUGUST 1978 ISSN 0303-2515

507.6%

YF THE SOUTH AFRICAN
MUSEUM

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

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names of joint authors connected by ampersand
et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).

Examples (note capitalization and punctuation)

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

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100—140.

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.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Koun, A. J. 19606. 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.

(continued inside back cover)

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

Volume 76 Band
August 1978 Augustus
Part 4 Deel

THREE NEW SPECIES OF HARPACTICOIDA
(CRUSTACEA, COPEPODA) FROM SANDY
BEACHES IN ALGOA BAY, SOUTH AFRICA, WITH
ies 10 THE GENERA ARENOSETELLA,
HASTIGERELLA, LEPTASTACUS AND
PSAMMASTACUS

By

ANTON McLACHLAN
&
COLIN G. MOORE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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

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van stof

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

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The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THREE NEW SPECIES OF HARPACTICOIDA (CRUSTACEA, COPE-

PODA) FROM SANDY BEACHES IN ALGOA BAY, SOUTH AFRICA,

mete KEYS TO THE GENERA ARENOSETELLA, HASTIGERELLA,
LEPTASTACUS AND PSAMMASTACUS

By
ANTON MCLACHLAN

Department of Marine Biology, University of Liverpool, Port Erin,
Isle of Man, and Zoology Department, University of Port Elizabeth,
South Africa

&
COLIN G. MOORE

Department of Marine Biology, University of Liverpool, Port Erin,
Isle of Man

(With 7 figures)

[MS. accepted 7 June 1978]

ABSTRACT

A new species of harpacticoid copepod of the family Ectinosomatidae, Arenosetella
fimbriaticauda sp. nov., is described from Algoa Bay, South Africa. It appears related to A.
duriensis Galhano and A. littoralis Bodin from which it differs in the setation of the endopo-
dites of the walking legs. It is proposed that Hastigerella palpilabra Nicholls and A. monensis
Moore are synonymous with H. tenuissima (Klie) which is transferred to the genus Arenosetella.
Two new species of the family Cylindropsyllidae are also described. Leptastacus naylori sp.
nov. differs from all described species of Leptastacus in the setation of the walking legs, while
Psammastacus erasmusi sp. nov. appears closely related to P. ghanai (Chappuis & Rouch)
from which it differs in the setation of the fourth leg and the structure of the fifth leg and
furcal rami. Keys to all these genera are provided.

CONTENTS

PAGE

Part 1
Introduction .. NER rege ct eo cn DO) aL OD
Systematic description Patmos. slow Fare 192
Discussion. . ae oe ees eee 197
Key to vereseie Wilson . ees eo ee ee ee lee OS
Key tovlasticerella Nicholls... =. 4} ve 2 199

Part 2
Introduction .. SEY SENT ASE ke WOO)
Systematic descriptions faa en Rhee aes eS)
Discussion. . ast 208
Key to the females of Leptostaeus T. Scott to DOS
Key to Psammasiacus Nicholls... . <2 4. 209
ANcknowiedsements:.< 0.) When. mele atis. oak ee) ) 209
INC ICRCMCE Seam fre esc) te, Ais held Rete eh ie 8 DOO

19]

Ann. S. Afr. Mus. 76 (4), 1978: 191-211, 7 figs.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

PART 1
INTRODUCTION

During investigations of the meiofauna of sandy beaches in Algoa Bay,
South Africa, specimens were collected of an interstitial harpacticoid of the
family Ectonisomatidae Sars, 1903, genus Arenosetella Wilson, 1932, thought
to be new to science. This species, referred to as Hastigerella sp. A by McLachlan
& Furstenberg (1977), is abundant around the high tide level on King’s Beach
and has also been recorded on Sunday’s River Beach, Algoa Bay. The physical
characteristics of these exposed sandy beaches have been described by McLachlan
(1977):

SYSTEMATIC DESCRIPTION

Family Ectinosomatidae Sars
Arenosetella fimbriaticauda sp. nov.

Material

A number of specimens were extracted from fine sand collected at the
spring high tide level ori King’s Beach (25°39’E 33°57’S) and preserved in 5 per
cent buffered formalin. For examination adult male and female specimens
were dissected in lactic acid and mounted in polyvinyl lactophenol.

Holotype
1 dissected 9 (SAM-A15708) deposited with the South African Museum
Cape Town, South Africa.

Allotype
1 adult ¢ (SAM-—A15712) in the South African Museum.

Paratypes
2 3dg (SAM-A15710 and 15711) and 1 2 (SAM-A15709) in the South

African Museum.
Other specimens are in the first author’s collection.

Description of adult female

Length 0,30-0,32 mm from base of rostrum to base of furcae. Body (cf
Fig. 1A) vermiform, cylindrical, with subtriangular pointed rostrum. Cephalo-
thoracic shield rectangular in dorsal view, with striations and bearing fine
hairs anteriorly. Cephalothorax about half as long as four free thoracic somites
combined; cephalothorax and thorax together slightly longer than abdomen.
Hyaline frill striated; cephalothorax and first two free thoracic somites fully-
incised obtusidigitate becoming acutidigitate (Moore 1976a) posteriorly;
penultimate somite (Fig. 1C—D) semi-incised subulate with small pseudoper-
culum dorsally and similar lobe ventrally; remaining somites deeply-incised
subulate (Fig. 1B). Cuticle of anal somite drawn out into rows of weak spinuli-

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 193

Fig. 1. Arenosetella fimbriaticauda sp. nov.

A. 6 dorsal. B. 2 abdominal hyaline frill.

C. 2 furcae dorsal.

lOO 7am

D. 2 furcae ventral.

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

form lappets; anterior row present dorsally and ventrally and apparently con-
tinuous inside median somitic cleft; posterior row of larger lappets curving from
the inner surface of the cleft onto the dorsal surface. Posterior margin of anal
somite with row of minute spinules. Genital double somite without obvious
signs of subdivision.

Furcal ramus (Fig 1C-D). Striated, slightly wider than long and tapering
distally (the specimen from which the drawings were made was slightly flattened).
Two principal terminal setae, inner one about as long as abdomen and free
thoracic somites combined and twice as long as outer one; accompanied to the
outside by one long and one short fine seta and to the inside by one short and
one longer, distally-plumose, fine seta. Distal margin of furca drawn out into
one dorsal and one ventral triangular lappet.

Antennule (Fig. 2A). Five-segmented, but segment five may possibly be
composed of a short proximal and a longer distal component. First segment
with a large plumose seta at anterior distal corner and a transverse row of fine
spinules on anterior surface. Third and terminal segments each supplied with
one aesthetasc.

Antenna (Fig. 2B). Exopodite three-segmented; first segment bearing one
terminal seta; second segment shortest and with one terminal seta; distal segment
with two terminal setae and a transverse row of spinules. Endopodite two-
segmented; first segment longer and slightly curved; second segment with two
transverse rows of spinules, anterior margin with some spinules near proximal
corner and two juxtaposed, spinulose setae; distal margin with six spinulose
spiniform setae.

Mandible (Fig. 2C). Coxa-basis with three setae near anterior distal corner.
Exopodite furnished with one lateral and two terminal setae. Endopodite same
length as coxa-basis and armed with three outer, three distal and two inner setae.

Maxillula (Fig. 2D). Arthrite of praecosa armed with one slender and four
strong unguiform spines. Basis with four setae, endopodite with four setae,
exopodite with two setae.

Maxilla (Fig. 2E). Syncoxa with two endites, each with two setae. Basis
with three setae near proximal inner edge. Endopodite with two strong genicu-
late setae and four slender setae.

Maxillipede (Fig. 2F). First endopodite segment longer than second and
with a row of long hairs along inner edge. Second segment with two fine setae
on inner edge, the proximal one longer, and two fine apical setae of different
lengths.

Leg 1 (Fig. 3A). Coxa and basis each with outer distal spinule row and
basis with strong inner seta. Rami three-segmented, spinulose along outer
edges. Exopodite extending to middle of distal endopodite segment; second
segment with inner seta modified with branched tip. Endopodite with median
and distal transverse rows of spinules on first segment and short distal row on
second segment.

Legs 2-4 (Fig. 3B-D). Coxa and basis each with outer distal spinule row.

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 195

SOM

Fig. 2. Arenosetella fimbriaticauda sp. nov. 2
A. Antennule. B. Antenna. C. Mandible. D. Maxillula. E. Maxilla. F. Maxillipede.

A. 9 Pi. B. 2 P2.. C. 9 Ps. D. 9 PA. Boo PS. FE. .g antenn

A
~\)
i \ /
Vy Ee NA NY
g V x ‘ Y
) , ()
: AA, /
om ca
. :
‘ |
==
E SOum
——

Fig. 3. Arenosetella fimbriaticauda sp. no

Vv.
ule. G.@ PS, Hie Ps:

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 197

Rami three-segmented, more or less spinulose along outer margins. Exopodite
extending past second endopodite segment; middle segment with modified
inner seta, such a seta also being present on the inner margin of the distal
segment of the P4. Endopodite with transverse row of spinules on first segment;
second segment bears, in addition to the inner seta, a short, thick, weakly-
chitinized seta on the posterior surface just inside the inner margin.

The setal formula is:

Exopodite Endopodite
| 2 3 1 2 3
Se 0 1 [oe aes 1 1 27 All
Sees. . | 1 eae. 2 ] 2, bet
P5 it 1 ee 2 1 2 lee ao
P4 1 ] 2 ae! ] 2 [e

Leg 5 (Fig. 3E). Rami indistinctly defined. Accessory seta issuing near
base of limb. Exopodite with three marginal setae; outer seta issuing from small
lobule, middle seta longest. Inner expansion of basoendopodite extending almost
to end of exopodite; inner seta more than twice as long as outer seta.

Description of adult male

Length 0,32—0,34 mm. Agrees with female apart from the following features.

Antennule (Fig. 3F). Eight-segmented; first segment bearing stout plumose
seta at anterior distal corner; second segment very short; fifth segment longest
and furnished with a large aesthetasc; terminal segment with long, slender
aesthetasc.

Leg 5 (Fig. 3G). The structure could not be definitively interpreted. All
setae subequal. Inner part of basoendopodite apparently distinct, bearing two
distal setae and extending to about the end of the expodite. Exopodite with
three marginal setae and no sign of an accessory seta.

Leg 6 (Fig. 3H). Bearing two inner spines and two outer setae.

Variability
Only slight variation in size was noted.

Etymology

The specific name alludes to the presence of the rows of fringing lappets
on the anal somite.

DISCUSSION

Arenosetella fimbriaticauda shows affinities with A. duriensis (Galhano
1970) and A. littoralis (Bodin 1978) from which it differs by the setation of the
distal endopodite segments of P2—P4. These three species, with their anal rows
of spinules or lappets, and A. balakrishnani (Bozic 1966) with four dorsal pairs
of spines on the anal somite, differ from most other members of the genus

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

which have claw-like structures. Lang (1965) distinguished Arenosetella from
Hastigerella solely by the presence of these anal claws. Wells (1976), however,
broadened the definition to include under Arenosetella forms with setae or
spines (as well as claws) on the dorsal surface of the anal somite as opposed to
the naked state in Hastigerella.

Whether all these anal structures are homologous is open to question, as 1s
their derivation. Chappuis (1953) believed that the anal claws are homologous
to the anal operculum present in most other harpacticoids. Thus, the last
copepodite stage of Arenosetella incerta Chappuis, 1953, possesses an oper-
culum armed with long spines, which in the adult are replaced by the claws.
The ornamentation of A. fimbriaticauda, however, suggests an alternative
origin: the position of the inner posterior row of lappets in A. fimbriaticauda
is occupied by a row of hairs or spinules in many harpacticoid species in several
families (e.g. Pseudobradya pulchera Lang, 1965; Arenolaophonte stygia Lang,
1965). Arenosetella fimbriaticauda may well represent an intermediate stage
in the development of this row, which is further modified into the laminar spines
of A. kaiseri Lang, 1965, and eventually into the unguiform claws of A. germanica
Kunz, 1938, and several other species.

The anal ornamentation was examined in several species of Hastigerella,
viz. H. palpilabra Nicholls, 1935, from the type locality (Millport, Scotland),
H. tenuissima (Klie, 1929) and H. leptoderma (Klie, 1929) from the Island of
Sylt, Germany. Careful examination, using phase contrast microscopy, failed
to reveal any dorsal armature of the anal somite in H. leptoderma; however,
both H. palpilabra and H. tenuissima were found to exhibit a pair of bifid
appendages identical to those of A. monensis Moore, 1976b: These are markedly
weaker than those found in A. germanica (see Moore 19766) and are conse-
quently easily overlooked. Further examination showed these three species to
be identical, the apparent setational differences of the pereiopods recorded in
the literature being due to variance of interpretation. A. incerta sensu Bodin,
1971, also appears conspecific. The senior synonym, H. tenuissima must,
therefore, be placed in the genus Arenosetella and it is suggested that H. lepto-
derma replaces H. palpilabra as the type-species of Hastigerella.

The following are a key to Arenosetella updated from Moore (1976b) and
a key to Hastigerella based partly on the keys of Lang (1965), Apostolov (1974)
and Wells (1976).

KEY TO ARENOSETELLA WILSON

1. Last somite dorsally with two simple appendages . . 2
— Last somite dorsally with two bifid appendages, each appendage furnished ‘with an
accessory spine . ey antes 3

— Last somite dorsally with two irifid appendaces Pag S06. at eae Noode, 1958
— Last somite dorsally either with two bifid appendages without accessory spines or with
two overlapping spines on each side of middle line > 92) 2))-2) ee aetna

— Last somite dorsally with four pairs of spines . . . . .balakrishnani Bozic, 1966

— Last somite with two rows of spinuliform lappets or spinules entirely or partly on the
dorsal‘sutfacé= : 2 s0 Se wo we ke)

=

ee eels Neal

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 199

Appendages not fused at base; last segment exopodite P4 with 2 inner setae
spinicauda Wilson, 1932
Appendages fused at base; last segment exopodite P4 without inner setae
indica Krishnaswamy, 1957
Accessory spine situated on outer side above bifurcation; last segment endopodite P4

with 4 setae and spinesinall . . . . . fissilis Wilson, 1932
Accessory spine situated on inner side ona evel with bitureation: last segment endopodite
P4 with 3 setae and spines inall . = incerta‘ Chappuis, 1953

Last segment endopodite P3 and P4 sadn “fh 5 setae and spines in all
madagascariensis Lang, 1965

Last segment endopodite P3 and P4 each with 4 setae and spines Mcae. Coon ee 3S
Last segment endopodite P3 and P4 each with 3 setae and spines inall . . if
Middle segment endopodite P4 with | seta . . . . . . . bidentata Ito, 1972
Middle seement endopodite P4 with 2 setae . . . . .. . 6
Appendages of last somite bifid, unguiform . . . . . . cerertan Kunz, 1938
Appendages of last somite straight. 2 2}. ‘kharser waneel9o5
First segment exopodite P2 and P3 each vit 1 inner r seta ~~ s = = rouchiLangs 1965
First segment exopodite P2 and P3 each with no inner seta. . ftenuissima (Klie, 1929)
Last segment endopodite P2—P4 with 4 setae and spines in all. fimbriaticauda sp. nov.
Last segment endopodite P2-P4 with 5 setae and spines inall. . . . . . . 9
Middle segment exopodite P2—P4 with | inner seta . . . . duriensis Galhano, 1970
Middle segment exopodite P2-P4 with 2 inner setae . . ._ . JUittoralis Bodin, 1978

KEY TO HASTIGERELLA NICHOLLS

Last segment exopodite P3 with 8 setae and spines inall . . . soyeri Bodin, 1976
Masmsccrinent.exopodite P3 with 7 setae and spmesinall .-. . . . . 4" 2
iastscement exopodite P3 with 6 setae and spmesinall . .. . . .°..5 +3
Last segment exopodite P3 with 5 setae and spines inall . . Ni a Gil eae Ay

Last segment exopodite P2—P4 with 7-7-7 setae and spines in Ait
meridionalis (Chappuis, 1954)

Last segment exopodite P2—P4 with 6-7-7 setae and spines in all .unisetosa (Wells, 1965)
Last segment exopodite P2—P4 with 7-6-6 setae and spines in all . chappuisi Soyer, 1974

Last segment endopodite P2-P4 with 4 setae and spines inall. . . . . . . 4
Last segment endopodite P2—P4 with 5 setae and spines inall. . . . . . . #5
Last segment endopodite P2—P4 with 6 setae and spines in all. . scheibeli Mielke, 1975
Al with 6 segments; penultimate abdominal somite with posterior ventral spinule
row ' ~ (2) 4. ) abbot: Lange. 1965
Al with 7 Posen: Be auliniate shdowinal somite with no posterior ventral spinule
row : : S).-bozier Soyer, 1974
Middle —— sndanedite P2-P4 ith 2 inner ieihe 3 neodts (Rao & Ganapati, 1969)
Muddictsesment endopodite P2—P4 with Linnerseta . 9. . .°>. . =. « .«. 6
hurcaonger than broad ~~... .. . :. . . .». . . bodini Apostolov, 1974
Furca broader thanlong . . . . . . .« psammae (Noodt, 1955)
Middle segment endopodite P1 with 2 inner sche oa ee et Pe Oe oe ee 8
Middle segment endopodite P! with | inner seta pes: AL OS Se nD

8. Last segment endopodite P1—P4 with 3 setae and spines in alll

clavata (Rao & Ganapati, 1969)

Last segment endopodite P1—P4 with 5 setae and spines in all . Jeptoderma (Klie, 1929)
Last segment exopodite Pl with 6 setae and spines in all

monniotae (Guille & Soyer, 1966)

Last segment exopodite PI with 5 setae and spines inall . . . Nass Perel |G)

Last segment exopodite Pl with 4 setae and spines inall . . Banwelenss Rao, 1972

Middle segment endopodite P2—P4 with | inner seta. . setosa (Rao & Ganapati, 1969)

Middle segment endopodite P2—P4 with 2 inner setae . grandimandibularis Wells, 1967

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

PARW 2
INTRODUCTION

Two new species of harpacticoid copepod of the family Cylindropsyllidae
Sars, Lang, thought to be new to science, have been collected between mean and
low tide levels on a high energy sandy beach in Algoa Bay. A species of
Leptastacus T Scott is common around the mean tide level and a species of
Psammastacus Nicholls is abundant around the spring low tide level (McLachlan
& Furstenberg 1977).

SYSTEMATIC DESCRIPTIONS
Family Cylindropsyllidae Sars, Lang
Leptastacus naylori sp. nov.
Material

Specimens were extracted from fine sand (median particle diameter
240 »m) on Sunday’s River beach (25°53’ E 33°43’S) and preserved in buffered
formalin. Six adults were examined, two being dissected in lactic acid and
mounted in polyvinyl lactophenol.

Holotype

1 9 (SAM-A15713) deposited in the South African Museum Cape Town,
South Africa.

Allotype m
1 adult § (SAM-A15716) in the South African Museum.
Paratypes
1 9(SAM-A15714) and 1 ¢ (SAM-A15715) in the South African Museum.

Description of adult female

Length 0,32-0,36 mm from base of rostum to base of furcae. Body (Fig. 4A)
vermiform, cylindrical with a short rostrum. Cephalothorax rectangular in
dorsal view. Genital double-somite without obvious signs of subdivision. Anal
operculum a rectangular plate with spinulose posterior margin.

Somitic ornamentation. Somites with a fine fully-incised subulate hyaline
frill. Anal somite with ventral rows of spinules along posterior edge.

Furcal ramus (Figs 4B, 41). Two times as long as broad and with an inner
row of ventrally-directed spinules; two terminal setae, inner much longer than
outer; one inner seta and two dorsal setae; distal end with a ventrally-directed
spine.

Antennule (Fig. 4C). Seven-segmented, with an aesthetasc on segments
four and seven. Segment two the longest and segments five and six the shortest.

Antenna (Fig. 4D). Coxa and allobasis devoid of ornamentation. Exopodite

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 201

Fig. 4. Leptastacus naylori sp. nov. 2

A. Habitus. B. Furcal rami. C. Antennule. D. Antenna. E. Labrum. F. Mandible.
G. Maxillula. H. Maxilla. I. Maxillipede.

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

one-segmented, with two small apical setae. Endopodite furnished with two
transverse rows of spinules on surface, two juxtaposed spines on anterior edge
and distal edge with two plain setae and three geniculate setae, the most
posterior sharing its base with a setule.

Mandible (Fig. 4F). Praecoxa with unidentate pars incisiva and a number
of small teeth along cutting edge. Palp two-segmented, segment one with one
spinule and segment two with one terminal and two lateral setae.

Maxillula (Fig. 4G). Arthrite of praecoxa with one claw and five spines.
Coxa with two apical setae. Basis with three terminal and two lateral setae.

Maxilla (Fig. 4H). Syncoxa with two endites, proximal one with two setae,
distal one with an unguiform spine and two setae. Basis with a large unguiform
spine. Endopodite two-segmented; first segment with one seta, second segment
with two apical setae.

Maxillipede (Fig. 41). First endopodite segment elongate; second segment
with a long, slender, pennate claw and a slender seta.

Leg | (Fig. 5A). Exopodite three-segmented, shorter than endopodite.
Endopodite of two subequal segments.

Legs 2-4 (Fig. 5B—D). Exopodite three-segmented, spinulose along outer
margin of first two segments. Endopodite of two subequal segments, shorter
than exopodite and spinulose along outer margin.

The setal formula is:

Exopodite Endopodite

l 2 3 l re
Pl 0 0 Ose ] Ociae
P2 0 0 O20 | O- 120:
P3 0 0 iD ale: | OMe
P4 0 | Dee 0 OF 0:

Leg (Fig. 5E). A triangular plate with one outer seta and spinule (or
setule), two distal setae and some spinules along inner margin.

Description of adult male

Length 0,28-0,33 mm. Agrees with female apart from the following
features. First two abdominal somites distinct.

Antennule (Fig. 5F). Haplocerate, eight-segmented. A very large aesthetasc
on segment four and a small one on segment eight. Segment two the longest
and segment five the shortest.

Leg 3. Second endopodite short and with one terminal seta.

Leg 5 (Fig. 5G). A plate with two marginal setae and a spinuliform
projection.

Leg 6 (Fig. 5H). A plate with two setae.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

This species is named in honour of Prof. Ernest Naylor of the Department
of Marine Biology, University of Liverpool, Isle of Man.

Psammastacus erasmusi sp. nov.

Material

Numerous specimens collected from medium sand (median particle diameter
260 ~m) at low tide level on Sunday’s River beach, preserved in 5 per cent
formalin and mounted in polyvinyl lactophenol.

Holotype
1 2 (SAM-A15717) in the South African Museum.

Allotype
| § (SAM-A15718) in the South African Museum.

Paratypes
7 92 (SAM-A15719) in the South African Museum.

Description of adult female

Length 0,38-0,49 mm from base of rostrum to base of furcae. Body (Fig.
6A) vermiform, cylindrical with an elongate pointed rostrum (Fig. 6B).
Cephalothorax rectangular in dorsal view. Genital double-somite without
obvious signs of subdivision. Anal operculum a rectangular plate with spinulose
posterior margin not easily visible. 7

Somitic ornamentation. Somites with a fine fully-incised subulate hyaline
frill and circumscribed by a row of rectangular thickenings on the abdomen.

Furcal ramus (Fig. 7A). 1,5 times as long as broad and with an inner row
of ventrally directed spinules; two terminal setae of unequal lengths; one inner
articulated seta, one middorsal seta and one outer lateral seta; distal end with
a strong, straight spine ventrally and a smaller, hooked spine dorsally.

Antennule (Fig. 6C). Seven-segmented with a large, annulated aesthetasc on
segment four. Segment two the longest and segment five the shortest.

Antenna (Fig. 6D). Coxa and allobasis devoid of ornamentation. Exopodite
one-segmented with two small apical setae. Endopodite furnished with two
transverse rows of spinules on surface, two outer juxtaposed spines and two
spines and three geniculate setae along distal edge; largest geniculate seta
confluent at base with a fine seta.

Mandible (Fig. 6F). Praecoxa with unidentate pars incisiva and a number
of small teeth along cutting edge. Palp two-segmented with one lateral and two
terminal setae on second segment.

Maxillula (Fig. 6G). Praecoxal arthrite with one seta and five spines.
Palp of one segment with two lateral and three terminal setae.

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 205

Fig. 6. Psammastacus erasmusi sp. nov.

A. 2 habitus. B. 2 rostrum. C. 9 antennule. D. 2 antenna. E. 2 labrum. F. 2 mandible.
G. ? maxillula. H. 2 maxilla. I. 2 maxillipede. J. J antennule.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

50 sum

Fig. 7. Psammastacus erasmusi sp. nov.
A. © furcal rami. B. 2 Pl. C.2 P2. D.¢ P3. E. 2 P4.. FB. 2 Pd. (Ga ese G4.
I, 6 25s Ja 26:

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 207

Maxilla (Fig. 6H). Syncoxa with two endites; smaller one with two small
setae; larger one with a seta and an unguiform spine. Basis with a terminal
unguiform spine. Endopodite with one lateral and two terminal fine setae.

Maxillipede (Fig. 61). First endopodite segment elongate; second segment
with a long, slender, pennate claw and a slender seta.

Leg | (Fig. 7B). Exopodite a single segment slightly longer than first
endopodite segment; with three terminal setae and one outer spine midway
along ramus. Endopodite of two subequal segments, the first with an inner
median spine and the second with two terminal setae.

Legs 2-4 (Fig. 7C—E). Rami more or less spinulose along outer margins.
Exopodite three segmented, with distal row of spinules; segments with a well-
developed appendicular hyaline frill. Endopodite a little shorter, two-segmented.
Fourth leg much the largest; inner distal spine of endopodite very stout and
partly confluent at base.

The setal formula is:

Exopodite Endopodite

i 2 3 1 2
Pl — — OR O22 0:
P2 0 0 On2e1 0 ORO)
235 a | 0 LeZell 0 Oni0%
Eee. .-.0 1 Dale 0 OR250:

Leg 5 (Fig. 7F). A small plate with four setae.

Description of adult male

Length 0,37-0,47 mm. Agrees with female apart from the following features.
First two abdominal somites distinct.

Antennule (Fig. 6J). Haplocerate, eight-segmented with an annulated
aesthetasc on segment four. Segment six with three chitinous thickenings along
anterior margin.

Leg 3 (Fig. 7G). As in female but without terminal seta on distal endopodite
segment.

Leg 4 (Fig. 7H). As in female but inner distal spine of endopodite of
different construction.

Leg 5 (Fig. 71). A small plate with 3 setae and a short spinuliform projection
at inner distal corner.

Leg 6 (Fig. 7J). A very small plate with two setae.

Etymology

This species is named in honour of Prof. Theunus Erasmus of the Zoology
Department, University of Port Elizabeth.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

DISCUSSION

Leptastacus naylori differs in the setation of the swimming legs from all the
described species of Leptastacus T. Scott. In this respect it is closest to L.
macronyx (T. Scott, 1892) and L. laticaudatus Nicholls, 1935, from which it
differs in the presence of an extra inner seta on the distal segment of the endopo-
dite of the third leg.

The following key to the females of the genus Leptastacus is based partly
on the key of Lang (1965) and the setal formula table of Lindgren (1975).
Here, because of inadequate descriptions, L. nichollsi Krishnaswamy, 1951,
L. acuticaudatus Krishnaswamy, 1957, L. euryhalinus Krishnaswamy, 1957, and
L. macronyx pontica Griga, 1962, have been omitted.

Apostolov (1973) gives drawings of a form from the Black Sea he ascribes
to L. laticaudatus Nicholls, 1935 intermedius Kunz, 1938. However, as the fifth
leg clearly differs from Kunz’s (1938) original description of this subspecies,
the Black Sea form is given in the key as L. laticaudatus intermedius sensu
Apostolovy.

KEY TO THE FEMALES OF LEPTASTACUS T. SCOTT

1. P2 endopodite with an inner seta on'segment-) 2 2 9.2) ee
— P2 endopodite without an inner seta on segment 1 2 92 7) ee LL
2. P3 endopodite with an mmner seta on seement 1 S57 =) eee eS
— P3 endopodite without an inner setaon segment! . . . . minutus Chappuis,1954
3. P4 endopodite with | seta on terminal segment . : raee a
=. P4 endopodite with 2 setae on tetmimal segment. ~~ 3 ieee
4. P3 endopodite with 1 seta on terminallsegment “: «2 0) *5 S)
— P3 endopodite with 2 setae on terminal segment. . . . . . . naylorisp. nov.
5. P5 foot-shaped at tip MPP r er GG lel lw
— PS not foot-shaped at tip . . » 2. a 2 % °s 5 6)
6. PS with2 inner setae. . . . . . . . . laticaudatus laticaudatus Nicholls; 1935
— P5with3innersetae. . . . . . . laticaudatus intermedius Kunz, 1938
7. P5 produced distally into short projection with rounded tip; furca about 3 times as long
as wide. . . . laticaudatus intermedius sensu Apostolov, 1973

— PS produced distally into long pointed projection; furca at least 4 times as long as
wide . . oo. 6 macronyx CE, Scott, 1892)

oy LES) produced distally into long pointed projection ee
— P5 not produced distally into long pointed projection . . . incurvatus Lang, 1965
9. P3 endopodite with 1 seta on terminal segment . . rostratus rostratus Nicholls, 1939
— P3 endopodite with 2 setae on terminal segment. . . rostratus taurica Marinov, 1973
10. Segment 2 exopodite P4 with no inner seta . . . . . . aberrans Chappuis, 1953
— Segment 2 exopodite P4 with 1 inner seta Pee
11. Terminal segment endopodite P4 with 2 setae and spines in all operculatus Masry 1970
— Terminal segment endopodite P4 with 1 seta. . : teen 12
12. Terminal segment endopodite P3 with 2 setae and spines in all Me eae
= ‘Terminal sesment endopodite P3 with lseta. | Ys) ee 13
(32° PS Wath: Setae.s, Ho, A) ce) Arye ee erat) eee PRIS Ais delamaret Rouch, 1962
— PSwith4setae . . a ena cePRtoiak Ny itoienes Rao & Ganapati, 1969
14. PS with no setae on inner ¢ margin erry (277) 02,77) Linclensi, ITs
— P5 with 1 seta on inner margin ce RS LE)
— P5with2setaeoninnermargin . ... . : . . . wieseri Chappuis, 1958
15. Principal furcal setae confluent at base =... ...2 0.2 <4 | (3 (anewesomers ne 1968

Principal furcal setae not confluent at basé . . . =. . . : IG

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY 209

16. Inner distal corner of furca produced into spiniform projection
mozambicus Wells, 1967
— Inner distal corner of furca not produced Se a PS oneonsinictus Lanesl965

The setation of legs one to four of Psammastacus erasmusi agrees only
with P. spinicaudatus Rao & Ganapati, 1969, and P. remanei Noodt, 1964.
However, the sexual dimorphism of the third and fourth legs shows the species
to be closest to P. ghanai Chappuis & Rouch, 1960. These two species differ in
the setation of the fourth leg and in the structure of the fifth leg and furcal rami.

The modification of the endopodite of the fourth leg of the female in
P. erasmusi is unique in the genus. Moreover, the annulated structure of the
antennular aesthetascs has not been observed by the authors in any other species
of harpacticoid copepod.

The following key to the genus is valid for both sexes.

KEY TO PSAMMASTACUS NICHOLLS

Pebniaescement expodite P4 with 2inner setae . . .-. 5 «© w= +» « « « 2
aihirdscament expodite P4 with 1 mnerseta. . . . «© « = » « « » « 6
2. Exopodite Pl with 3 setae and spinesinall . . . . perplexus Wells & Clark, 1965
Exe npoditer! with 4setaeandspinesinall .. . . .- . . «-» =m » « «» 3
3. Exopodite Pl with outer spine midway alongramus. . ........ 4
— Exopodite Pl with no such spine midway along ramus. . . spinicaudus Wells, 1967
4, Inner distal corner exopodite P4 prolonged into stout spine . . . erasmusi Sp. nov.
— Inner distal corner exopodite P4 with no such spine. 5
5. Fifth legs partly fused along midline . . . . spinicaudatus Rao & IGanapatt 1969
— Fifth legs distinct along midline . ... . . . . ramanei Noodt, 1964
memiopedie b4 with)! apicalseta . 4. 2 we 3 ew we es ew tC
SP AMOnMmC E4 With 2 apical Stace. 2 9. 6s mw ew ee ee 8
feeeuccarbroadet thanlone . . . . . . =. +. «+ . brevicaudata Nicholls, 1935
—wenorcaienserthan-broad . . . . « « « « « ¢« ~.,-€onfluens Nicholls, 1935
Seesavitder setae . .-. <« « . = » » » « @cuticaudatus Krishnaswamy, 1957
Sammiimitmaasetae . <«-. . « .« « »« « « « ghanai (Chappuis & Rouch, 1960)
ACKNOWLEDGEMENTS

We thank Dr W. Scheibel who originally identified to genus specimens of
all three species, Drs W. Mielke and I. Barclay for supplying specimens, and
Dr Ph. Bodin for critical reading of part of the initial manuscript. The first
author gratefully acknowledges financial assistance from the British Council,
the South African Council for Scientific and Industrial Research and the Univer-
sity of Port Elizabeth.

REFERENCES

AposToLoyv, A. 1973. Sur divers Harpacticoides (Copépodes) de la Mer Noire. Zool. Anz.
190: 88-110.

ApostoLov, A. 1974. Copépodes Harpacticoides de la Mer Noire. Trav. Mus. Hist. Nat.
‘Gr. Antipa’ 15: 131-139.

BopIn, P. 1971. Copépodes Harpacticoides marins des environs de La Rochelle 2. Espéces
de la zone intertidale d’Yves. Téthys 3: 411-433.

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

BopIN, P. 1976. Catalogue of new marine harpacticoid copepods: Supplement 2. Téthys 7:
265-278.

Bonin, P. 1978. Copépodes Harpacticoides marins des environs de La Rochelle 5.—Espéces
nouvelles ou incertaines. Vie Milieu (in press).

Bozic, B. 1966. Deux copépodes harpacticoides de l’Inde. Bull. Mus. natn. Hist. nat., Paris
38: 869-873.

CuaAppulis, P. A. 1953. Harpacticides psammiques récoltés par Cl. Delamare Deboutteville en
Méditerranée. Vie Milieu 4: 254-276.

CHApPuIsS, P. A. 1954. Copépodes psammiques des plages du Rousillon. Archs Zool. exp. gén.
91: 35-50.

CHAPPUIS, P. A. 1958. Harpacticoides psammiques marins des environs de Seattle (Washington,
USA). Vie Milieu 8: 409-422.

CHAPPUIS, P. A. & Roucn, R. 1960. Arenotopa ghanai n.g., n.sp. Harpacticoide psammique
des cdtes de l’Afrique. Bull. Inst. fr. Afr. noire 22A: 1248-1251.

GALHANO, M. H. 1970. Contribuicao paro o conhecimento da fauna intersticial em Portugal.
Publ. goes Inst. Zool. Dr. Auguste Nobre 110: 1-106.

GRIGA, R. E. 1962. Copepoda of the benthonic biocenoses in the region of Eupatoria of the
Black Sea. Trudy sevastopol’. biol. Sta. 15: 101-117.

GuILLE, A. & Soyer, J. 1966. Copépodes Harpacticoides de Banyuls-sur-Mer. 4. Quelques
formes des gravelles 4 Amphioxus. Vie Milieu 17: 345-387.

Ito, T. 1968. Descriptions and records of marine harpacticoid copepods from Hokkaido I.
J. Fac. Sci. Hokkaido Univ. ser 6, Zool. 16: 369-381.

Ito, T. 1972. Descriptions and records of marine harpacticoid copepods from Hokkaido, IV.
J. Fac. Sci. Hokkaido Univ. 6: 305-336.

Kure, W. 1929. Die Copepoda Harpacticoida der stidlichen und mittleren Ostsee mit beson-
derer Beriicksichtigung der Sandfauna der Kielier Bucht. Zool. Jb. Syst. 57: 329-386.
KRISHNASWAMY, S. 1951. Three new species of sand dwelling copepods from the Madras coast.

Ann. Mag. nat. Hist. ser 12 4: 273-280.

KRISHNASWAMY, S. 1957. Studies on the Copepoda of Madras. Unpublished Ph. D. thesis,
University of Madras, India.

Kunz, H. 1938. Zur Kenntnis der Harpacticoiden des Kustengrundwassers der Kieler Forde.
Kieler Meeresforsch. 2: 95-115.

LANG, K. 1965. Copepoda Harpacticoidea from the Californian Pacific Coast. K. svenska
Vetensk Akad. Handl. 10: 1-560.

LINDGREN, E. W. 1975. Six meiobenthic Harpacticoidea (Crustacea) from North Carolina
beaches. Cah. Biol. mar. 16: 445-473.

Marinov, T. 1973. Quelques harpacticides psammophiles inconnus pour le bassin de la mer
Noire. Vie Milieu 23: 309-326.

Masry, D. 1970. Ecological study of some sandy beaches along the Israeli Mediterranean
coast, with a description of the interstitial harpacticoids (Crustacea, Copepoda). Cah.
Biol. mar. 11: 229-258.

McLacu.an, A. 1977. Studies on the psammolittoral meiofauna of Algoa Bay. I. Physical
and chemical evaluation of the beaches. Zoologica afr. 12: 15-32.

McLACHLAN, A. & FURSTENBERG, J. P. 1977. Studies on the psammolittoral meiofauna of
Algoa Bay. III. A quantitative analysis of the nematode and crustacean communities.
Zoologica afr. 12: 61-71.

MIELKE, W. 1975. Systematik der Copepoda eines Sandstrandes der Nordseeinsel Sylt. Abh.
math, naturw.e KI. Akad. Wiss. Mainz 52: 1-134.

Moore, C. G. 1976a. The form and significance of the hyaline frill in harpacticoid copepod
taxonomy. J. nat. Hist. 10: 451-456.

Moore, C. G. 1976b. The harpacticoid families Ectinosomatidae and Diosaccidae (Crustacea,
Copepoda) from the Isle of Man. J. nat. Hist. 10: 131-155.

NICHOLLS, A. G. 1935. Copepods from the interstitial fauna of a sandy beach. J. mar. biol.
Ass. U.K. 20: 379-404.

NIcHOLLs, A. G. 1939. Marine harpacticoids and cyclopoids from the shores of the St.
Lawrence. Naturaliste can. 66: 241-316.

Noopt, W. 1955. Copepoda Harpacticoida von Teneriffa (Kanarische Inseln). Zool. Anz.
154: 200-202.

THREE NEW SPECIES OF HARPACTICOIDA FROM ALGOA BAY DI

Noopt, W. 1958. Die Copopoda Harpacticoidea des Brandungsstrandes von Teneriffa
(Kanarische Inseln). Abh. math.-naturw. Kl. Akad. Wiss. Mainz 2: 51-116.

Noopt, W. 1964. Copepoda Harpacticoidea aus dem Litoral des Roten Meeres. Kieler.
Meeresforsch. 20: 128-154.

Rao, G. C. 1972. Some new interstitial harpacticoid copepods from Andhra coast, India.
Cah. Biol. mar. 13: 305-319.

Rao, G. C. & GANAPATI, P. N. 1969. Some new interstitial copepods from Waltair Coast.
Proc. Indian Acad. Sci. 69B: 1-14.

Roucu, R. 1962. Harpacticoides (Crustacés Copépodes) d’Amérique du Sud. Jn: DELAMARE-
DEBOUTTEVILLE, CL. & RApPoporT, E., eds. Biologie Amérique Australe: 237-280. Paris:
Centre National de la Recherche Scientifique.

Scott, T. 1892. Additions to the fauna of the Firth of Forth. Rep. Fishery Bd Scotl. 3: 244-272.

Soyer, J. 1974. Contribution a lVetude des Copépodes Harpacticoides de Méditerranée
Occidentale. 9. Le genre Hastigerella Nicholls (Ectinosomidae Sars, Olofsson). Systé-
matique, écologie. Vie Milieu 24B: 175-192.

WELLS, J. B. J. 1965. Copepoda from the meiobenthos of some Scottish marine sublittoral
muds. Proc. R. Soc. Edinb. 69B: 1|-33.

WELLS, J. B. J. 1967. The littoral Copepoda (Crustacea) of Inhaca Island, Mozambique.
Trans. R. Soc. Edinb. 67: 189-358.

WELLS, J. B. J. 1976. Keys to aid in the identification of marine harpacticoid copepods. Aber-
deen: Dept. Zoology, Univ. Aberdeen.

WELLS, J. B. J. & CLARK, M. E. 1965. The interstitial Crustacea of two beaches in Portugal.
Reyta Biol., Lisb. 5: 87-108.

WILSON, C. B. 1932. The copepods of the Woods Hole region, Massachusetts. Bull. U.S.
natn. Mus. 158: 1-135.

6. SYSTEMATIC papers must conform to the J/nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *.. . the Figure depicting C. namacolus...’; *. . . in C. namacolus (Fig. 10)...’

(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 but F. von Huene

(c) Scientific names, but not their vernacular derivatives
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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
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‘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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

ANTON McLACHLAN
&
COLIN G. MOORE

THREE NEW SPECIES OF HARPACTICOIDA
(CRUSTACEA, COPEPODA) FROM SANDY BEACHES
IN ALGOA BAY, SOUTH AFRICA, WITH KEYS TO
THE GENERA ARENOSETELLA, HASTIGERELLA,
LEPTASTACUS AND PSAMMASTACUS

CAPE TOWN

|VOLUME 76 PART 5 SEPTEMBER 1978 ISSN 0303-2515

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

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. Ta: 7 634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. 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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September 1978 September
Part.) es “Deel

THE SKELETON OF THE MAMMAL-LIKE REPTILE
CmstECEPHALUS WITH: EVIDENCE FOR A
FOSSORIAL MODE OF LIFE

By

MICHAEL A. CLUVER

Cape Town Kaapstad

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THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS
WITH EVIDENCE FOR A FOSSORIAL MODE OF LIFE

By
MICHAEL A. CLUVER

South African Museum, Cape Town
(With 20 figures)

[MS. accepted 21 June 1978]

ABSTRACT

A full description of the skeleton of Cistecephalus, based on a number of specimens, is
given. A skeletal reconstruction shows that Cistecephalus was probably the most aberrant
member of the infraorder Dicynodontia, and comparison with living animals suggests that
the osteological modifications of, in particular, the shoulder girdle and forelimb represent
adaptations to powerful and frequent digging activities.

CONTENTS
PAGE
Introduction : é ; ‘ : : : : > 213
Material . ' : : : ; a oa
Skeleton of ieceohalas
Skull and lower jaw. : ike Teeth hs free AEDS
Vertebral column . F : ‘ ; : DNS
Ribs . é : ; : - 22h
Pectoral eile and forelimb : F ‘ : 28S)
Pelvic girdle and hind limb . ‘ 5 PvE
Range of movements and musculature of the faaee «.5| 236
Pectoral girdle and forelimb : ‘ ; ; «7236
Pelvic girdle and hind limb . : ; : 5 3h 2.239
Discussion . : ‘ : , : : : 3 2) |
Summary . : : ‘ 3 f : 5 . 244
Acknowledgements : ; : k : : : . 244
References . : : : 5 2 , 4 ; . 244
Abbreviations . ‘ ; 4 3 ‘ E , ww), 245
INTRODUCTION

The cranial morphology of the Upper Permian dicynodont Cistecephalus
has been made well known through a succession of papers following Owen’s
original description of the genus in 1876. Chief among these are those of Seeley
(1894), Broom (1932, 1948), Broili & Schréder (1935), Brink (1950, 1952),
Keyser (1973) and Cluver (1974). As details of the skull structure were estab-
lished, it became clear that Cistecephalus was a highly specialized animal
showing a number of fundamental departures from the usual dicynodont
condition. As these specializations must represent adaptations to a specific
way ot life, Cistecephalus has been the subject of some speculation by a number

213

Ann. S. Afr. Mus. 76 (5), 1978: 213-246, 20 figs.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

of authors. Aquatic habits were proposed by Broom (1948) and Brink (1950),
but the latter author later (Brink 1952) maintained that the structure of the
manus indicated adaptations to digging activities. Keyser (1973) thought that
the orientation of the orbits and possible opposability of digits of the manus
pointed to commitments to arboreality, but included digging activities as part
of the animal’s general way of life. Cluver (1974) compared the skull of Ciste-
cephalus with that of the related Kawingasaurus, for which genus Cox (1972)
had produced convincing evidence of powerful digging or fossorial habits,
and also with skulls in living and extinct fossorial mammal groups, and con-
cluded that Cistecephalus was in all likelihood an accomplished digger.

Since only tantalizingly little of the cistecephalid postcranial skeleton has
been described in the literature (Brink 1952, Keyser 1973, Von Huene 1942,
Cox 1972), it seemed worth while to investigate available material of the genus
in order to determine whether the peculiarities in the cranial structure are
matched by specializations in the axial and appendicular parts of the skeleton.
As a result of this exercise, it has been possible to establish a complete recon-
struction of the cistecephalid skeleton, which shows that the living animal was
probably the most specialized and aberrant member of the infraorder Dicyno-
dontia, in itself a highly modified division of the order Therapsida.

Studies on the postcranial skeleton of Permian dicynodonts are regrettably
few; in this investigation use was made chiefly of the accounts by Watson (1960),
Cox (1959, 1972) and Boonstra (1966).

MATERIAL

SAM-10665. Cistecephalus sp. Ou Plaas, Murraysburg, Cape Province. The
skull, complete but slightly compressed dorsoventrally, has been fully prepared.
The lower jaw, lacking the posterior part of the right ramus, has been separated
from the skull and also fully prepared.

BPI 4086. Cistecephalus sp. Bloukop, Roggevlei boundary between Richmond
and Victoria West, Cape Province. An almost complete skeleton, showing the
skull and lower jaw, the vertebral column and ribs, the right scapula, humerus,
radius and partial manus, a complete left hind limb and the right femur and
distal ends of the tibia and fibula. The atlas and last caudal vertebrae are absent.

BPI 696. Cistecephalus sp. Aasvogelkrans, Murraysburg, Cape Province. A skull
and lower jaw with associated but largely disarticulated postcranial skeleton.
The right scapula, left humerus ‘and ulna and partial left manus are well dis-
played, as well as the pelvic girdle, sacral and caudal vertebrae.

BPI 2915. Cistecephalus sp. Beeldhouersfontein, Murraysburg, Cape Province.
A skull and lower jaw with cervical vertebrae, pectoral girdle and complete
left forelimb and manus.

BPI 506. Cistecephalus sp. Towerwater, Murraysburg, Cape Province. A skull
and lower jaw with anterior vertebrae, a complete pectoral girdle and a good
left humerus.

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS DS

BPI 2450. Cistecephalus sp. Klipplaat, Murraysburg, Cape Province. A skull
and partial lower jaw with isolated ribs and the right half of the pelvic girdle.
BPI 4120. Cistecephalus sp. Modderfontein, Victoria West, Cape Province.
Skull and anterior part of vertebral column, pectoral girdle and articulated
forelimbs.

RC 298. Cistecephalus sp. Tweefontein, Graaff-Reinet, Cape Province. Good
skull and lower jaw with anterior part of vertebral column and ribs, a nearly
complete pectoral girdle and a good left humerus with the proximal part of the
ulna in articulation.

BPI 2124. Cistecephalus sp. Kraaifontein, Murraysburg. Skull and lower jaw
with pectoral girdle and right and left forelimbs in near-natural articulation.
Left manus seen in ventral view.

GS K224. Cistecephalus sp. Steilkranz, New Bethesda. Skull and lower jaw
with articulated anterior part of postcranial skeleton. Manus of left and right
sides seen in ventral view.

SAM-11114. Oudenodon sp. Melton Wold, Victoria West. Skull, lower jaw,
pectoral girdle and left and right forelimbs.
All specimens are from Upper Permian, Cistecephalus zone localities.

THE SKELETON OF CISTECEPHALUS

Skull and lower jaw

The Cistecephalus skull and lower jaw have been fully described by Keyser
(1973) and need not be considered in detail here. Some features of functional
significance may, however, be mentioned. As suggested by Cluver (1974), the
broad intertemporal region and rounded occiput may reflect forward extension
and hence increased size of the neck and shoulder musculature, such as is
found in many burrowing mammals. The basicranial region is strengthened by
a meeting in the ventral midline between the pterygoids, vomer and the base
of the presphenoid, with the obliteration of the interpterygoidal vacuity.

The lower jaw is short, deep and robust. The dentary symphysis is extended
as a Sharp transverse blade which met the premaxillary part of the secondary
palate during mastication. This scoop-like anterior edge of the lower jaw is
narrower than the secondary palate, and it is likely that the lower jaw could be
moved laterally across the secondary palate while crushing food in the mouth.
The short lower jaw did not meet the blunt anterior edge of the premaxilla at
any time during the masticatory cycle.

In Figure 20 skull details are derived from SAM-—10665 and BPI 4086.
Detailed drawings of cistecephalid skulls have been published by Keyser (1973)
and Cluver (1974).

Vertebral column :

In RC 298 the atlas, axis and succeeding 6 cervical vertebrae are preserved
in natural or near-natural articulation (Figs 1-2). The right half of the atlas
neural arch lies slightly displaced upon the odontoid process of the axis, and

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

t. proc.

Fig. 1. Cistecephalus sp. RC 298. A. Cervical vertebrae in right lateral view. B. Axis and
3rd cervical vertebra in ventral view. C. Atlas intercentrum in ventral view.

carries a strong anteroposteriorly elongated transverse process. The facet for
the proatlas is situated on a distinct raised platform. The body of the atlas arch
carries a large posteromedially facing facet that articulates with the odontoid
process, and an anteromedially facing facet that meets the occipital condyle.
An isolated atlas intercentrum is present. The element is transversely expanded
and carries a ventral longitudinal crest. Little can be seen of the articulatory
facets for the odontoid and occipital condyle, but it appears that the bone is
flatter than that of Lystrosaurus (Cluver 1971).

The axis is well preserved. The neural arch is indistinguishably fused to
the centrum, and bears a broad spine, higher than those of the succeeding
vertebrae. The postzygapophysis is tightly articulated with the prezygapophysis
of the next vertebra, but the prezygapophyseal region is damaged, evidently as
a result of the displacement of the atlas. The centrum of the axis carries a
distinct transverse process and is apparently fused with the odontoid element.
There is a smaller process, the parapophysis, low down on the centrum directly
beneath the transverse process, serving for the articulation of the capitulum of
a short but stout axial rib.

Six vertebrae are preserved in articulated position behind the axis, the
last two incompletely exposed. The three immediately behind the axis have
narrower spines than the fourth, and could represent true cervicals. Including
atlas and axis, this would mean a total of five cervical vertebrae in the column.
The third cervical carries a long, backwardly and dorsally directed transverse
process, much more prominent than those of the axis or the succeeding verte-
brae; the transverse processes of the last two cervicals are directed laterally and

217

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS

only slightly posteriorly. The zygapophyses of the cervicals, particularly the
last two, are set close to the neural spine and are tightly interlocked. In what is
regarded as the first dorsal vertebra, the transverse process is short and exca-
vated posteriorly, so that in lateral view the distal surface of the process is
arcuate in outline.

In BPI 4086 the vertebral column is complete except for the atlas and the
most posterior tail vertebrae. In this specimen, too, a total of five cervicals can
be recognized. The first three cervicals, including the axis, are articulated, but

bt

et

vililinlintilinlin

eI

Fig. 2. Cistecephalus sp. RC 298. Stereophotographs of anterior cervical
vertebrae. A. Right lateral view. B. Ventral view.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

the fourth and fifth in the row are loose from each other and from the sixth,
from which point the column is again fully articulated. The fourth in the row
(fifth cervical) is more complete than the anterior vertebrae; the spine appears
narrower than that of the succeeding vertebra, which differs, too, in the stouter
transverse process carried higher on the neural arch. This latter vertebra can
be regarded as the first dorsal (see p. 219). A total of twenty-five dorsals is
present. All are damaged to some extent, and the twenty-first and twenty-
second dorsals have become separated from each other. As far as can be made
out, the dorsals have broader neural spines than the cervicals, and in some
instances this width becomes extreme. The twenty-second dorsal has a very
broad anteroposteriorly inflated spine; the adjoining vertebrae are too damaged
to give any indication of the breadth of their spines. In all the vertebrae,
including the cervicals, the zygapophyses articulate at steep angles to the
horizontal.

Three sacral vertebrae are present in BPI 4086, each apparently indis-
tinguishably fused to its pair of ribs (Fig. 16A). The first sacral rib is very deep,
as is the second, but the third is less strongly developed. All three sacral vertebrae
are damaged and many details are lacking, but it seems that the spine of the
second slopes back to a much greater degree than does that of the last dorsal.

In BPI 696 the two halves of the pelvis are in near-natural position relative
to each other and sacral and caudal vertebrae are preserved. The first sacral
vertebra is partially exposed and its right rib is seen in anteroventral view. The
rib is short and wide distally and is not fused to the vertebra, which it meets
close to the body of the centrum. The centra of three vertebrae are visible in
ventral view behind the first sacral; the last of these is that of the first caudal.
In this specimen the remaining caudal vertebrae have folded over to lie upon
the dorsal spines of the sacral vertebrae, obscuring them from view. Only the
ventral surfaces of the centra, and in some cases the ribs, of the caudal vertebrae
are visible. A total of fifteen caudals is present, the last being a mere fleck of
bone and it seems likely that this represents a complete or near-complete
series, although it is possible that members of the series may be missing at the
point of dislocation from the sacral series.

In BPI 4086 four caudal vertebrae, all damaged and the last very incomplete,
are present. The second caudal has a fairly long laterally directed rib.

As far as can be seen, the zygapophyses of all vertebrae examined meet
each other at steep angles to the horizontal, and it is likely that only dorso-
ventral flexure or rotation between vertebrae could have been possible, even in
the neck region. No horizontal or near-horizontal articulations are present
anywhere in the column.

Ribs
The anterior ribs are best seen in RC 298. There is no indication of an

atlantal rib, but a short, stout and double-headed axis rib is present (Fig. 1).
The second rib present, that of the third cervical, is poorly preserved and it is

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 219

not possible to determine whether a double head was present; there is, how-
ever, no clear parapophysis for attachment of the capitulum, as in the axis. In
the fourth cervical a short and relatively slender double-headed rib is present.
In the fifth cervical the rib is incomplete but longer than that of the fourth. The
capitulum and tuberculum lie close to each other and are separated by a very
shallow notch. The next two ribs are almost complete, and lie crossed over each
other with the ventral ends meeting the side of the sternum in what appears
to be a natural association. The sixth vertebra carries the anterior of these
two ribs and can be regarded as the first dorsal, an identification supported by
characters of the vertebrae themselves (see p. 218). The head of the first dorsal
rib is incomplete, but that of the second dorsal is wide and a separate capitulum
and tuberculum cannot be made out. It seems likely (see below) that in this
specimen at least three ribs are attached to the sternum.

In BPI 4086 ribs are preserved from the first dorsal vertebra to the twenty-
fourth dorsal vertebra. Ribs are very short in the posterior region and are
longest between the sixth and eleventh dorsal, but it should be noted that none
of the ribs is absolutely complete. As shown above, the second caudal vertebra
carries a simple, laterally directed rib. In BPI 696 the first three caudals in the
dislocated portion of the row have short, slender ribs; there is no sign of haemal
arches in this series, and the base of the tail was probably clearly demarcated
from the rest of the body.

Pectoral girdle and forelimb

In RC 298 there is an almost complete left scapula, procoracoid and
coracoid, a complete clavicle on both sides, an interclavicle and a sternum
(Figs 3-5). The left humerus is present in natural articulation with the glenoid,
but is damaged proximo-anteriorly. The olecranon portion of the ulna is pre-
served in contact with the left humerus. On the right side the clavicle articulates
with the acromian process of the scapula, which is complete below the level of
this process. The proximal part of the humerus is preserved in position in the
glenoid cavity. A complete coracoidal plate is also present, but is slightly
obscured by the overlying clavicle.

The scapula is a long, relatively slender bone. The acromian process,
situated low down close to the level of the glenoid, is a continuation of the
anterior edge of the scapula blade and is formed more by an excavation of the
anterior edge of the blade than by a separate bony projection (see Watson (1960)
fig. 12). The upper half of the scapular blade is bent back relative to the lower
half and the bone can thus be divided into an acromian section and an upper
section, lying at about 25° to each other. Below the acromian process the base
of the scapula is turned sharply inwards to meet the procoracoid. The glenoid is
a well-defined hollow bounded by a sharp, flared rim, and faces laterally rather
than posterolaterally as in other dicynodonts (Watson 1960).

The procoracoid/coracoid plate is more complete on the right side, but is
partially obscured by the clavicle. There are no clear boundaries between the

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

cla. cla.

Fig. 3. Cistecephalus sp. RC 298. Semi-articulated pectoral girdle in left lateral view.

two elements making up the plate, but a ridge running forward from the rear
of the glenoid cavity may indicate a large coracoid, forming most of the ventral
part of the glenoid cavity. The anterior edge of the procoracoid is expanded to
form a fairly robust process which abutted against the clavicle of its side. A
procoracoid foramen is present half-way between the ventral limit of the anterior
procoracoid process and the ventral rim of the glenoid cavity.

The clavicle is a fairly stout bone with a wide contact with the inner surface
of the scapula at the level of the acromian process, and a broad ventral meeting
with the clavicle of the other side in the ventral midline. Along the midline each
clavicle is dorsoventrally expanded, so that the pair meet over a wide area. This
connection seems to have been a strong one and is preserved in most specimens.
The median plate formed by the two clavicles in the ventral midline is overlain
by the front of the interclavicle, less robust in comparison. Half-way down the
posterolateral surface of its shaft the clavicle carries a recess, bounded laterally
by a thin crest of bone, which in size and position is suited for articulation with
the anterior procoracoidal process. This articulation would have served to
strengthen the connection between clavicle and scapulocoracoid, and prevent
forward and inward displacement of the scapulocoracoid resulting from forces
at the glenoid.

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 221

The partially exposed interclavicle is seen as a flat, rectangular sheet with
its anterior end obscured by the meeting between the clavicles, and its posterior
end lying above the sternum. The sternum is fully exposed and most of its
structure can be made out. It is wide and flat-edged anteriorly and tapers
posteriorly to a fairly narrow median spine. A prominent ventral median crest

14

ETAT) HE

13

i

i
hh

12

11

HEEL
ee)

\ Hy
10

;

Ou
|

Fig. 4. Cistecephalus sp. RC 298. Stereophotographs. A. Left scapulocoracoid
in lateral view. B. Pectoral girdle in ventral view.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Cistecephalus sp. RC 298. A. Left scapulocoracoid in lateral view. B. Pectoral girdle
in ventral view.

runs the length of the bone. On each side three processes for the attachment
of ribs can be made out, one behind the anterolateral corner of the bone, one
half-way down the side and one on the posterolateral corner. On the right side
two ribs are in near natural articulation with the two most anterior processes.
The distances separating the three processes are approximately the same.

Several additional specimens show further details of the pectoral girdle.
In BPI 2915 a good scapula, with cleithrum, is present, differing from that in
RC 298 in that it extends straight dorsally without a posterior twist in the
dorsal half. On the left side the coracoid and procoracoid have become detached
from the scapula, and the suture between the two ventral elements can be seen
running vertically and transversely through the middle of the ventral part of the
glenoid. The procoracoid foramen is situated in the centre of the procoracoid,
well away from the glenoid cavity. In BPI 696 the right scapula, of which the
dorsal part is damaged, shows the acromian region well. The procoracoid is
present, though apparently incomplete, and where the two bones have moved
slightly apart it can be seen that the scapula extends ventrally for a considerable
distance on the inside of the procoracoid. In BPI 4086 and BPI 2124 the scapula
is bent towards the rear as in RC 298.

In BPI 2915 a distinct cleithrum is present, and this appears to have been
the case in the other specimens as well. Von Huene (1942) described this element
in material later referred to Kawingasaurus by Cox (1972).

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 223

In GS K224 (Fig. 12A) the interclavicle is not completely preserved and
the sternum has apparently been eroded away. Seen in the midline above where
the sternum would have lain are the bases of the coracoids, meeting over the
anterior parts of their inner surfaces in what appears to be a natural relation-
ship; all the remaining postcranial elements in this specimen are in natural
articulation with each other. In BPI 2915 a similar meeting of the coracoids in
the midline, over the rear of the interclavicle and the front of the sternum, seems
likely.

The best humerus is that of the right side in BPI 4086 (Figs 7-8), but the
bone is also well preserved in BPI 506 (Fig. 9) and BPI 696 (Fig. 10). The bone
is almost as broad as it is long, and is characterized by greatly enlarged processes
serving for muscle attachment. There is a clearly demarcated head (caput
humeri), forming the most proximal corner of the bone and facing dorsally as
a prominent, slightly rounded condyle. The head is especially strongly developed
in BPI 506. Behind this the posterior corner of the humerus is extended as a
powerful posteromedial process, which consists of a neck portion and an
expanded terminal portion. In front of the humeral head the inner, anterior
edge of the bone is drawn forward as a broad, thin deltopectoral crest, with a
thickened leading edge.

rad.con.

uln.

pro. for

Fig. 6. Cistecephalus sp. BPI 2915. Partial pectoral girdle and left forelimb. Manus seen in
ventral view.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

pm. proc.

cap. hum.

cap.hum.

ent.con.

ect.con.
ent.con.

uln.con.

uln. con. ect.con.

rad.con.

pm. proc.
dp. cr

rad.con. Y= ent. con.

uln. con.

Fig. 7. Cistecephalus sp. BPI 4086. Right humerus. A. Dorsal view. B. Posterior view.
C. Ventral view.

The proximal part of the humerus is convex dorsally and concave ventrally.
The distal half is twisted so that the plane in which the distal edge lies is at right
angles to the plane in which the deltopectoral crest and posteromedial process
lie. In this way, when the proximal part of the humerus is brought to lie in the
horizontal plane, the articulatory surfaces for the radius and ulna on the distal
edge face forward and laterally. With the distal edge in the horizontal plane,
the most anterior part is formed by the broad ectepicondyle. A thin crest leads
medially from the ectepicondyle to merge with the body of the humerus near
the waist of the bone. Below and behind the ectepicondyle is a prominent,
rounded condyle for the radius, separated from the ectepicondyle by a distinct
groove. Behind this is a smaller but equally distinct condyle for the ulna; the

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 225

distal edge of the humerus is thinner in this region, and the ulnar condyle
stands clear of both the dorsal and ventral surfaces of the bone, while on the
dorsal surface there is a slight depression medial to the condyle. This depression
is deepest in BPI 506, where the condyles are very powerfully developed.
Posterior to the ulnar condyle the humerus is extended as a long, relatively
slender entepicondyle. At the base of this process, near the waist of the bone,
the humerus is pierced by the entepicondylar foramen.

Fig. 8. Cistecephalus sp. BPI 4086. Stereophotographs of right humerus.
A. Posterior view. B. Dorsal view.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

cap.hum.
a Pp. cap. hum.

enaN

Fig. 9. Cistecephalus sp. BPI 506. Left humerus. A. Dorsal view. B. Posterior view.
C. Ventral view.

The radius is well preserved in BPI 2915 (Fig. 6) and BPI 2124 (Fig. 11).
The length of the bone in BPI 4086 is equal to that of the humerus between the
humeral head and the outside surface of the radial condyle. The ulna is fairly
robust, widened proximally for the meeting with the humerus and flared distally
to meet the bones of the carpus over a fairly broad area. A posterior crest on
the distal end gives the bone a triangular cross-section where it meets the carpus.

The ulna (Figs 6, 10-11) is present in BPI 2915, BPI 696 and BPI 2124.
It is characterized by a wide and powerful olecranon process extending up above
a notch for articulation with the ulnar condyle on the humerus. The shaft of
the ulna is slender, and the distal extremity is rounded in section and narrower
than that of the radius. On the posterior surface a deep groove extends down
the shaft from the level of the articulatory facet for the humerus.

The carpus is best seen in BPI 2915 (Figs 6, 13), where the ventral surfaces
of the bones are exposed. The radius and ulna are separated from the carpus

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS DOM,

by a slight gap, within which three small bones lie alongside the ulna. The one
nearest the ulna can be regarded as a pisiform, drawn medially during the
pulling apart of the wrist, while of the other two the lateral one probably
represents an ulnare, and the medial element the intermedium. The remainder
of the carpus is still undisturbed and the bones can be identified with a fair
amount of certainty. There are a large medial radiale and a centrale, followed
distally by a row of four distalia. In BPI 2124, the specimen described by Brink
(1952), an ulnare, an intermedium and a large radiale are present, as well as
four, possibly five, distalia (Fig. 11). A centrale is also present. The doubtful
fifth distal is the most median one, which may actually be part of the radiale.
The middle distale is smaller than figured by Brink, who mistook part of the
third metacarpal as being part of it. The most lateral distale, behind the fourth

pm. proc.

pm.proc.

dp.cr

ect.con.

ent.con. ue ae zm re
uln.con. rad.con. ent.con. uln.con.

1 cm.

olec. proc. +

Fig. 10. Cistecephalus sp. BPI 696. Left humerus. A. Anteroventral view. B. Ventral view.
C. Left ulna in anterior view.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

1cm.

Fig. 11. Cistecephalus sp. BPI 2124. Left and right forelimbs as preserved. Left manus in
ventral view, radius and ulna of both sides in posterior view.

metacarpal, was regarded by Brink as metacarpal V. In BPI 4120 a row of four
distalia is seen.

In BPI 4120 (Fig. 12B) four metacarpals are present in the right manus,
the two middle ones being the largest. These are interpreted as being, from
medial to lateral, the first to fourth metacarpals; a small bone close inside the
fourth metacarpal may be a reduced fifth metacarpal. Four metacarpals are
present in BPI 2124 (Fig. 11), Brink’s (1952) specimen, these being nos. I, II,
III and IV. In BPI 696, where the carpus is very incomplete, a splint of bone
alongside the fourth digit may represent a fifth metacarpal. Three metacarpals
are in position in BPI 2915 (Figs. 6, 13A). These have been identified as nos. I,
III and IV. A loose, damaged element lying medial to the rest of the wrist bones
may possibly be a dislodged first metacarpal.

A digital formula can be obtained in GS K224, the specimen described by
Keyser (1973). This is the only specimen to show a hand with five digits
(Figs 12A, 13B). The left hand, seen in ventral view, is best preserved. The first
digit is reduced but appears complete, with the two phalanges separated by
only a narrow line of fusion. There is a small metacarpal. Digits 2 and 3 are
robust, also with the first and second phalanges fused, and are provided with
powerful ungual phalanges. The ungual phalanx of digit 2 is damaged but was
clearly of a size comparable with that of 3. The fourth digit is reduced in size,

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 229

and about the size of the first, while the fifth is also small. Here there is an
ungual phalanx smaller than that of 4, and a single phalanx is preserved behind
this. The right manus is also preserved, and preparation of digit 5 shows that
it is complete; it is smaller than the others but consists of an ungual phalanx
and fused first and second phalanges. The digital formula of this specimen is
fous 2,3,3,3,3.

Several other specimens show details of the manus. In BPI 2915 (Figs 6,
13A) three digits are present, two of them nearly complete. In two digits,
regarded as the second and third, the first and second phalanges are immovably
united, and can be distinguished from each other only by a thin line of fusion.
Medially a third fused pair of phalanges lies separated from the radiale by a
space large enough to accommodate a metacarpal; this probably represents
the first digit, with the first and second phalanges fused. The ungual phalanges
of the second and third digits are damaged, but that of the third digit is of

Fig. 12. Cistecephalus sp. A. GS K224. Left and right manus in ventral view, as preserved.
B. BPI 4120. Right manus in ventral view.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Cistecephalus sp. A. Stereophotographs of BPI 2915, left manus in
ventral view. B. Stereophotograph of GS K224. Left and right manus in ventral
view.

considerable length. Only a fragment of the first phalanx of the fourth digit
is preserved. In BPI 4120 (Fig. 12B) four digits are present, these being the
first, second, third and fourth. In all the first and second phalanges are closely
united but in only the first digit have the two reached the stage of semi-fusion
found in the case of BPI 2915. The ungual phalanges are long and sharp-pointed,
particularly that of the second digit. In the second, third and fourth digits the

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 231

first phalanx is shortened to almost disc-like proportions. In overall appearance
the manus is short, wide and powerful.

In BPI 696 a left manus, with a very incomplete wrist, is preserved. As
interpreted, the first, second, third and fourth digits are present. In the first
digit the first and second phalanges are fused, while in the other digits this is
the condition in the first two phalanges, the ungual phalanx being movable on
the distal tip of the second phalanx. In Brink’s (1952) specimen, BPI 2124, a
part of the first metacarpal is preserved, and possibly part of the first phalanx
(Fig. 11). Digits 2 and 3 are complete, and the first two phalanges of each are
firmly united and separated by a line of fusion. The fourth digit is very incom-
plete, but at least one phalanx is present.

The absence of a fifth digit in all but one of the specimens available is
probably best regarded as being due to accidents of preservation resulting
from this digit’s relatively delicate structure.

Pelvic girdle and hind limb

The pelvic girdle is preserved in BPI 696 (Fig. 14B), BPI 2450 (Fig. 16B)
and BPI 4086 (Figs 14A, 16A). The ilium lies well forward in relation to the
other pelvic bones with the blade forming a strong anterior process. A posterior
process is clearly demarcated from the base of the bone. Above the acetabulum
the base of the ilium is drawn out as a buttress terminating in a laterally directed
process, while in front of the acetabulum its anterior surface is broad and flat.
The pubis forms the ventral part of the acetabulum and the anterior wall of the
obturator foramen. Ventrally it carries a low keel set off laterally to the ventral
border of the bone. The ischium forms a lower rim for the acetabular socket
which is not as strongly developed as the iliac buttress. The acetabulum is
shallowest on the ischtum and deepest on the ilium, where the iliac buttress
overhangs the socket. All three bones are firmly united and the puboischiadic
plate lies at an angle to the illum, suggesting that the two ischia approached
each other in the ventral midline. There is, however, no indication of a bony or
cartilaginous symphysis between the two halves of the pelvis.

In BPI 2450 two clear depressions and a third less distinct one on the
inside of the iliac blade mark the points of attachment of the sacral ribs. The
most anterior of these lies some way behind the anterior tip of the ilium and is
shallow and fairly long. The second is deep in comparison and lies above the
anterior border of the acetabulum. The third depression may mark the attach-
ment of the last sacral; it lies close behind the preceding one. If the sacral
vertebrae were held horizontally, these depressions, which lie some distance
below the dorsal border of the iliac blade, show that the pelvis must have been
rotated so that the puboischiadic plate lay largely behind the acetabulum.

The ilium of BPI 2450 (Fig. 16B) differs in some respects from that of
BPI 696 and BPI 4096. The iliac blade is shorter than in BPI 696, and the anterior
border of the anterior process is turned out laterally and descends to meet the
highest point of the acetabular buttress well behind the anterior edge of the

PEW ANNALS OF THE SOUTH AFRICAN MUSEUM

bone. As a result, the iliac blade is laterally concave. There is no sign of a
separate posterior process at the base of the iltum, but the dorsal crest of the
bone does extend back a short way. In overall view, therefore, the ilium is a
high, narrow bone. The acetabulum appears to be deeper than in BPI 696, and
bounded by a more prominent rim; the posteroventral, ischial part of the rim
is almost as strongly developed as the anterodorsal, iliac part.

The structure of the hind limb can be determined in BPI 4096 (Figs 15, 16).
Both femora are preserved, the right one being the better exposed. The bone is
a remarkable one, with a strongly developed head carried medial to the inner
border of the shaft and standing well forward of the anterior surface of the bone.
The greater trochanter is confined to the lateral corner of the proximal part of

Fig. 14. Cistecephalus sp. Pelvic girdle in right lateral view. A. BPI 4086. B. BPI 696.

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 233

gr. tr

_. cap. fem.

Cn.cr

: iceman;

Fig. 15. Cistecephalus sp. BPI 4086. A. Right femur in anterior view. B. Right tibia in medial
view. C.Right tibia in anterior view. D. Left tibia in lateral view. E. Left fibula in ?lateral
view.

the bone, and does not extend down the shaft as a separate crest. The antero-
posteriorly compressed shaft is slightly rounded anteriorly but almost perfectly
flat behind. Distally the bone is expanded for the articulatory areas with the
tibia and fibula. The proximal end of the bone between the head and expanded
greater trochanter is essentially as narrow as the shaft.

The proximal ends of the tibia and fibula are preserved on the right side
but both bones are fully exposed on the left side, except where the distal end
of the tibia is partially obscured by the foot. The tibia is a well-ossified bone
with the proximal articular surface roughly triangular in outline. There are two
clear areas for articulation with the condyles of the femur, forming two corners
of a triangle completed by an anterior cnemial crest. Proximally the lateral

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. Cistecephalus sp. A. Stereophotograph of BPI 4086. Right ilium,

femur and proximal part of tibia and fibula. Left astragalo-calcaneum in

proximal (dorsal) view. B. Stereophotograph of BPI 2450. Right half of pelvic
girdle in lateral view.

surface of the bone bears a groove which may have received the dorsal part
of the fibula (see Watson 1960). Distally the shaft becomes narrower and
lateromedially compressed, but it widens again to form a rounded condyle for
articulation with the tarsus.

The slender fibula is slightly expanded proximally where it lay up against
the lateral side of the tibial head. It could not have had more than a small area

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 235

of articulation with the femur, but distally it widens to form a small, flat articu-
latory surface for the tarsus.

The left tarsus and pes, slightly separated from the tibia and fibula, are
preserved nearly complete and are seen in ventral view (Figs 16-17). The astra-
galus and calcaneum appear to be unique among dicynodonts in that the two
bones are united and can be distinguished from each other only by a notch and
a line marking the fusion between them. Their proximal surfaces combine to
form a wide, concave articulatory area for the tibia and fibula. The largest

Fig. 17. Cistecephalus sp. BPI 4086. Stereophotographs. A. Left ilium, femur,
tibia, fibula and pes. B. Left astragalo-calcaneum and pes in ventral view.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

part of this surface is formed by the astragalus and is more concave than the
smaller calcaneal surface which met the fairly flat distal part of the fibula. The
distal surface of the calcaneum, more extensive than that of the astragalus, is
rounded for its meeting with the tarsal bones. The size and strength of the
structure, and its well-developed articulation with the tibia and fibula, are
surprising in a reptile such as Cistecephalus, and suggest considerable movement
at the ankle joint, including rotation between the astragalo-caleaneum and the
tibia and the fibula, and flexure and extension between astragalo-calcaneum
and the foot.

In front of the astragalo-calcaneum are two proximal tarsal bones, one
seen only very indistinctly. These probably met the medial part of the calcaneum
and the lateral part of the astragalus. Following on these is a row of four distal
tarsals. The most medial of these meets the first metatarsal and part of the
second metatarsal; the third meets the third metatarsal while the fourth and
largest touches the base of both fourth and fifth metatarsals. The metatarsals
of the second, third and fourth digits are short and fairly slender. Only in the
fifth digit can three phalanges be distinguished, and these are not very clearly
shown. The rest of the digits have suffered varying amounts of damage. Only
two phalanges are found in the second, third and fourth digits, while in the
first digit only one separate phalanx could be identified. In this digit, a small
bony flake might represent part of a terminal phalanx.

RANGE OF MOVEMENTS AND MUSCULATURE OF THE LIMBS

Attempts at reconstructing musculature and limb movements in extinct
reptiles, particularly in the case of forms which have no living descendants,
inevitably involve some degree of speculation, but if this shortcoming is clearly
borne in mind the exercise can be a useful and informative one. In the following,
only general features in the bony structure and only those main muscle groups
which are expected to be present in all terrestrial reptiles are considered (see
Romer 1922, 1944). Cox (1972) has given a more detailed account of the muscu-
lature and possible range of movement in the forelimb of Kawingasaurus, where
the material included separated bones which could be manipulated and where
the relations between the joints could be better determined than was possible
in the case of the Cistecephalus material available for the present study.

Pectoral girdle and forelimb

The pectoral girdle of Cistecephalus is unusual in several respects. The
girdle as a whole lies well forward, leaving a short neck with the scapula rela-
tively close behind the skull. The glenoid cavity faces almost straight laterally
while the blade itself, instead of sloping slightly forward, is vertical or even
posteriorly inclined in its upper half. The base of the scapula has rotated
forward and the acromian process, rather than being a separate projection, is
part of the general anterior edge of the scapular blade. The clavicle appears
to have lain at an angle of approximately 45°, from an acromian connection

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS VSG

ast.

Fig. 18. Cistecephalus sp. BPI 4086. A. Left astragalo-calcaneum in proximal (dorsal) view.
B. Left pes in ventral view.

straight down to a second support at the anterior edge of the procoracoid. This
may be associated with the forward position of the shoulder girdle as a whole
relative to the vertebral column and skull. Besides being strongly braced by
scapulae and procoracoids, the clavicles also meet in the ventral mid-line in an
extensive face-to-face contact.

These features of the pectoral girdle can be interpreted in terms of resistance
to strong lateral forces which were imposed upon the forwardly rotated glenoid
region of the scapulocoracoid. Such forces would have been transmitted
internally on to the clavicle which, instead of bracing the scapulocoracoid
against stress from a posterolateral direction, had to support that bone against
forces of almost purely lateral origin. To help counter these forces, the acromian
articulation was extended dorsally so that the clavicle enjoyed an extensive
dorsoventral overlap with the scapula; at the same time the ventral part of the
clavicle rotated back to meet the anterior edge of the procoracoid in an extra
point of support. The substantial meeting between the clavicles in the ventral
mid-line may be regarded as a further strengthening device.

The relationship between the glenoid cavity and humerus is also of signifi-
cance. The glenoid is a well-finished socket, with a sharp dorsal rim on the
supraglenoid buttress. The humerus carries a distinct humeral head, set off as
a dorsal condyle on the proximal part of the bone, and it is evident that all
movement between humerus and glenoid took place about its slightly convex
articulatory surface. With the humeral head dorsally oriented the deltopectoral
crest faces forwards, and the twist of the humerus along its long axis is such
that the condyles for the radius and ulna are directed forward at an approxi-
mate angle of 45° to the horizontal. With the head of the humerus facing
forward in the glenoid cavity the distal condyles of the humerus face straight

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

down. Judging by the relations between the head of the humerus and the
glenoid, this 45° rotation, accompanied by forward and back swing, was the
most general movement of the upper arm.

The highly developed articulations between the radius and ulna and the
humerus indicate that considerable movement between upper and lower fore-
arm was characteristic of the animal. While the radius seems restricted to a
rocking motion against the radial condyle of the humerus, the ulna is notched
for articulation with the ulnar condyle and distal edge of the humerus. The
depression on the humerus immediately proximal to the ulnar articulation
suggests, too, that the ulna could be straightened on the upper arm by the
action of muscles attaching on its high olecranon process.

Powerful extension of the forearm on the humerus, best explained as a
digging or scraping movement for which the broad, powerfully constructed
manus with its fused phalanges and strong claws was clearly suited, would
account for the lateral forces directed on to the glenoid region of the scapulo-
coracoid. A consideration of some of the chief muscles associated with the
pectoral girdle and forelimb supports this interpretation.

In a dicynodont such as Cistecephalus it is expected that the main muscles
bracing the scapula on the side of the trunk were the trapezius muscle anteriorly,
inserting on the anterior edge of the bone and on the cleithrum, and the levator
scapulae superficialis muscle. These arose from the posterior and posterodorsal
part of the rounded occipital area of the skull and from the ligamentum nuchae.
Posteriorly the scapula would have been held in position by the serratus anterior
superficialis muscle, arising from the spines of the dorsal vertebrae and inserting
on the dorsal edge of the scapula. The backward slope of the-dorsal part of the
scapula in some specimens may indicate a division of the insertional areas of
these three muscles: the levator scapulae superficialis anteroventrally, the
trapezius anterior anterodorsally and the serratus anterior superficialis
posterodorsally.

Muscles which are generally responsible for movement of the humerus
are, superficially, the latissimus dorsi, arising from the surface of the back and
flank behind the pectoral girdle, and the deltoideus group, arising from the
scapula and clavicle as the scapular and clavicular deltoid muscles. In Ciste-
cephalus the area for insertion of the deltoideus is clearly marked off from the
head and more posterior part of the humerus by a crest running from the base
of the head to the ectepicondyle. The pectoralis muscle normally arises from
the sternum and ribs and inserts on a process below the proximal end of the
humerus. In Cistecephalus the insertion areas of both the deltoideus muscles,
pulling the humerus forward and up, and the pectoral muscle, pulling the
humerus back and down, are greatly expanded as a powerful deltopectoral
crest, giving both muscle groups added leverage on the upper arm and reflecting
their increased size. Another muscle important in locomotion is the sub-
coracoscapularis, which arises from inside the girdle to insert near the head of
the humerus and which serves to pull the humerus back—in Cistecephalus the

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 239

prominent medially oriented internal process marks the point of insertion of
this muscle, and is evidence of its increased importance during locomotion.
Cox (1972) suggests that due to the similar medial position of the internal
process relative to the condyle in Kawingasaurus, contraction of the subcoraco-
brachialis would also have resulted in a downward thrust of the humerus.
Other muscles inserting on the humerus would have been the scapuichumeralis
anterior, arising from the forward part of the girdle and inserting near the head
of the bone, and the supracoracoideus, extending from the anterior part of
the coracoid plate to the underside of the humerus.

Movement of the forelimb is brought about by the triceps muscle dorsally
and the biceps and brachialis muscles ventrally. In Cistecephalus the areas of
origin of the triceps on the humerus were the posteromedially extended internal
process and the expanded area behind the ectepicondyle which, as shown
above, is demarcated from the deltoideus insertion area by a crest. The third
triceps head arose from the scapula. In Cistecephalus the insertional area of the
muscle, the olecranon process, is greatly enlarged over the normal dicynodont
condition, and it is evident that the triceps, though short, was a muscle of
considerable size and able to exert powerful extension forces on the lower arm.

Besides extension and flexion movements of the forearm on the humerus,
it is likely that anteroposterior swing of the forearm was possible (see Cox 1972).
Such movements would have been controlled by flexor and extensor muscles
arising from the ectepicondyle and entepicondyle of the humerus; in Ciste-
cephalus these are powerfully developed and extend well in front of and behind
the condyles for articulation with the radius and ulna.

Seen in its entirety, the structure of the pectoral girdle and forelimb in
Cistecephalus, characterized by strong processes for muscle attachments and
well-finished joints and articulations carrying a minimum of cartilaginous
lining, suggests that the animal was capable of using its forelimbs in controlled
movements of considerable power. As suggested above, these movements may
be interpreted as forming part of habitual digging or scraping activities.

Pelvic girdle and hind limb

In the hind limb all joints and articulations are, as in the forelimb, charac-
terized by well-ossified and smooth bony surfaces. The relationships of the femur
to the pelvic girdle and lower hind limb are of particular significance. The
prominent, rounded femoral head projects medially and forward on the narrow
proximal part of the bone, and it is clear that a considerable range of movement
was possible at the acetabulum. The offset head suggests that the femur was
held with the lower end drawn in close to the body in a mammal-like fashion,
with movement taking place mainly in a vertical, parasagittal plane. In addition,
the fact that much of the articulatory surface on the head faces forward as well
as inward suggests that the femur could rotate medially along its long axis so
that the anterior part of the head came to lie deeper within the acetabulum.
In this position the greater trochanter on the proximolateral corner of the bone

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

is turned forward, while the condyles for the tibia face slightly outward, turning
the lower leg away from the body. Since the condyles on the femur for articula-
tion with the tibia face distally as well as posteriorly, it is likely that the lower
limb could have been extended far forward on the femur.

The strengthened astragalo-calcaneum provided a single, large articulatory
surface for movement between the foot and the tibia and fibula. The rounded
distal end of the tibia has a radius of curvature smaller than that of the broadly
concave astragalus, but the fibula terminates in a relatively flat articulatory
surface which coincides more closely with the surface of the calcaneum. Move-
ment between the astragalo-calcaneum and the tibia and fibula must have
involved considerable rotation and even sliding between the bones of the lower
limb and tarsus. )

Increased rotatory movement between the lower leg and the pes could be
the result of the femoral rotation discussed above, during which the lower leg
was directed away from the long axis of the body. For example, if the femur
were rotated inwards along its long axis while in the forward position, the
lower leg could have been flexed back and away from the body in a scraping
movement; however, rotation of the femur would have resulted in lateral dis-
placement of the lower end of the tibia and fibula, and could have occurred
only if compensatory movements were possible between the lower leg and pes
while the pes was firmly placed on the ground. Thus, longitudinal rotation of the
femur and the development of a broad articulatory surface on the astragalo-
calcaneum in Cistecephalus are probably related phenomena, and are indications
of unusual hind limb function in the living animal.

Although areas of muscle attachment are less clearly marked on the bones
of the hind limb than on those of the forelimb, some observations can neverthe-
less be made regarding the possible arrangement of the hind limb musculature.
In reptiles several muscles are responsible for movement of the femur. The
puboischiofemoralis internus arises from the lumbar region and inner surface
of the girdle and inserts on the femur near its head. In Cistecephalus a large,
forward-facing area on the base of the ilium probably served for the origin of
part of this muscle, which served to pull the femur forward. The iliofemoralis
muscle in reptiles arises from the ilium and inserts on the upper surface of the
femur, drawing it back; in Cistecephalus a distinct greater trochanter on the
proximolateral corner of the bone shows that the muscle inserted in a primitive
mammal-like fashion. Other muscles which generally draw the femur back are
the caudifemoralis muscles arising from the tail, but the tail in Cistecephalus
and other dicynodonts is weak and it is unlikely that these muscles were very
important—no trochanter for their insertion is found on the femur. However,
backward pull on the femur would have been provided by the ischiotrochan-
tericus (obturator internus) running from the inside of the ischium to the head
of the femur. In addition, an adductor femoralis arising from the pubo-ischiadic
plate would have attached down the ventral surface of the femur and served
to pull the leg back.

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 241

For the lower leg a group of extensor muscles, the iliotibialis and femuro-
tibialis, run from the girdle and femur to insert on the tibia. A distinct cnemial
crest on the Cistecephalus tibia indicates a well-developed femurotibialis, while
the distally placed tibial condyles on the femur show that the lower limb could
have been almost fully straightened upon the femur. Flexor muscles for the
lower leg would have included the puboischiotibialis, running from the pelvic
girdle to the lower leg.

The hind limb has clearly been modified in a different way to the forelimb.
While the humerus has become broad and robust, with enlarged processes for
the attachment of muscles, the femur is slender in comparison with other
dicynodonts. The manus is relatively immobile and powerfully constructed,
while the pes again is small but very mobile on the lower leg. In terms of length
there are also significant differences between the bones of the forelimbs and
hind limbs. Thus, the femur and tibia are both longer than their counterparts
in the forelimb, the total length of the humerus and radius being approximately
73 per cent of the total length of femur and tibia. Although the point of articula-
tion between the humerus and glenoid cavity is lower on the body than is the
articulation between the almost vertically held femur and the acetabulum, it
appears that the pelvic region of the body was held higher off the ground than
the pectoral region during ordinary locomotion.

DISCUSSION

In its skull and jaw, axial skeleton, pectoral and pelvic girdles as well as
in its appendicular skeleton, Cistecephalus shows a wide range of modifications
which together make it probably the most aberrant of dicynodonts. It follows
that Cistecephalus was adapted to a very distinctive mode of life, and that the
general nature of the animal’s habits should be indicated by these skeletal
modifications. The peculiarities of the skeleton are all related to strengthening
of individual bones or functionally integrated groups of bones and reflect
increase in the size and power of various parts of the musculature. The clearly
defined and in some cases enlarged areas of articulation between bones of
the limbs and girdles, and the distinct articular condyles, with a minimum of
cartilaginous capping, on the humerus and femur, all point to a high degree of
muscular control over the limbs during movements of considerable power.
Indications are that the forces exerted during these movements were far in
excess of the requirements for normal locomotion, and it can be deduced that
the limbs were frequently used during additional activities that formed an
integral part of the animal’s life. Moreover, the overall similarities between
the skulls of Cistecephalus, Kawingasaurus and Cistecephaloides, and between
the pectoral girdle and forelimb of Cistecephalus and Kawingasaurus, suggest
that the family Cistecephalidae was adapted to a broadly uniform way of life.

Cox (1972) concluded on the basis of available material that Kawingasaurus
was an active digger, and it is now possible to assign a similar way of life to
Cistecephalus and, by inference, to Cistecephaloides. Compared with living

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

au i
(TS
[Qe

Fig. 19. Left scapulocoracoids, humeri and ulnae. Not to scale. A. Oudenodon sp.,
SAM-11114: a—scapulocoracoid in lateral view, b—dorsal view of humerus with proximal
end horizontal, c—dorsal view of humerus with distal end horizontal, d—ulna in anterior
view. B. Cistecephalus sp., from several specimens: a—scapulocoracoid in lateral view, from
RC 298, b—dorsal view of humerus with proximal end horizontal, from BPI 4086 (drawn
from right side), c—dorsal view of humerus with distal end horizontal, from BPI 4086 (drawn
from right side), d—ulna in anterior view, from BPI 696. C. Talpa europaea: a—scapula in
lateral view, b—humerus in posterior view, c—humerus in anterior view, d—ulna in anterior
view.

b Cc

mammal groups, the skeletal anatomy of Cistecephalus bears the closest overall
resemblances with those forms which are adapted to a digging or burrow-
ing way of life (Hisaw 1923, Yalden 1966, Reed & Turnbull 1965). Such
features as a rounded occipital region, the broad humerus with powerful
processes for muscle attachment, and a greatly enlarged olecranon process in
Cistecephalus are found in analogous form in the European mole Talpa europaea
and the Cape Golden mole Chrysochloris asiatica (Fig. 19). In these living
forms the rounded occiput is related to increased size of the shoulder and neck
muscles, while the robust humerus and high olecranon process reflect the
increased power of muscles used during burrowing. The broad manus of Ciste-

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 243

cephalus, with three enlarged digits and fused phalanges, is suited to scraping
or digging, with the highly mobile pes shovelling loosened soil to the rear and
side of the animal.

The limbs and limb girdles provide the strongest evidence for digging or
burrowing activities, but other features of the skeleton can also be interpreted
in terms of this way of life. Thus strengthening of the skull by broadening the
skull roof and eliminating the interpterygoidal vacuity in the basicranial girder,
and loss of mobility between the cervical and dorsal vertebrae could indicate
resistance to forces encountered by the skull and transmitted to the vertebral
column during burrowing. The anterior position of the pectoral girdle relative
to the vertebral column is also a feature found in both the true moles (Talpidae)
and the Cape Golden mole (Chrysochloridae) (Campbell 1938, 1939) and it
is significant that outward rotation of the hind limb is a characteristic of the
true mole Jalpa (Yalden 1966).

The family Cistecephalidae, including Cistecephalus, Kawingasaurus and
Cistecephaloides, therefore represents a dicynodont radiation into a burrowing
or fossorial way of life. Food sources such as plant roots and small invertebrates
become available to animals capable of powerful digging, while the true bur-

PEARL MN
rT

\\

aN

Fig. 20. Formal reconstruction of Cistecephalus in lateral and anterior views, mainly from
BPI 4086 and SAM-—10665. Approximately 2 natural size.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

rower has available an effective means of escape from predators. The degree of
burrowing characteristic of the various cistecephalid genera will remain
uncertain until further more definite discoveries are made; for instance, the
discovery of a complete skeleton in an in-filled burrow would establish a
highly-developed fossorial way of life for the particular species. Also, material
at present available does not allow the actual nature of the cistecephalid burrow-
ing activity to be identified and compared with the ‘swimming’ burrowing style
of Talpa and its allies, or the ‘running’ style of the Chrysochloridae (Hisaw 1923,
Yalden 1966, Campbell 1938). It is, however, clear at this stage that the Ciste-
cephalidae were committed to intensification of digging or burrowing activities
which probably formed only a minor part of the general way of life of other
dicynodont groups. |

SUMMARY

In both its cranial and postcranial skeleton Cistecephalus shows evidence
of adaptations to a very specific mode of life. The skull is structured within
the characteristic dicynodont framework, but its many substantial modifications
appear to be linked with specialized features of the postcranial skeleton. Seen
in its entirety, the skeleton of Cistecephalus represents an osteological extreme
in dicynodont evolution, and all indications are that the living animal, com-
mitted as it was to at least semi-fossorial habits, was a highly unusual member
of South Africa’s Upper Permian fauna.

ACKNOWLEDGEMENTS

Iam particularly grateful to Dr J. W. Kitching of the Bernard Price Institute,
University of the Witwatersrand, Dr A. W. Keyser, Geological Survey, Pretoria,
and Mr and Mrs R. Rubidge of Wellwood, Graaff-Reinet, for the loan of
specimens. Mrs Kathleen Riall of the Department of Palaeontology, South
African Museum, was responsible for the skilful preparation of several speci-
mens; for the photographs I am indebted to Mr Neville Eden of the same
department.

REFERENCES

BoonsTrRA, L. D. 1966. The girdles and limbs of the Dicynodontia of the Tapinocephalus zone.
Ann. S. Afr. Mus. 50: 1-11.

BRINK, A. S. 1950. On a new species of Cistecephalus Owen. Ann. Mag. nat. Hist. (12) 3:
985-997.

BRINK, A. S. 1952. Studies on Karoo reptiles. III. The manus of Cistecephalus. S. Afr. J. Sci. 49:
13-15.
BroILi, F. & SCHRODER, J. 1935. Beobachtungen an Wirbeltieren der Karoo-formation. VI.
Uber den Schadel von Cistecephalus Owen. Sber. bayer. Akad. Wiss. 1935: 1-20.
Broom, R. 1932. The mammal-like reptiles of South Africa and the origin of mammals. London:
Witherby.

Broom, R. 1948. A contribution to our knowledge of the vertebrates of the Karroo Beds of
South Africa. Trans. R. Soc. Edinb. 61: 577-929.

CAMPBELL, B. 1938. A reconsideration of the shoulder musculature of the Cape Golden mole.
J. Mammal. 19: 234-240.

THE SKELETON OF THE MAMMAL-LIKE REPTILE CISTECEPHALUS 245

CAMPBELL, B. 1939. The shoulder anatomy of the moles. A study in phylogeny and adaptation.
Am. J. Anat. 64: 1-39.

CLuver, M. A. 1971. The cranial morphology of the dicynodont genus Lystrosaurus.
Ann. S. Afr. Mus. 56: 155-274.

CLuvER, M. A. 1974. The skull and mandible of a new cistecephalid dicynodont. Ann. S. Afr.
Mus. 64: 137-155.

Cox, C. B. 1959. On the anatomy of a new dicynodont genus with evidence of the position
of the tympanum. Proc. zool. Soc. Lond. 132: 321-367.

Cox, C. B. 1972. A new digging dicynodont from the Upper Permian of Tanzania. In: JoyseEy,
K. A. & Kemp, T. S. eds. Studies in vertebrate evolution: 173-189. Edinburgh: Oliver &
Boyd.

Hisaw, F. L. 1923. Observations on the burrowing habits of moles (Scalopus aquaticus machri-
noides). J. Mammal. 4: 79.

HUENE, F. von. 1942. Die Anomodontier des Ruhuhu-Gebietes in der Tubinger Sammlung.
Palaeontographica 94: 154-184.

Keyser, A. W. 1973. A preliminary study of the type area of the Cistecephalus zone of the
Beaufort Series, and a revision of the anomodont family Cistecephalidae. Mem. geol.
Surv. S. Afr. 62: 1-71.

OwEN, R. 1876. Descriptive and illustrated catalogue of the fossil reptilia of South Africa in
the collection of the British Museum of Natural History. London: British Museum.
REED, C. A. & TURNBULL, W. D. 1965. The mammalian genera Arctoryctes and Cryptoryctes
from the Oligocene and Miocene of North America. Fieldiana, Geology 15: 99-170.
Romer, A. S. 1922. The locomotor apparatus of certain primitive and mammal-like reptiles.

Bull. Am. Mus. nat. Hist. 46: 517-606.

Romer, A. S. 1944. The development of tetrapod limb musculature—the shoulder region of
Lacerta. J. Morph. 74: \-4l.

SEELEY, H. G. 1894. Researches on the structure, organisation and classification of the fossil
Reptilia. Part IX, section 1: On the Therosuchia. Phil. Trans. R. Soc. (B) 185: 987-1018.

WaATson, D. M. S. 1960. The anomodont skeleton. Trans. zool. Soc. Lond. 29: 131-208.

YALDEN, D. W. 1966. The anatomy of mole locomotion. J. Zool. Lond. 149: 55-64.

ABBREVIATIONS
acet. acetabulum
acr. proc. acromian process
a.l. atlas intercentrum
ant. pro. anterior process of ilium
ast. astragalus
at. atlas
ax. axis
ax. f. axis rib
butt. buttress of ilium
e: centrale
cal. calcaneum
cap. fem. caput femoris
cap. hum. caput humeri
cla. clavicle
en. Cr. cnemial crest
cor. coracoid
cr. crest
d. distal carpal, tarsal
dp. cr. deltopectoral crest
ect. con. ectepicondyle
ent. con. entepicondyle
ent. for entepicondylar foramen
fac. hum facet for humerus
gl. glenoid
gr. groove

1c

il.

isch.

lat. pro.
N.S.

obt. for.
od.

olec. proc.

pa. fac.
par.

pi.

pm. proc.

post. proc.

pro.
pro. for.
pro. p.
pub.

iP;

ra.

(PA.

rad.

rad. con.

humerus

ANNALS OF THE SOUTH AFRICAN MUSEUM

intermedium
interclavicle

ilium
ischium

lateral process
neural spine
obturator foramen

odontoid

olecranon process
facet for proatlas
parapophysis

pisiform

posteromedial process
posterior process of ilium
procoracoid

procoracoid foramen
procoracoid process

pubis
rib
radiale

rib articulation

radius

radial condyle

scapula
sternum
ulnare
ulna

ulnar condyle

Bernard Price Institute for Palaeontological Research, Witwatersrand University,

Johannesburg
Geological Survey, Pretoria
Rubidge Collection, Wellwood, Graaff-Reinet
South African Museum, Cape Town.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA
Nucula (Leda) see Gould, 1845: 37.
Leda plicifera A. Adams, : 50.
Laeda bicuspidata Boa 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
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Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
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>) ‘

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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

MICHAEL A. CLUVER

THE SKELETON OF THE MAMMAL-LIKE REPTILE
CISTECEPHALUS WITH EVIDENCE FOR A
FOSSORIAL MODE OF LIFE

a
VOLUME 76 PART 6 OCTOBER 1978 ISSN 0803-2515

1 507.68

~ ANNALS

‘OF THE SOUTH AFRICAN
MUSEUM

CAPE TOWN

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1. MATERIAL should be original and not published elsewhere, in whole or in part.
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BULLOUGH, W, S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, p. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires, des littorines. Archs
Zool. exp. gen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg! eat and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4):

THIELE, J. 1910. Mollusca: B. Polyplaeantiprs Gastropoda marina, Bivalvia. Jn: 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.

(continued inside back cover)

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

Volume 76 + Band
October 1978 Oktober
Part 6 Deel

x iy) SN
1 er
@ -B.9.B:3.2

74
“Woug np WS

THE DEVELOPMENT OF XENOPUS GILLI
ROSE & HEWITT (ANURA, PIPIDAE)

By

Rea ben Re

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT
(ANURA, PIPIDAE)

By
R. E. Rau
South African Museum, Cape Town
(With 10 figures)

[MS. accepted 6 July 1978]

ABSTRACT

Tadpoles of the Cape clawed frog Xenopus gilli can be distinguished from those of
Xenopus laevis by the distribution of melanophores and differences in dimensions. Xenopus gilli
breeds in cooler water and is less tolerant of rising water temperatures. The species seems to
be more abundant in mountainous terrain than in the low-lying Cape Flats. The breeding
period largely coincides with that of Xenopus laevis, but indications are that it starts and ends
earlier in the year. Where both species occur together some specimens appear to be hybrids.

CONTENTS

PAGE
TER OGG HOM tere 8 cra eR ue van we od, we SEA
Materials andmethed ~. . . . ". . . 249
Wevelopmentvese ti.” > 2) a8 od a Se eg
Further observations on living larvae . . . 259
Summary of distinguishing features . . . 263
Acknowledgements .. 2 © « : « « +. 263
References a ns ee eee oe Sees lek 26S

INTRODUCTION

The Cape clawed frog, as it should preferably be called (Mertens 1970),
was recognized as a new species and described as Xenopus gilli by Rose & Hewitt
in 1927. The type locality of this pretty and easily distinguishable species is the
Silvermine Stream near Clovelly, Cape Peninsula. A smaller, more pointed head,
the absence of the subocular tentacle and the distinctive coloration render it
quite different from the more common Xenopus laevis, which reaches a much
greater maximum size. Both species often occur together, and the validity of the
species X. gilli has sometimes been questioned.

Recently X. gilli has been confirmed as a true species. H. Kobel (in Jitt.
1977) of the Zoology Department of the University at Geneva succeeded in
breeding Xenopus gilli, and also in cross-breeding X. gilli with X. laevis. The
male hybrids proved to be sterile, while the female crosses could reproduce
when mated with either YX. gilli, laevis or muelleri (see Kobel & Du Pasquier
1975; Wabl & Du Pasquier 1976).

In the sandy Cape Flats only a few metres above sea-level, one specimen of

247

Ann. S. Afr. Mus. 76 (6), 1978: 247-263, 10 figs.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Xenopus gilli is occasionally found amongst approximately 1000 YX. laevis
which are being caught commercially for export as laboratory animals (J. Wood
pers. comm.). Sometimes, during the dry season, several specimens of X. gilli
have been found hibernating together under logs, etc., or encapsulated into the
mud of dried-up vleis in the Cape Flats.

There are two specimens, male and female, in the collection of the South
African Museum (ZR18914), which were collected at Citrusdal, in approxi-
mately 1937. Several Xenopus specimens, originally considered to be X. laevis,
were reidentified after the establishment of YX. gilli as belonging to that species
(Mertens 1970), as well as a male specimen at the South African Museum
(ZR2346) collected at Willemsrivier (31°21’S 19°06’E) west of Calvinia in 1898.

Both Citrusdal and Willemsrivier are within the winter-rainfall area of the
south-western Cape. These localities have not been reconfirmed, however, nor
has Xenopus gilli been recorded between the Cape Flats and the already-
mentioned northern localities. The reason might be that a thorough search for
this species has not yet been conducted in the area. In addition, Xenopus gilli is
usually not easily found as the frogs hide in the mud or leaf-layer at the bottom
of the pond. Occasional migration over land of this aquatic species, as is known
to happen with Xenopus laevis, probably does occur. This could perhaps
explain why in a given locality Xenopus gilli is found in one year and not in the
following years.

It is also possible that the two northern localities represent isolated
occurrences, especially if Xenopus gilli is a relict of a formerly more widely
distributed form. Xenopus laevis lives in both acidic and alkaline waters
(Nieuwkoop & Faber 1956) ranging from clear, cold, fast-running mountain
streams to shallow, warm, muddy vleis. If, indeed, the essentially tropical
Xenopus laevis, which occurs in most of eastern and southern Africa from the
Red Sea to the western Cape (Mertens 1970), has invaded the range of the
winter-rainfall species Xenopus gilli, the obviously greater tolerance to environ-
mental changes of XY. /aevis might well be detrimental to the more specialized
Xenopus gilli.

A clarification of the present range of Xenopus gilli and the question of
possible competition with Xenopus laevis, especially in view of possible protective
measures, is desirable. As described above, the presence of Xenopus gilli is not
easy to establish when depending, as hitherto, on the fully developed frog alone.
This suggested the use of the larval stages for distribution investigation. While
the development of Xenopus laevis is fully documented (Nieuwkoop & Faber
1956), both eggs and larvae of Xenopus gilli remained unknown (Wager 1965).

Investigations during the past few years revealed that in several ponds and
dams within the plateau of the mountainous Cape Point Nature Reserve
Xenopus gilli is comparatively common. This in all probability is due to the
curious fact that Xenopus laevis is comparatively rare in these localities.

To establish whether YX. gi//i can be identified in its larval stages and also to
prove or disprove the species validity through cross-breading with Xenopus

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 249

laevis, breeding experiments were undertaken by the author during 1976 at the
South African Museum and by S. McVeigh at the Fauna and Flora Section of
the Cape Department of Nature and Environmental Conservation. While the
females responded to the hormone treatment by laying eggs, the males showed
no reaction. One female, 5,3 cm in length (snout to vent), laid approximately
270 eggs within a few hours with a yolk-size of 1,4-1,8 mm. Thus the larval
stages still remained undescribed.

MATERIALS AND METHOD

The natural breeding season of Xenopus laevis is given as September to
December for the Stellenbosch-Cape Town area (Nieuwkoop & Faber 1956).
It was thought that, since Xenopus gilli is a true western Cape winter-rainfall
species, its breeding season could well start as early as winter. Xenopus tadpoles
were caught during metamorphosis in January—February 1976 in the Cape Point
Reserve. They completed development in an aquarium and proved to be
Xenopus gilli. This, and the dominance of Xenopus gilli over Xenopus laevis in
the reserve, led to the checking of various water accumulations in the reserve on
31 July 1977. Xenopus larvae, approximately 12-15 mm in length, were found
in coffee-coloured water with a pH of 5, and with a temperature of 12°C at
11h00 at a depth of 25 cm.

Some of these larvae were reared in an aquarium in the open, and the ponds
in the Cape Point Reserve were checked periodically for new spawnings and to
record the development of the tadpoles in natural conditions. Several tadpoles
of various stages were kept in a plastic gauze cage 1m xX 0,5m x 0,5m,
which was fixed between poles at the edge of a pond, leaving the upper edge
10 cm above the water. This facilitated observations on the development of
specific, free-living individuals. In addition, approximately 100 preserved
larvae and young frogs of Xenopus gilli, collected in the same area and fixed in
Lenhossek fluid or alcohol, were used for the study.

For comparison the following material of Xenopus laevis was examined:
preserved larvae and young frogs collected by N. A. H. Millard in the Cape
Flats during the 1940s and by the author in the Cape Flats during 1977; a series
of stage-determined larvae collected by Hubrecht Laboratory, Utrecht, Holland,
at Stellenbosch in 1949-50 and housed in the South African Museum; and live
larvae reared in an aquarium during 1977.

In the following description, which is the object of this paper, the stage
numbers refer to those of the Normal Table of Xenopus laevis laid down by
Nieuwkoop & Faber (1956).

DEVELOPMENT

While there are changes in body-proportions and pigment-distribution
throughout development, free-swimming larvae of Xenopus gilli can be
distinguished from those of Xenopus laevis by their unpigmented longitudinal

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

bands on the dorsal surface of the head (Fig. 1A). These occur, one on either
side, a short distance lateral to the central nervous system (seen by transparency).
The bands embrace the eyes from where they run in an S-shape inwards then
backwards towards the trunk. The two bands together form a lyre shape, which,
even in the water, is very conspicuous, the rest of the body being very dark.
From approximately Stage 45 onwards isolated melanophores begin to appear

4
at:

7st

sient ene
fens Shai Lhe

Racoon
Seaway

er mANCO _
0

mm

ee ewes

V. BRANCO

Fig. 1. A. Xenopus gilli larva, Stage 45-46, dorsal. B. Xenopus laevis larva, Stage 45-46, dorsal.

251

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT

Aa
Sh

WU

Lateral.

Fig. 2. Xenopus gilli larva, Stage 51. A. Dorsal. B

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

B. X. laevis.

illi,

Fig. 3. Photographs of heads of live larvae, Stage 51, dorsal. A. Xenopus

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 253

in the lyre in the thymus gland region. They increase in number during further
development, but never seem to reach the density of the centre portion and the
sides of the head (Figs 2, 3A). The posterior third of the lyre remains
unpigmented until at least Stage 56, while the areas surrounding the eyes can
become somewhat pigmented from Stage 55 onwards (Fig. 5). However, in most
specimens the lyre, although partly obscured, remains distinguishable well into
metamorphosis (Fig. 5).

In contrast, Xenopus laevis larvae from an early stage usually have a more or
less uniform distribution of pigment on the dorsal surface of the head (Fig. 1B),
except sometimes in a position corresponding to the last portion of the lyre,
which may be unpigmented. However, in very young Xenopus laevis larvae
(free-swimming, but before Stage 44) there may be some unpigmented areas on
either side of the central nervous system, which remind one of the lyre of
X. gilli, though the anterior part is quite different (Fig. 6). This condition was
found in ten out of twenty specimens of the same batch and size. But even in
these specimens the posterior portion of the ‘lyre’ usually becomes pigmented
very early and is often indistinct at Stage 46, when the areas between eyes and
central nervous system have usually also become pigmented (Fig. 1B). In the
water the only dark portions which show up are the central nervous system, the
eyes and the abdomen.

The thymus gland, which is situated between eye and ear just below the
surface, and which becomes pigmented in both species at Stage 49, is more
conspicuous in Xenopus laevis, appearing almost like a second pair of eyes
(Figs 3, 4). In Xenopus gilli, this gland is situated just outside the lyre and is
somewhat obscured by the heavy pigmentation in the layers above it.

The pigmentation in the dorsal and ventral tail-fins starts at the hind end
and increases in an anterior direction. In Xenopus gilli the pigmentation of the
ventral fin extends as a narrow band along the edge, and reaches the cloaca at
Stages 55-56 (Fig. 5), while in Xenopus laevis the only fin pigmentation near the
cloaca is an isolated patch which appears at Stage 56 (Nieuwkoop & Faber
1956).

While melanophores appear on the hind-limb buds of Xenopus laevis at
Stage 51, in Xenopus gilli the hind-limb buds are already pigmented at Stage 48.
At the same stage in X. gi/li, and sometimes earlier, a whitish spot becomes
visible above the anterior portion of the central nervous system. In later stages
the ‘Stirnorgan’, in the form of a spherical opaque structure which pushes up
the skin above it, can be observed in this spot (Fig. 5). In Xenopus laevis an
unpigmented spot is said to appear above the ‘Stirnorgan’ at Stage 57. The
‘Stirnorgan’ is developed from the pineal body and lies between the epidermis
and the skull (Nieuwkoop & Faber 1956).

The naso-lachrymal duct is indicated in Xenopus gilli tadpoles at
Stages 55-56 when the pigment begins to part along a line roughly parallel to
the upper lip, from the outer corner of the nostril to where it meets a triangular,
unpigmented area in front of the eye (Fig. 5). This unpigmented line is very

ANNALS OF THE SOUTH AFRICAN MUSEUM

254

Dorsal. B. Lateral.

Xenopus laevis larva, Stage 51. A.

4,

Fig

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 255

Fig. 5. Xenopus gilli larva, Stage 57. A. Dorsal. B. Lateral.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

clear and narrower than the olfactory nerve at Stages 57 and 58, after which
melanophores shift over it again. At Stage 60 it is nearly invisible. A somewhat
irregular protuberance appears at Stage 61 at the point where the nasolachrymal
duct meets the pre-orbital triangle on the lateral wall of the head. The pro-
tuberance shifts in a ventro-caudal direction, coming closer to the eye, and
reaches its final position ventral to the eye, almost in line with the circumorbital
lateral line sensory organs, at Stage 64 (Fig. 7). Two depressions appear on it
which at the end of metamorphosis have become oval apertures, facing in a
slightly caudal direction. Due to its size and position this raised area bearing the

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VP Atpen gn
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Fig. 6. Xenopus laevis larva, pre Stage 44, dorsal.

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 2511

=

Fig. 7. Development of nictitating membrane and external aperture of nasolachrymal duct,
Stages 62, 64, 66 (metamorphosis completed). A-C. Xenopus laevis. D-F. Xenopus gilli.

external openings of the nasolachrymal duct can easily be mistaken for one of
the sensory organs. However, in adult stages the protuberance is usually
flattened out and only two openings remain.

Féske (1934 from Paterson 1939b) believed that the nasolachrymal duct
develops within a very short period from the lower layer of the epidermis. He
was, however, unable to find a developmental stage in support of this view.
It is likely that the parting of the pigment above the future nasolachrymal duct
in X. gilli, as described above, supports Féske’s view.

The nictitating membrance (‘eyelid’) forms parallel with this development.
In Xenopus gilli at Stage 59 the posterior margin of the unpigmented pre-orbital
triangle becomes raised and shows a vertical fold close to the eye. The fold
lengthens during further development and shifts along the ventral margin of the
eye in a caudal direction, absorbs the unpigmented triangle and becomes the
nictitating membrane (Fig. 7).

In Xenopus laevis the development of the ‘eyelid’ is similar. The posterior
end of the nasolachrymal duct is first visible at Stage 62, when a small, indented
semicircular protuberance appears near the anterior tip of the unpigmented
pre-orbital triangle (Fig. 7A). At Stage 64 this protuberance is longer than wide,
oval in cross-section and has two apertures at its tip, separated by a short
septum. It increases in length, comes closer to the eye and, together with the
nictitating membrance, shifts ventrad to the eye, where it reaches its final
position at Stage 65, forming the subocular tentacle (Fig. 7).

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Head and nostril, Stage 59 onwards, dorsal.
A-B. Xenopus laevis. C-D. Xenopus gilli.

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 259

This so-called subocular tentacle, which is not a true tentacle (Paterson
1939a, 1939b) sometimes retains both openings in mature frogs. Often, possibly
through wear, the short septum disappears so that the two openings unite
partially or fully.

Contrary to expectation the nasolachrymal duct in both species has no
contact with the orbital capsule. Paterson (19395) concluded that had there
been movable eyelids the duct would have been directed inwards towards the
eye and its aperture would then have been comparable with the punctum
lacrimale of higher forms. Observation of living adult Xenopus laevis reveals
that the slightly opaque nictitating membrane can move in an upward direction,
over the surface of the eye, to its upper margin. This, however, seems to happen
only when irritation is felt.

At Stages 55-56 of X. gilli the nostril has its outer corner well sunk below
the surface of the head. Its caudal margin begins to develop a cone-like pro-
trusion, roughly where the olfactory nerve meets it. This protrusion shifts
outwards and becomes situated at the outer corner at Stage 59, where it remains.
During these stages the margins of the nostril become raised progressively
towards the outer corner, so that at Stage 59 the inner corner is level with the
head-surface, while the outer is raised well above it. The lip-like raised margins
are now pigmented while the ‘knob’ is unpigmented (Fig. 8C—D). Soon after the
completion of metamorphosis the ‘knobs’ at the outer corners of the nostrils
become pigmented and the darkish belly-dotting appears. At the completion of
metamorphosis the frogs measure approximately 15 mm from snout to vent.

This development of nostril and nasolachrymal duct is much the same as
in Xenopus laevis. However, in the iatter the ‘knob’ is usually pigmented, while
the ‘lips’ are almost unpigmented (Fig. 8A—B).

Thus, while identification of Xenopus gilli tadpoles by the lyre becomes less
easy in advanced stages, the reversed pigment-pattern of the nostril-margins
provides a useful character.

At Stage 60 a light dorsal midline appears on the head and body of Xenopus
gilli, beginning roughly between the eyes and ending roughly between the hind
legs. The typical gi/li-pattern of more or less complete longitudinal dark bands,
two on the back and a weaker, more broken one, on either side, is fully
differentiated at Stage 62 (Fig. 9A).

FURTHER OBSERVATIONS ON LIVING LARVAE

The duration of development from the fertilized egg to the end of meta-
morphosis probably varies. For Xenopus laevis, reared in the laboratory, it is
given as approximately 58 days (Nieuwkoop & Faber 1956). Xenopus gilli
larvae examined here were first seen when at Stage 45-46 (probably one week
old) on 31 July 1977. On 6 November 1977 approximately 15 per cent of the
larvae in the ponds had reached Stage 62. On 20 November some had completed
metamorphosis. The development thus required approximately 120 days. This
much longer period was probably partly due to the low water temperature, which

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. A. Xenopus gilli larva, Stage 62, dorsal.
B. Xenopus laevis larva, Stage 62, dorsal.

THE DEVELOPMENT OF XYENOPUS GILLI ROSE & HEWITT 261

on 6 November 1977, a warm, sunny day, measured only 20°C at 11h00 in
25 cm depth, whereas Nieuwkoop & Faber retained the water temperatures for
their developing larvae at 22—24°C throughout. All larvae observed in three
different ponds in the Cape Point Reserve on 31 July and 7 August 1977 were
of roughly the same size and stage. During subsequent examinations small and
more advanced larvae were encountered together, indicating additional
spawning. However, on 6 November the smallest larvae were in Stage 48-49,
probably indicating that spawning had ceased. In February 1976 Xenopus gilli
larvae from Stage 54 to the end of metamorphosis had been collected in the
biggest pond where the temperature might have remained sufficiently low.
Xenopus gilli larvae show signs of discomfort when the water temperature
reaches 27°C. They remain at the bottom and try to go even deeper. At 31°C
they make sporadic attempts to swim, but balance-disturbance causes them to
tumble, often sinking to the bottom, where they remain motionless, sometimes
lying on the side or back. In contrast, Xenopus laevis larvae still behave normally
at a temperature of 34°C. Only when 36°C is reached, do they show signs of
dying.

During the night the larvae of both species darken considerably. The
tail-fin especially appears practically black when suddenly brought into light.

Larvae reared in an aquarium show retarded growth, although the develop-
ment seems to progress normally, thus producing smaller frogs. Two Xenopus
gilli, which were caught near the end of metamorphosis in February 1976 in the
Cape Point Reserve, were kept indoors thereafter and did not develop the
typical gilli belly-dotting and ochrous undersurfaces of the thighs. The nostril
‘knob’, too, remained unpigmented as in advanced Xenopus gilli larvae. In all
other respects they showed the gi//i characteristics and measured approximately
43 mm from snout to vent in November 1977.

Occasionally, when both species occur together, some specimens appear to
be hybrids. Their size and general shape is that of Xenopus laevis, and so is the
presence of the subocular tentacle just ventral to the eye. Both dorsal and
ventral coloration is that of Xenopus gilli, the longitudinal dark bands being
sometimes more complete than in some specimens of Xenopus gilli.

A specimen (SAM-ZR44317), ccllected by B. Deyer in March 1975 in
the Lotus River near the entry into Zeekoevlei and donated to the museum by
R. Boycott in March 1977, is peculiar in some aspects: its general shape, the
lack of a subocular tentacle, the dense pigmentation of the webbing of the feet
and the dark belly dotting are typical of Xenopus gilli; the teeth in the upper
jaw are, as in Xenopus gilli, long and protrude well beyond the edge of the
mouth; however, the dorsal surface of the animal is of a more or less even,
greyish colour with no pattern; the general colour is lighter than is usual in
Xenopus gilli and the claws on both feet are unpigmented (some unpigmented
claws do occur occasionally in both species). The specimen, with a snout-to-vent
length of 67 mm, exceeds the maximum size given for Xenopus gilli (Poynton
1964).

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

interorbital
distance

internasal
distance
! |

| | eye diameter

Fig. 10. Diagram to illustrate dimensions used to distinguish
Xenopus gilli from Xenopus laevis larvae.

THE DEVELOPMENT OF XENOPUS GILLI ROSE & HEWITT 263

SUMMARY OF DISTINGUISHING FEATURES OF X. GILLI AND
X. LAEVIS LARVAE

It should be noted that the material on which the dimensions are based consists of freshly
preserved specimens in the case of X. gilli, and in X. laevis of both freshly preserved specimens

and specimens which have been preserved for about 30 years.

X. gilli
. Tadpoles very dark with unpigmented lyre.

. Thymus gland and blood-vessels on head-
body not conspicuous.

. Eyes comparatively small; eye diameter/
interorbital distance,

Stage 45—46: 0,20-0,32 X55 0,26;

Stage 50-51: 0,18-0,22 x,; 0,20

(see Fig. 10).

. Nasal capsules comparatively far apart;
internasal distance/eye diameter,

Stage 45-46: 0,83-1,40 x5, 1,04;

Stage 50-51: 0,62-0,90 x,; 0,74

(see Fig. 10).

. Nostril ‘knob’ unpigmented; raised margin
of nostril pigmented.

. Pigmentation of ventral tail-fin reaching
cloaca from Stage 55 onwards.

X. laevis
Tadpoles light with more or less even
pigmentation.
Thymus gland and blood-vessels on head-
body conspicuous.
Eyes comparatively large; eye diameter/
interorbital distance,
Stage 45-46: 0,30-0,54 x,, 0,34;
Stage 50-51: 0,23-0,46 x, 0,33.

Nasal capsules comparatively close
together; internasal distance/eye diameter,
Stage 45-46: 0,43-0,88 xj, 0,63;
Stage 50-51: 0,38—0,80 x,,; 0,53.

Nostril ‘knob’ pigmented; raised margin
of nostril unpigmented.

Pigmentation of ventral tail-fin not
reaching cloaca but forming an isolated

patch near cloaca at Stage 56.

ACKNOWLEDGEMENTS

I wish to express my thanks to the chief warden, Mr G. E. P. Wright, and
the rangers of the Cape Point Nature Reserve for co-operation; to my colleagues,
Mr L. R. Swartz and Mr G. Esau for assistance in the field; to Mr V. Branco
for the drawings made with the aid of a camera lucida; to Mr G. X. Kannemeyer
for the photographing of tadpoles; and to Dr N. A. H. Millard for encouraging
discussion and help with the manuscript.

REFERENCES

Koper, H. R. & Du Pasquier, L. 1975. Production of large clones of histocompatible fully
identical clawed toads (Xenopus). Immunogenetics 2: 87-91.

MERTENS, R. 1970. Uber den Kapkrallenfrosch, Xenopus gilli. Aquar.-u. Terrar.-z. 23 (1):
21-23.

Nieuwkoop, P. D. & FABER, J. eds. 1956. Normal table of Xenopus laevis (Daudin). Utrecht:
Hubrecht Laboratory, Amsterdam: North Holland Publishing Company.

PATERSON, N. F., 1939a. The head of Xenopus laevis. Q. JI microsc. Sci (NS) 81: 161-234.

PaTeErRSON, N. F., 1939b. The olfactory organ and tentacles of Xenopus laevis. S. Afr. J. Sci.
36: 390-404.

Poynton, J. C. 1964. The Amphibia of southern Africa. Ann. Natal Mus. 17: 1-334.

Rose, W. & HewitT, J. 1927. Description of a new species of Xenopus from the Cape Peninsula.
Trans. R. Soc. S. Afr. 14: 343-346.

Wasi, M. R. & Du Pasquier, L. 1976. Antibody patterns in genetically identical frogs.
Nature, Lond: 264: 642.

Wacer, V. A. 1965. The frogs of South Africa. Cape Town, Johannesburg: Purnell & Sons.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-1SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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In describing new species, one specimen must be designated as the holotype; other speci-
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Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
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R. EB, BAU

THE DEVELOPMENT OF XENOPUS GILLI
ROSE & HEWITT (ANURA, PIPIDAE)

— f“°7 Yys™

VOLUME 76 PART 7 SEPTEMBER 1978 ISSN 0303-2515

OF THE SOUTH AFRICAN
MUSE

UM

CAPE TOWN

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

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

KOMN, A. J. 19606. 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.
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(continued inside back cover)

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

Volume 76 Band
September 1978 September
Parte. .) ace

LATE TERTIARY HYAENIDAE FROM
LANGEBAANWEG, SOUTH AFRICA, AND THEIR
MME YANCE TO THE PHYLOGENY OF THE
FAMILY

By

Q. B. HENDEY

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH
AFRICA, AND THEIR RELEVANCE TO THE PHYLOGENY OF THE
FAMILY

By
Q. B. HENDEY
South African Museum, Cape Town

(With 11 figures and 4 tables)
[MS. accepted 11 July 1978]

ABSTRACT

The Hyaenidae are divided on phylogenetic grounds into the ‘Hyaena group’ and the
‘Percrocuta group’, both of which are represented by fossil forms from the latest Miocene/early
Pliocene Varswater Formation at Langebaanweg, South Africa, four species being assigned
to the Hyaena group and one to the Percrocuta group. The latter species is Adcrocuta australis,
while the others are Ictitherium preforfex, Hyaena abronia, Hyaenictitherium namaquense and
an unnamed species of Euryboas. They reflect a pattern of representation established during the
late Miocene, although individual species are more advanced than their Eurasian late Miocene
counterparts, and they fill an important temporal gap in the recorded history of the family.
Chasmaporthetes is regarded as a member of the Percrocuta group and not as a close relative
of Euryboas.

CONTENTS

PAGE

Introduction . : p : =) 2605
The Langebaanweg Hyaenidae ‘ : a 270
Introduction . 3 : ; . 270
The Hyaena group. . : AAS
The Percrocuta group . : ‘ SAD
Nomenclature . : . 280
Relationships of Oyasnanar dere: ‘ » 283
Acknowledgements . ; : ; 1 296
References . 5 ; : : : ~ 296

INTRODUCTION

There are only four hyaenid species still extant and they comprise a
relatively uncommon element in the faunas of Africa and southern Eurasia.
The species are the striped hyaena (Hyaena hyaena), the brown hyaena (Hyaena
brunnea), the spotted hyaena (Crocuta crocuta) and the aardwolf (Proteles
cristatus). The latter is an aberrant insectivorous species, while the others are
well adapted to a scavenging role, although they may also be actively predacious.

The family, which stemmed from the Viverridae during the Miocene, was
formerly more diverse and widespread, and, although hyaenas are rare as
fossils in North America, they are sometimes abundantly represented in late

265

Ann. S. Afr. Mus. 76 (7), 1978: 265-297, 11 figs, 4 tables.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cenozoic deposits in the Old World. In spite of this, opinions on hyaenid inter-
relationships and phylogeny are almost as numerous as papers dealing with
these subjects, some recent examples being Thenius (1966), De Beaumont
(1967), Ficcarelli & Torre (1970), Hendey (1974a), Schmidt-Kittler (1976), and
Galiano & Frailey (1977). There is, however, one important point about which
there does appear to be general agreement. Evidently hyaenids evolved from
viverrids on more than one occasion, and on phylogenetic grounds the family is
divisible into at least two major groups. These two groups have yet to be
accorded formal nomenclatural recognition, and they are here informally
termed the ‘Percrocuta’ and ‘Hyaena’ groups.

The former was the first to be differentiated, having evolved from an as yet
undetermined viverrid ancestor (or ancestors) early in the Miocene. The genera
(or subgenera) constituting this group are Percrocuta, Dinocrocuta and
Adcrocuta (see Schmidt-Kittler 1976). For reasons which will be explained later,
the genus Chasmaporthetes is also included here. Members of the group appear
in Africa and Eurasia as progressive hyaenas late in the middle Miocene. They
may have had their origins in Africa where the Miocene fossil record is com-
paratively poor. Certainly the superior Eurasian record includes no appropriate
links with the Viverridae.

By contrast, the evolution of the Hyaena group is well documented by
Eurasian fossils. The stem genera were the ‘Vindobonian’ Protictitherium and
Miohyaena (Schmidt-Kittler 1976), which were apparently descended from the
late Oligocene Herpestides (De Beaumont 1967). The group radiated during the
late Miocene and a number of small- to medium-sized species occurred together
as a characteristic element in Eurasian ‘Hipparion faunas’ (i.e. Turolian in
Europe and Turolian equivalent in Asia). There is as yet no consensus on the
nomenclature of the late Miocene taxa. The genera recognized here are
Plioviverrops, Ictitherium, Palhyaena, Hyaenictitherium, Hyaenictis, and Lycy-
aena. Numbers of species and subspecies have been named, but the complexities
of the situation will not be explored.

Many of the recorded late Miocene hyaenids of Eurasia were separated
from one another temporally and/or geographically, but there are well-
documented examples of the contemporary occurrence of several species at a
single locality. Pikermi in Greece is perhaps the most extreme case, its fauna
including five members of the Hyaena group (i.e. Plioviverrops orbignyi,
Ictitherium robustum, Palhyaena hipparionum, Hyaenictis graeca, and Lycyaena
chaeretis), and a percrocuta (Adcrocuta eximia) (Pilgrim 1931). A similar
association is evident at several localities in China, where there are three
Hyaena group species (Uctitherium gaudryi, Palhyaena wongii, and Hyaen-
ictitherium hyaenoides), and a percrocuta (a variety of A. eximia) (Zdansky 1924;
Kurtén 1953).

The diversity of hyaenids in Eurasia during the late Miocene is a reflection
of the wealth of the fauna as a whole at that time. It is a period of which Kurtén
(1971: 135) has said ‘may well be regarded as the climax of the entire Age of

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 267

Mammals’. This probably applies in the case of Africa as well, but the late
Miocene fauna of the continent is not well known.

It is, however, becoming increasingly clear that African faunas of the very
late Miocene and early Pliocene were as spectacular in character as the Eurasian
‘Hipparion faunas’, which predate them. The African faunas in question include
those from Mpesida (Bishop et al. 1971), Lothagam 1 (Smart 1976), Lukeino
(Pickford 1975), Sahabi (Petrocchi 1952), ‘E’ Quarry at Langebaanweg (Hendey
1976) and Kanapoi (Behrensmeyer 1976). Smart (1976) has already pointed
out that climatic and environmental changes, which adversely affected Eurasian
faunas towards the end of the Miocene, had less of an impact in Africa. Indeed,
they may have had the effect of increasing mammalian migration from Eurasia
to Africa.

In spite of the improving situation in Africa, the very late Miocene/early
Pliocene fossil record in the Old World is generally poor. This was a period
for which recorded fossil occurrences are few in number, widely dispersed and
have often yielded faunas of limited size. There is a considerable improvement
in the record throughout the Old World during the late Pliocene, a tendency
which continues into the succeeding Pleistocene epoch.

The Old World late Tertiary record is thus characterized by both temporal
and geographical irregularities, which complicate interpretations of the origins
and history of mammals that lived during this period. The problem is particu-
larly acute in the case of those mammalian groups, such as the Hyaenidae,
which were both diverse and widespread. Consequently any fossil occurrence
which dates from a poorly known part of the record, and which produces good
samples of material, could make a significant contribution towards the under-
standing of the history of such groups.

‘E’ Quarry at Langebaanweg is an occurrence of this kind. Deposits
exposed in this quarry are comprised largely of the latest Miocene/early Pliocene
Varswater Formation, which has produced a fossil assemblage of size and
diversity unequalled by contemporary occurrences elsewhere in Africa (Hendey
1976). One of the more remarkable features of this fauna is the number and
variety of carnivores which are represented (Hendey 1974a, 1976, 1977), and,
of the terrestrial species, Hyaenidae occur most commonly. At least five hyaenid
species are represented, which is in marked contrast to the situation at other
late Miocene/early Pliocene localities in Africa (Table 1).

Before proceeding to an account of the hyaenids, a few further observations
on the ‘E’ Quarry fauna and deposits are necessary to place them in perspective.

Evidence which indicates an early Pliocene age for the Varswater Formation
was recently reviewed (Hendey 1978: 2). The dating of this formation is based
on comparisons between certain of its taxa with their counterparts in the east
African sequence, for which there are some radiometric dates. The suggested
age limits of 4 to 5 million years (m.y.) for the Varswater Formation are,
however, not securely established. The 4 m.y. limit has already been questioned
(Hendey 1978), and there is now reason to believe that the older limit may have

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE |

Late Miocene/early Pliocene Hyaenidae of Africa.

LOCALITY AGE TAXA REFERENCES
Kanapoi early Pliocene Hyaena sp. Behrensmeyer 1976
Langebaanweg latest Miocene/ Ictitherium preforfex This report

(‘E’ Quarry) early Pliocene Hyaena abronia

Hyaenictitherium namaquense
Euryboas sp.
Adcrocuta australis

Sahabi late Miocene or Hyaenidae gen. et sp. indet. —

early Pliocene
Lukeino late Miocene cf. Crocuta sp. Pickford 1975
Lothagam 1 late Miocene aff. Euryboas sp. Smart 1976

been underestimated. This matter will be dealt with in detail elsewhere, but in
the meantime the former practice of referring to the age of the Varswater
Formation and its fossils as ‘early Pliocene’ is here replaced by the reference,
‘latest Miocene/early Pliocene’.

It is nevertheless perfectly clear that the Varswater Formation fauna as a
whole postdates the classic Eurasian late Miocene faunas such as those from
Pikermi and lower Samos, which are between 7 and 10 m.y. old, but predates
the Old World late Pliocene/early Pleistocene faunas, which are younger than
3,5 m.y. In other words, it dates from a period towards the end of the Tertiary
when the Old World fossil record is generally poor.

Another important point about the Langebaanweg (CE Quarry) fauna
relevant to the present study is that it comes from the most southerly part of
Africa and is thus geographically far removed from those faunas with which it
is here compared and contrasted. It is therefore possible that the ‘E’ Quarry
fauna, like that of the near-by Baard’s Quarry, includes regional variants of
more widespread taxa, endemic taxa and even late surviving members of
lineages which were elsewhere extinct (Hendey 1978). If there are regional
peculiarities in the fauna and these are not recognized, this could result in
incorrect interpretations of the relationships of such taxa. Furthermore, the
populations to which the Langebaanweg species belonged were probably not
directly ancestral to later ones elsewhere.

The Cenozoic terrestrial faunas of the most southerly parts of Africa may
characteristically have contributed little in the way of emigrants to regions
further north. Instead the composition of the local faunas is likely to have been
changed largely by immigration from the north and some endemic speciation,
while local extinctions were not necessarily coincident with those elsewhere.
The region may thus be viewed as a terminus in a zoogeographic sense. Neverthe-
less, local populations may have exhibited all the characteristics of their more
northerly counterparts from which subsequent populations did evolve. Thus,

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 269

when Langebaanweg species are suggested as ancestors of later ones, they were

themselves not directly ancestral, but simply represent the kinds of animals

from which later ones were derived (i.e. ‘structural ancestors’).

The observations in the preceding paragraphs are conveniently summarized
if the Langebaanweg (‘E’ Quarry) fauna is visualized as a younger southern
African counterpart of the classic “Hipparion faunas’ of Eurasia. It differs from
such faunas because it is separated from them by the width of a continent and
a few million years in time. In spite of this, its composition is of an essentially
similar pattern. The ‘E’ Quarry fossil assemblage, with its eighty mammalian
species, provides the best evidence yet that the Old World late Tertiary ‘climax
of the entire Age of Mammals’ persisted over much, if not all, of Africa long
after it had passed in Eurasia.

A final point about the ‘E’ Quarry fauna relevant to the present report is
that it includes species from a variety of habitats (Hendey 1976: 222-230).
As a general rule terrestrial carnivores such as hyaenids are not as tied to
particular habitats as are, for example, herbivores such as bovids. However,
the contemporary occurrence at Langebaanweg of several closely related
hyaenids suggests that the species concerned did occupy different habitats.
A similar association of three hyaenid species in China during the late Miocene
led to the suggestion by Kurtén (1953: 45) that one was a ‘steppe form’, one a
‘forest form’, and the third an ‘intermediate’.

The situation at Langebaanweg, and indeed elsewhere, cannot be completely
explained by habitat preference, since there were more hyaenid species than
major habitat types. The co-existence of more than one species in a given
environment is made possible by different behaviour patterns in the species
concerned. This no doubt applies in the case of certain of the Langebaanweg
hyaenids, but the habitat preference factor is almost certainly significant as well,
since certain species occur commonly in, or are restricted to, deposits of
particular kinds (e.g. river channel, floodplain). It is thus possible that while
some species lived in the immediate vicinity of what was then a river estuary,
others lived further inland and the remains of individuals were transported to
the area of deposition by the river. While it is not possible to relate individual
species to major habitat types (e.g. riverine woodlands, open plains), it is
nevertheless significant that elements of the faunas from such habitats were
incorporated in the Varswater Formation.

To sum up, the ‘E’ Quarry fossil occurrences are a potentially important
source of information on Hyaenidae for the following reasons:

1. the fauna dates from a period which has a comparatively poor fossil record
in the Old World;

2. the occurrences are situated on a continent for which the whole late Tertiary
record is poor, but which may well have played a more important role in
hyaenid evolution than has hitherto been supposed;

3. the sample sizes of individual taxa are reasonably good;

4. elements from more than one major habitat type are included in the fauna.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE LANGEBAANWEG HYAENIDAE

INTRODUCTION

Five hyaenid species were described in a recent study of material from
‘E’? Quarry (Hendey 1974a). They were identified as follows: Hyaenictis
preforfex, Hyaena abronia, Hyaena sp. B, Hyaena sp. E, and Percrocuta australis.

Hyaena sp. E is known only from a mandible fragment of an immature
individual and, since its status is uncertain, it is excluded from the discussions
which follow. Each of the remaining four species is known from cranial remains
of three or more individuals and incomplete postcranial skeletons of at least
two individuals. In addition, there is one undescribed species, a Euryboas,
which is represented by the cranial remains of several individuals and an
incomplete postcranial skeleton of one individual. The present situation in
respect of individual species is now reviewed.

The smallest of the Langebaanweg hyaenids, that identified previously as
Hyaenictis preforfex, has proved to be a problematical species. The type
specimen is the damaged skull and incomplete postcranial skeleton of an aged
individual (Hendey 1974a) from bed 3aS of the Pelletal Phosphorite Member
(Hendey 1976: 226-230, 1978: 3). Additional specimens assigned to this species
have since been found in bed 3aN of the same member and, although of
comparable size, the new specimens differ from the holotype in certain dental
characters. The differences are here regarded as more apparent than real and
are ascribed to the aged condition of the holotype. Certain of the characteristics
of the species mentioned in this report are evident in the bed 3aN sample but
not in the holotype. They include a prominent M, metaconid, a feature which
suggests that the genus concerned was not Hyaenictis. The reassessment of this
species also suggests that it may not have been ancestral to the Transvaal
‘Hyaenictis’ forfex as indicated earlier.

The additional ‘Hyaenictis’ preforfex specimens show that this species was
more similar to the second of the Langebaanweg hyaenids, Hyaena abronia,
than had previously been supposed. Specimens which are unequivocally assigned
to H. abronia are from the Quartzose Sand Member, which underlies the
Pelletal Phosphorite Member, and are from floodplain deposits, whereas all the
‘H. preforfex specimens are from river channel deposits. H. abronia is perhaps
the least problematical of the Langebaanweg hyaenids in terms of its status and
relationships.

The third, and next largest, of the species is the unnamed Hyaena sp. B,
which is known only from the floodplain deposits of the Quartzose Sand
Member. At least two additional individuals of this species are now represented
and there is no longer any doubt that it is distinct from H. abronia. The possibility
that species B and the poorly known ‘Hyaena’ namaquensis from Kleinzee
(Stromer 1931) were closely related was mentioned earlier (Hendey 1974a: 147).
The additional species B specimens have made a close relationship seem more
than just possible. The only observable difference between ‘H.’ namaquensis and

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA D/A

species B is that the M, of the former is slightly longer and narrower. In spite

of this, species B is here identified with ‘A.’ namaquensis since the small M, size

difference is outweighed by the general size similarity, comparable tooth
morphology, the proximity of the Langebaanweg and Kleinzee occurrences and
the likelihood that they are broadly contemporaneous.

Many of the specimens belonging to the three species already mentioned
were excluded from the present study because the general similarity in the
characteristics of the species creates the potential for incorrect identification.
Only the best preserved and most complete specimens, that is, those which
undoubtedly belong to the species concerned, were taken into account. In
addition, each sample was limited to specimens from a single stratigraphic unit.
The specimens examined were as follows:

‘Hyaenictis’ preforfex—7 individuals from bed 3aN of the Pelletal Phosphorite
Member (SAM-—PQ-L33046, L31028, L31333, L32893, L33520, L33842,
L34778).

Hyaena abronia—4 individuals from the Quartzose Sand Member (L14186,
L20984, L21009, L22202).

‘Hyaena namaquensis—3 individuals from the Quartzose Sand Member
(L12848, L21008, L25026).

These three species are clearly closely related members of the Hyaena
group and are distinguished from one another principally on the basis of size.
They are characterized by the presence of P, and M3, although M? is sometimes
absent in H. abronia, and are generally similar in terms of tooth morphology
(Figs 1-2).

The fourth species is the undescribed Euryboas. It is similar in overall size
to ‘H.’ namaquensis, but is distinguished by the absence of P,, the occasional
absence of M,, shorter M!, reduction or loss of the M, metaconid, smaller and
simpler M, talonid and longer but narrower Bs to 2 and M, (Fig. 6, Table 2).
The Langebaanweg Euryboas, which is apparently the earliest and most
primitive known member of the genus, will be fully described elsewhere.
Remains of only two individuals were taken into account in the present study
(L21000, L21788). Both are from deposits in the Quartzose Sand Member
which were probably laid down close to, or even in, a river channel. Other
specimens probably belonging to this species are from river channel deposits of
the Pelletal Phosphorite Member.

The fifth and largest of the ‘E’ Quarry hyaenids is a member of the
Percrocuta group and, although described as a Percrocuta, it is now referred to
Adcrocuta (i.e. A. australis). This is done because a close relative and possible
ancestor, the Eurasian ‘Percrocuta’ eximia, is now generally regarded as an
Adcrocuta (Ficcarelli & Torre 1970; Schmidt-Kittler 1976). A. australis has
some characters in common with the Euryboas, but is distinguished by its
larger size, lower crowned and stouter canines, occasional presence of anterior
accessory cusps on P, and Ps, longer carnassials, and unicuspid M, talonid.
The A. australis sample used in this study was derived from the Quartzose Sand

Di2 ANNALS OF THE SOUTH AFRICAN MUSEUM

ST WNIT

CN

ht
4 215

mn I

unl
3 2

An A

ui

mn

MN UNL IN

Fig. 1. Lateral views of hyaenid hemimandibles from Langebaanweg. A. “Hyaenictis’
preforfex (L33046). B. Hyaena abronia (L14186) (reversed). C. ‘Hyaena’ namaquensis
(L25026).

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 213

Cs:
qe

HHI

ui

nA a

2}

i

i

Fig. 2. Occlusal views of hyaenid hemimandibles from Langebaanweg. A. ‘Hyaenictis’
preforfex (L33046). B. Hyaena abronia (L14186) (reversed). C. ‘Hyaena’ namaquensis (25026).

Member, but it is also known from the Pelletal Phosphorite Member and is
apparently not confined to deposits of a particular type.

The ‘E’ Quarry hyaenids appear to be unique in providing evidence of a
progression in an evolutionary sense from Eurasian late Miocene hyaenids,
while at the same time maintaining the pattern of their representation. In other
words, there is apparently no other Old World fauna of comparable age which
includes a variety of hyaenid species which are reminiscent of the Eurasian late
Miocene. In order to justify this observation recorded Eurasian late Miocene
taxa are compared and contrasted with those from Langebaanweg.

THE HYAENA GROUP

Plioviverrops orbignyi is the smallest of the Eurasian species and one which
evidently does not have a counterpart at Langebaanweg. This species may have
become extinct without issue, although Thenius (1966) suggested it as a possible
ancestor of Proteles cristatus, a species whose fossil history is largely unknown
except for a Pleistocene species from the Transvaal (Hendey 19745).

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

The European Ictitherium robustum and Chinese I. gaudryi have long been
recognized as closely related forms and they may be conspecific. The situation
in respect of Palhyaena hipparionum and P. wongii is similar. By contrast, the
Chinese Hyaenictitherium hyaenoides apparently had no European counterpart,
but together with [ctitherium and Palhyaena is part of a close-knit combination
within the Hyaena group which has been thoroughly examined by Kurtén
(1954), amongst others.

Schmidt-Kittler (1976) cast some doubt on the status of Hyaenictitherium,
claiming that it does not even merit separate subgeneric status. H. hyaenoides
is nevertheless distinguishable from the classic Palhyaena, the taxon which
Schmidt-Kittler regards as the stem form. The classification of taxa in such
situations is prone to be controversial and Schmidt-Kittler’s ‘lumping’ is
justifiable. However, the evolution of this group of hyaenids is here interpreted
as successive branching from a primary lineage, with each additional branch
representing a new genus (Fig. 3).

This arrangement can be justified only if later, well differentiated repre-
sentatives of individual branches are recognized. Thus the earliest member of
a new lineage, although essentially similar to the stem form, is distinguished at
the genus level if it can be established that it had descendants whose charac-
teristics are clearly different from those of members of the stem lineage. Certain
of the Langebaanweg hyaenids are relevant to the recognition here of Ictitherium,
Palhyaena and Hyaenictitherium as distinct genera.

The ‘Hyaenictis’ preforfex/Hyaena abronia/*Hyaena namaquensis combina-
tion at Langebaanweg is reminiscent of the Eurasian trio referred to above. In

Ictitherium Palhyaena Hyaenictitherium

Protictitherium

Fig. 3. Suggested phylogenetic relationships of some late Tertiary members of the Hyaena
group.

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 213

each case size differences of no great magnitude are combined with differences
in tooth proportions to distinguish associated species of the two combinations
of taxa. In general, the Langebaanweg species are more advanced than those
of the Eurasian late Miocene. F or example, in the Langebaanweg species the
molars (i.e. M*, M, talonid, M,) are reduced relative to those of the Eurasian
species. In addition, the species of the Langebaanweg series are larger than those
of the Eurasian series, that is, ‘H.’ preforfex is larger than Ictitherium, H. abronia
is larger than Palhyaena and ‘H.’ namaquensis is larger than Hyaenictitherium.

The simplest phylogenetic interpretation of the preceding observations
would be to regard Ictitherium as the ancestor of ‘H.’ preforfex, Palhyaena the
ancestor of H. abronia, and Hyaenictitherium the ancestor of ‘H.’ namaquensis,
with the ancestral forms possibly being African counterparts of the recorded
Eurasian species.

Having established this as a working hypothesis, the suggested inter-
relationships can be examined in more detail. Superficially there is nothing
which would positively preclude the suggested relationships. Indeed they seem
eminently feasible. For example, one of the more striking features of the two
series of taxa is that in each case it is the smallest member which is the most
primitive (i.e. viverrid-like).

In a study of the Eurasian taxa Kurtén (1954: 16-17, Fig. 9) used a ratio
diagram of certain tooth lengths to illustrate similarities and differences.
A similar ratio diagram for the Langebaanweg species is equally revealing
(Fig. 4). Kurtén used Palhyaena wongii as a standard and in the case of the
Langebaanweg series Hyaena abronia was selected since it is the suggested
descendant of Palhyaena. It is worth noting that a ratio diagram using tooth
breadths revealed an essentially similar pattern.

There are some remarkable similarities between the ratio diagrams of the
Langebaanweg and Eurasian series. In their proportions the teeth of ‘Hyaenictus
preforfex and “Hyaena’ namaquensis differ from the Hyaena abronia standard
in much the same way as Ictitherium robustum and Hyaenictitherium hyaenoides
differ from Palhyaena wongii. Interestingly, a much closer approach to the
‘H. preforfex graph is achieved when the J. robustum and I. gaudryi samples are
combined by drawing a new graph on Kurtén’s Figure 9 equidistant from the
graphs of the two Ictitherium species. This may not be a valid statistical pro-
cedure since the unpublished primary data for the two species should be
combined in order to calculate accurate ratios for the combination. Nevertheless,
the experiment must have produced a graph which is approximately correct.

The similarity between the ratio diagrams of the Langebaanweg and
Eurasian samples strongly suggests that the three Langebaanweg species are
interrelated in a manner which is comparable to the interrelationships between
the Eurasian Ictitherium, Palhyaena and Hyaenictitherium. This is here inter-
preted as evidence in support of the ancestor/descendant relationships postulated
earlier.

Not surprisingly the two ratio diagrams also differ in certain respects. For

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

90 110 120 a
|

X -Py

“Pa

“Py

X -p4

Fig. 4. Ratio diagram comparing mean lengths of

lower teeth and P* in certain Langebaanweg

Hyaenidae: ‘Hyaenictis’ preforfex (x), ‘Hyaena’

namaquensis (/\). Standard of comparison (100%):
Hyaena abronia (0).

example, in terms of its tooth proportions ‘H.’ preforfex is closer to H. abronia
than are either of the Eurasian Jctitherium species to P. wongii. This applies
particularly in the case of the posterior premolars (Ps, P,). This is also illustrated
by a second ratio diagram (Fig. 5), in which the lower cheektooth lengths of
the Langebaanweg species are plotted against an J. robustum sample as standard.

Figure 5 illustrates even more clearly than Figure 4 that the tooth propor-
tions of ‘H.’ preforfex and ‘H.’ namaquensis are essentially similar and that
H. abronia differs from them principally in its premolar development. An increase
in premolar size is one of the more general rules in hyaenid evolution and their
marked size increase. in ‘H.’ preforfex and ‘H. namaquensis relative to the
condition in J. robustum is not surprising, since the latter species is a generalized
form which presumably resembles the common ancestor of this group of
hyaenids. Evidently the situation in respect of premolar development in
H. abronia was somewhat different. A possible explanation for this situation
emerges from an examination of the hypothetical lineage which includes
H. abronia.

This lineage, which has Jctitherium as the stem form and with Palhyaena,
H. abronia and the living Hyaena hyaena as subsequent members (Hendey 1974a,
this report), may well prove to be less controversial than others suggested here.

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 277

There is little difference in overall body size and tooth characters in later
members of the lineage, that is, H. abronia through to living H. hyaena. This
suggests that the evolution of at least some characters had virtually ceased by
the time an H. abronia-like stage had been reached. Since some recorded
Palhyaena specimens seem to be little different from the Langebaanweg
H. abronia, the lineage had evidently reached an ‘optimum’ stage relatively
early. In other words, the ‘Hyaena’ condition was achieved precociously.

Evidently the same did not apply to the Ictitherium—‘H.’ preforfex and
Hyaenictitherium—H.’ namaquensis lineages, where development of characters
such as premolar size continued. They were thus slower in reaching an ‘optimum’
condition.

Another important point which is clearly illustrated by the accompanying
ratio diagrams concerns the development of the canines. Kurtén (1954: 16)
found Hyaenictitherium hyaenoides to be ‘fairly similar’ in tooth proportions to
Palhyaena ‘with the exception of the powerful canines’. This is also the character
which most readily distinguishes the Langebaanweg ‘H.’ namaquensis from its
two contemporary near relatives. The implication is that H. hyaenoides and
‘H. namaquensis had the development of large canines as a character in common
and, consequently, that they were, indeed, closely related.

There is, however, a complication with this interpretation of the data.
Judging from Figure 5, canine size in the Langebaanweg species is approxi-
mately proportional to the overall size of the species concerned, something
which is also evident, although perhaps less obvious, in Kurtén’s (1954) ratio

100 110

Fig. 5. Ratio diagram comparing mean lengths of lower teeth in certain Langebaanweg
Hyaenidae: ‘Hyaenictis’ preforfex (x), Hyaena abronia (©), “Hyaena’ namaquensis (/\).
Standard of comparison (100%): European Ictitherium robustum (Kurtén 1954).

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

diagram. It is therefore possible that the large canine size in H. hyaenoides and
‘H.’ namaquensis is due not to a phylogenetic connection, but simply to the fact
that they are the largest members of their respective series. This means that
canine size, and indeed the overall similarities in tooth proportions in the two
series, may be coincidental.

Coincidence can, of course, not be ruled out, but it must be highly improb-
able. Not only do the two series have similar patterns in tooth proportions
which go together with evolutionary advances in tooth morphology (e.g. reduced
molar size in the Langebaanweg series), but in the case of the Palhyaena-—
H. abronia \ineage at least, there is evidence of advances in the postcranial
skeleton as well (Hendey 1974a: 116, table 20). Furthermore, if the Langebaan-
weg series did not evolve from the suggested late Miocene taxa, alternatives
must be sought and certainly none are obvious.

Whereas the evolution of H. hyaena from the Palhyaena—H. abronia
combination appears likely, the subsequent histories of the JIctitherium-—
‘H. preforfex and Hyaenictitherium—H.’ namaquensis lineages are obscure. The
former almost certainly has no living descendant, but the latter could be
ancestral to Hyaena brunnea.

The first fully mature ‘H.’ namaquensis specimen from Langebaanweg
(SAM-—PQ-L25026) is similar in overall size to living H. brunnea. The fossil
species differs in some body proportions such as, for example, having longer
hind limbs. This difference is similar to one which distinguishes H. abronia from
H. hyaena (Hendey 1974a: 116-118). There are also some marked differences
in the dentitions of ‘H. namaquensis and H. brunnea. For example, the former
retains P, and M, and has lower crowned and more slender premolars. These,
and other differences, all indicate the primitive state of ‘H.’ namaquensis and
all are likely to have been present in an early ancestor of H. brunnea. Conse-
quently, there is apparently nothing to preclude an ancestor/descendant
relationship between these taxa.

The situation is reminiscent of that which exists between H. abronia and
H. hyaena, although in this instance the differences are less marked and there
is an appropriate late Pliocene/early Pleistocene intermediate form recorded
(i.e. H. hyaena makapani). It was suggested above that the hypothetical
Hyaenictitherium—H.’ namaquensis lineage had apparently not reached an
‘optimum’ evolutionary state by the early Pliocene. If evolutionary advances
continued during this epoch, they may well have been in the direction of an
H. brunnea-like species. This would account for the fact that H. brunnea is the
more specialized (advanced) of the two living Hyaena species. While a direct
relationship between ‘H.’ namaquensis and H. brunnea is here suggested as a
possibility, more evidence is needed to test the hypothesis.

Whereas Ictitherium, Palhyaena and Hyaenictitherium form a close-knit
unit within the late Miocene Hyaena group of Eurasia, the group apparently
has at least two additional members, namely, Hyaenictis and Lycyaena.

For a reason given earlier, the Langebaanweg ‘Hyaenictis’ preforfex is no

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 279

longer regarded as a member of that genus. There is, however, another of the
Langebaanweg species which may be related to Hyaenictis or Lycyaena. This
is the undescribed species of Euryboas.

The late Miocene ancestor of the Langebaanweg Euryboas is likely to have
had the following amongst its characteristics:

1. M? reduced or absent.

2. M, metaconid present, although possibly smaller than in IJctitherium,

Palhyaena and Hyaenictitherium;

M, talonid bicuspid;

M, persistently present, but P,; absent or sometimes absent;

5. Postcranial skeleton similar in overall size to larger members of the
Ictitherium/Palhyaena/Hyaenictitherium subgroup.

=

The last character is important in suggesting that Euryboas belongs to the
Hyaena group rather than the Percrocuta group, since the late Miocene
percrocutas were evidently relatively large, heavily-built animals. The other
characters combine to suggest that Euryboas was derived from Hyaenictis or
Lycyaena, rather than the Ictitherium/Palhyaena/Hyaenictitherium subgroup.
Both Lycyaena (De Beaumont 1967) and Hyaenictis (Thenius 1966) have
previously been suggested as possible Euryboas ancestors.

In the case of Hyaenictis, H. graeca is the species of appropriate age to be
ancestral to a primitive Euryboas. This species apparently fulfils most of the
required criteria for this role, but it does lack the M, metaconid. This is not
necessarily a serious objection since this cusp is sometimes variably developed
in hyaenid species (Kurtén 1956: 12-14), and the M, of A. graeca, like that of
the Langebaanweg Euryboas, may sometimes have had a small metaconid.
Alternatively, this cusp may have been present in an as yet unknown African
counterpart of H. graeca.

The Eurasian late Miocene representative of Lycyaena is L. chaeretis, for
which a similar complication exists. According to Pilgrim (1932) this species
lacks Mz, a tooth which is sometimes present in the Langebaanweg Euryboas.
Zapfe (1948) has, however, recorded a L. chaeretis from Austria in which a small
M. was present. The presence of M, may have characterized certain late
Miocene populations of Lycyaena, including an African one, if it existed.

On balance the other recorded characteristics of Lycyaena appear to be
more Euryboas-like than those of Hyaenictis and the former is here regarded as
the likely ancestor of Euryboas.

Later species of Euryboas are recorded from elsewhere in Africa and in
Europe. Some of the specimens previously included in this genus were recently
referred instead to Chasmaporthetes (Galiano & Frailey 1977).

THE PERCROCUTA GROUP

The last of the ‘E’ Quarry hyaenids, Adcrocuta australis, was once thought
to be the least problematical of the species, since its large size and certain
specialized dental characters readily distinguished it from its contemporaries.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

The increase in the sample size has revealed that there is appreciable variation
in this species, which may be due at least in part to sexual dimorphism, and
some cranial material apparently belonging to A. australis resembles specimens
assigned to Euryboas.

Even when allowance is made for marked sexual dimorphism, and
problematical fragmentary specimens are excluded from consideration, the
material assigned to A. australis is more variable than that belonging to other
Langebaanweg hyaenids. Indeed, there are grounds for suspecting that the
material may belong to two species. An essentially similar situation was
encountered by Zdansky (1924) when he studied the Chinese A. eximia variabilis.
His decision to recognize only one variable species has been generally supported
(e.g. Pilgrim 1931; Kurtén 1957). With this precedent in mind, and since
A. eximia is regarded as the structural ancestor of A. australis (Hendey 1974a),
the material assigned to the latter is left undivided.

The Langebaanweg A. australis and Euryboas have certain dental characters
in common. For example, their cheekteeth are generally similar in morphology
(Fig. 6) and in proportions (Fig. 7). These two species are readily distinguished
from the other Langebaanweg hyaenids, and this raises the possibility that the
Euryboas belongs in the Percrocuta, rather than Hyaena group. Although this
possibility cannot be dismissed, it is not favoured since A. australis does have
some specialized characters not evident in Euryboas, which suggests that the
two species belong to lineages which had had a long, separate history. For
example, A. australis was a large animal with robustly proportioned postcranial
bones, whereas the Euryboas was smaller and more lightly built. A. australis is
also distinguished by specialized dental characters such as sometimes having
prominent anterior accessory cusps on P, and P, and in having a unicuspid M,
talonid. |

The similarities between A. australis and the Euryboas, which are here
ascribed to parallel evolution, will be discussed again in a later section of this
report.

NOMENCLATURE

Having reviewed the status and relationships of the ‘E’ Quarry hyaenids,
the taxonomic implications of the conclusions reached here can be considered.

Since ‘Hyaenictis’ preforfex is regarded as a descendant of late Miocene
Ictitherium, and since the subsequent history of the lineage is not known, the
Langebaanweg species is referred to Ictitherium on the principle that members
of a single lineage are congeneric.

The situation in respect of Hyaena abronia remains unchanged, although a
new complication now arises. In this instance both ancestral (Palhyaena) and
descendant (H. hyaena) forms are recognized and, in order to conform to the
principle stated above, Palhyaena should be sunk into Hyaena, the latter being
the name which has priority. This step is, however, not formally proposed since
there is no point in synonymizing a name in common use if the reason for doing

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 281

eA

Mn

i

a
3 + sf 6i 7 8 9 Hy

Fig. 6. Lateral and occlusal views of hyaenid hemimandibles from Langebaanweg.
A. Euryboas sp. (L21000) Gmmature adult). B. Adcrocuta australis (L22204).

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

100 110 120 z 130 140 -p2
i %, _p3

x .p4

© ee ‘My

a
~

Fig. 7. Ratio diagram comparing mean lengths of upper
and lower teeth of Langebaanweg Euryboas sp. (©) and

Adcrocuta australis (x). Standard of comparison (100%):
Hyaena hyaena.

X “Py

so is not generally accepted. A decision can await reactions to the suggestion.

‘Hyaena namaquensis, like Ictitherium preforfex, is referred to its supposed
ancestral genus, in this instance, Hyaenictitherium. The name becomes Hyaen-
ictitherium namaquense. If substantiated, the suggestion that H. namaquense is
ancestral to Hyaena brunnea will provide a solution to the problem of the
generic name of the latter. At least twice in recent years it has been indicated
that the relationship between H. hyaena and H. brunnea is distant enough to
warrant nomenclatural distinction above the species level (Hendey 1974a:
148-149; Galiano & Frailey 1977: 11-12). Hyaenictitherium probably has
priority over any other generic name available for ‘Hyaena’ brunnea.

In spite of the problems and doubts about relationships mentioned earlier,
no name changes are proposed in the cases of Adcrocuta australis and the
unnamed Euryboas species.

The Langebaanweg (‘E’ Quarry) hyaenids now recognized are listed in
Table 1. The suggested relationships of the taxa already discussed are
summarized in Figure 8. In all instances the ancestral forms are merely recorded

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 283

species in an appropriate evolutionary state for this role and they need not
have been in the actual ancestral position. For example, the Chinese Hyaen-
ictitherium hyaenoides was almost certainly not a direct ancestor, but was
simply a structural ancestor of the South African H. namaquense. It is much
more likely that the latter stemmed from an as yet unknown African late
Miocene Hyaenictitherium which resembled H. hyaenoides, but which was not
necessarily conspecific with it.

The tooth sizes of the samples used in this study are recorded in Table 2.

RELATIONSHIPS OF CHASMAPORTHETES

In a recent note on the origins of North American Chasmaporthetes, the
Chinese late Miocene Adcrocuta eximia variabilis was suggested as a likely
ancestor (Hendey 1975). In recording what they regard as the first Asiatic
Chasmaporthetes (C. kani), Galiano & Frailey (1977: 9) concluded that this
genus “was apparently generically distinct before its entry into North America
and shares a number of derived characters with Euryboas rather than with
Percrocuta’. This conclusion was based on a cladistic analysis of the Hyaenidae
other than the percrocutas.

The omission of the percrocutas was justified on the grounds that they
retain ‘a number of primitive characters’ in association with others that are
‘highly derived’, which sets them apart from other hyaenids, including
Chasmaporthetes (Galiano & Frailey 1977: 9). The derived characters mentioned
are a reduced P* protocone and contact between the premaxillary and frontal

HYAENA GROUP PERCROCUTA GROUP

‘Hyaena’
brunnea
f

Hyaena
hyaena

PLEISTOCENE
AND
HOLOCENE

Euryboas
bielawskyi

PLIOCENE

Adcrocuta
australis*
1
i}

1
Hyaenictitherium Hyaena Ictitherium Euryboas
namaquense* abronia* preforfex* sp. is

PLIOCENE

' t
Hyaenictitherium Palhyaena Ictitherium Lycyaena

hyaenoides hipparionum robustum chaeretis

Om eemaamat

Protictitherium Miohyaena

i]
Adcrocuta
eximia

MIOCENE

*Langebaanweg species Herpestides

Fig. 8. Tentative phylogeny of some Hyaenidae.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 285

bones of the skull, while the only primitive character mentioned was the presence
of a large metacarpal I. While the omission of most percrocutas on these grounds
may be justified, this does not apply in the case of A. eximia variabilis.

A large metacarpal I, which amongst living taxa is found only in Proteles,
undoubtedly is a ‘primitive’ character, but it was a feature of most, if not all,
late Tertiary hyaenids. This certainly applies in the case of all five species
recorded from ‘E’ Quarry.

Although the P* protocone generally is reduced in the percrocutas (Kurtén
1957), A. eximia variabilis is exceptional in having this cusp of variable size
(Zdansky 1924: 96; Pilgrim 1931: 117; Kurtén 1957: 399). Evidently in the
case of certain individuals at least, the P* protocone was not necessarily any
smaller than that in C. kani, in which this cusp is also somewhat reduced
(Galiano & Frailey 1977: fig. 1).

The contact between the premaxillaries and frontals is not a constant
character in A. eximia variabilis, Zdansky (1924) having recorded one specimen
in the series he studied in which there was no such contact. This character is
also variable in some other hyaenids. For example, a series of 13 Crocuta
crocuta skulls in the South African Museum includes 7 specimens in which
there is no contact between these bones, | specimen in which there is contact
on one side only, and 5 specimens in which there is contact. Galiano & Frailey
were thus mistaken both in regarding the maxillary-frontal contact in A. eximia
variabilis as constant, and in regarding it as unique.

Since there are now no recorded grounds for dismissing the possible
phylogenetic link between A. eximia variabilis and Chasmaporthetes, it is here
examined in some detail.

North American Chasmaporthetes undoubtedly had an Asian ancestor,
and Galiano & Frailey (1977) have provided evidence that the Chinese C. kani
is an appropriate candidate for this role, at least in terms of morphology.
Consequently the theory that Chasmaporthetes and Adcrocuta are closely
related can most conveniently be tested by comparing C. kani with the Chinese
A. eximia variabilis described by Zdansky (1924). The comparison is facilitated
by the fact that both species are represented by fairly complete and well-
preserved specimens belonging to several individuals.

In the comparisons which follow account is also taken of the undescribed
Euryboas from Langebaanweg. It is relevant because if Galiano & Frailey are
correct in concluding that Chasmaporthetes and Euryboas were closely related,
then the Langebaanweg species should, because of its age relative to recorded
Chasmaporthetes, be a more appropriate structural ancestor of the latter than is
A. eximia variabilis. Mention will also be made of the Langebaanweg A. australis
since it is evidently intermediate in age between A. eximia variabilis and C. kani,
and, if the latter two taxa are phylogenetically related, A. australis should also
have characters in common with C. kani.

The skulls and mandibles of C. kani and A. eximia variabilis are superficially
similar morphologically and little different in size. The mandible of Euryboas

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

may, on average, be more slender than those of the other two taxa, but this
possible difference has yet to be substantiated. There is certainly a greater
similarity between the mandible of C. kani and certain of the Langebaanweg
A. australis specimens than between that of the former and the Langebaanweg
Euryboas.

One of the more striking characteristics of the mandible in Euryboas is that
the corpora are remarkably straight, the two halves being steadily divergent
from the symphysis. Viewed ventrally the mandible is V-shaped. In this respect
Euryboas resembles all other members of the Hyaena group, living and fossil,
examined during the course of the present study. By contrast, at least some of
the mandibles of A. australis, A. eximia (e.g. Schmidt-Kittler 1976: pl. 3, fig. 6)
and North American Chasmaporthetes (e.g. Stirton & Christian 1940: fig. 1)
have markedly curved corpora and, viewed ventrally, are almost U-shaped.
This also applies to some specimens in the available series of living C. crocuta
specimens, in which four out of seven specimens have curved jaws.

Since mandibular curvature is not constant in any one species, it is an
unreliable distinguishing character. However, it may be significant that curved
mandibles were observed only in species which are, or which might be, members
of the Percrocuta group. Adcrocuta and Chasmaporthetes are here regarded as
percrocutas, while C. crocuta may be, even though it is generally included in the
Hyaena group (e.g. Thenius 1966; Galiano & Frailey 1977). This may be an
instance where a character (i.e. curved mandibles) is indicative of one group
only (i.e. the percrocutas), whereas the opposite condition (i.e. straight
mandibles) is found in both groups of hyaenas. Further observations are
required to test this hypothesis.

Jaw shape is usually an indication of the shape of the cheektooth rows,
although there are instances where straight jaws (viewed ventrally) occur
together with curved cheektooth rows. This applies in the case of, for example,
‘Hyaena brunnea. Toothrow curvature has, therefore, to be considered as a
separate character.

Galiano & Frailey (1977) have pointed out that in Euwryboas the cheektooth
rows are straight, whereas in Chasmaporthetes they are curved. The latter
condition is evident in both A. eximia variabilis and A. australis, even when the
jaws are straight. The primitive Langebaanweg Euryboas, like later forms, has
straight cheektooth rows, which is an indication that this was a characteristic
of the lineage for most, and probably all, of its history. The implication is that
Euryboas and Chasmaporthetes had evolved independently at least since late in
the Miocene.

As with the mandibles, there is a greater similarity between the teeth of
C. kani and A. australis than between those of the former and Euryboas. In
addition, there is a general similarity between the teeth of these three taxa and
those of A. eximia variabilis, although it is on the differences which do exist
that the question of interrelationships hinges.

Ratio diagrams are used to illustrate some of the similarities and differences

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 287

in tooth proportions in A. eximia variabilis, C. kani and the Langebaanweg
Euryboas (Fig. 9). There is a basic similarity in the patterns of tooth length in
the three taxa. The premolars of C. kani are only slightly shorter than those of
A. eximia variabilis, whereas the carnassials are appreciably shorter (Fig. 9A).
By contrast, the carnassials of C. kani and the Euryboas are of comparable
length, whereas the anterior premolars of the latter are longer (Fig. 9B). Thus
if the Euryboas represents the structural ancestor of C. kani, then the evolu-
tionary trend in respect of cheektooth length was reduction of premolar length.
Alternatively, if A. eximia variabilis was the ancestor, then the trend was
reduction of carnassial length.

One of the metrical characters not illustrated in the accompanying ratio
diagrams is crown height of the canines. The canines of A. eximia and North
American Chasmaporthetes johnstoni, and possibly also C. kani, are unremark-
able, relatively low-crowned teeth, whereas the canines of the Langebaanweg
Euryboas are relatively high-crowned. The length: crown height ratio for the C
of specimen SAM-—PQ-L21000 is 1 : 2, which compares closely to the ratio

100 110 120 130 100 110 120

Fig. 9. Ratio diagrams comparing mean lengths of upper and lower teeth of Adcrocuta eximia
variabilis (+) (Kurtén 1957), Chasmaporthetes kani (C1) (Galiano & Frailey 1977), and the
Langebaanweg Euryboas sp. (©). Standard of comparison (100%): Hyaena hyaena.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

of 1 : 1,97 in the £. bielawskyi from Roccaneyra (Schaub 1941). The ratio in
C. johnstoni is 1 : 1,79, while in one of the Langebaanweg A. australis specimens
(22204) it is 1 : 1,63. The canine height in Euryboas is clearly a specialized
character, which is not shared by at least some, and perhaps all, species of
Adcrocuta and Chasmaporthetes. Thus in terms of canine development Adcrocuta
is more appropriate than Euryboas as an ancestor for Chasmaporthetes.

C. kani differs from A. eximia variabilis in having a slightly larger P?.
Galiano & Frailey (1977: 6) are of the opinion that the P! of C. kani ‘is
proportionally larger than in any other hyaenid genus’. No explanation of this
character is offered but, since it is apparently unique, it must be regarded as a
specialization. By curious contrast in view of their conclusion on relationships,
Galiano & Frailey record the absence of P! as one of the characteristics of
Euryboas. In fact, this tooth is present in the primitive Langebaanweg species.
Since this species was ancestral to forms which had lost P!, it is perhaps unlikely
that it should also be the ancestor of another (i.e. C. Kani) in which this tooth
was unusually well developed. No such anomaly exists in respect of Adcrocuta,
which would, therefore, be a more appropriate ancestral form for C. kani.

C. kani also differs from A. eximia variabilis in lacking P,, a tooth which is
usually present in the latter. The reduction and eventual loss of P, is a general
rule in hyaenid evolution, so in respect of this tooth C. kani could well have
evolved from A. eximia variabilis, a species in which loss of P, was already being
manifested. On the other hand, the Langebaanweg Euryboas would be an
equally appropriate structural ancestor since, like C. kani, it lacks P,.

The Pe P. and P, of C. kani and A. eximia variabilis are of comparable
length, but those of C. kani are narrower (Table 3), the latter being a difference

TABLE 3

Mean dimensions of Chasmaporthetes kani and Adcrocuta eximia variabilis teeth.

Pe PP Re Pp M?

Chasmaporthetes kani* 8,8 7,8 181 11,3 21,9 13,7 . 32,6. ASO eam:
Adcrocuta examia

variabilis* 7,1 7,4 18,1 . 12,5 22,6 16,0 3845 19:00 Gore
P, ES Eb P, M,
] b ] b ] b ] b | b
Chasmaporthetes kani+ 0 0 15,8 90 19,2 10;7 2233 - 1S e2see
Adcrocuta eximia
variabilis” a3 5.9 164 11,8 19,7 142 22.2 “iss ees

1 Galiano & Frailey 1977.
* Zdansky 1924.

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 289

not shown on the ratio diagram (Fig. 9A). The Langebaanweg Euryboas has
premolars comparable in breadth to those of C. kani but which are, with the
exception of P,, slightly longer. Advanced Chasmaporthetes, like Euryboas, has
narrow, sectorial premolars as a major distinguishing characteristic. It follows
that while derivation of C. kani from A. eximia variabilis would be logical in
terms of the relative development of their premolars, derivation of the former
from the Langebaanweg Euryboas is not, since even this primitive Euryboas was
already more specialized in terms of premolar proportions.

Also relevant here is the observation by Galiano & Frailey (1977: 2) that
the ‘anterior accessory cusps of P, and P, [in C. kani are] relatively weak as
compared with other Chasmaporthetes species’. In this respect C. kani occupies
a position intermediate between A. eximia variabilis and advanced Chasma-
porthetes, an appropriate position in the hypothetical Adcrocuta-North
American Chasmaporthetes lineage. Assuming that Galiano & Frailey were
correct in reassigning some Old World hyaenids to Chasmaporthetes, then it
appears that Euryboas never did develop prominent anterior accessory cusps
on the anterior premolars. This need not necessarily exclude the Langebaanweg
Euryboas from the role as structural ancestor of Chasmaporthetes but, as in the
case of P!, it would be another instance where descendant forms evolved
different characteristics.

The shapes of the anterior lower premolars are also significant. In Chasma-
porthetes these teeth tend to be ovate in occlusal view, whereas in Euryboas they
are more or less rectangular in outline. This is most obvious in the case of P3.
In A. eximia variabilis and A. australis the P,;’s are rectangular, and in this
respect they resemble most, if not all, contemporary hyaenids. Rectangularity in
the lower premolars is thus a primitive condition, and their ovate shape in
Chasmaporthetes, combined with the narrowness of these teeth, appears to be
unique amongst hyaenids.

The carnassials of C. kani differ from those of A. eximia variabilis by being
shorter and, in the case of M,, in having a simple unicuspid talonid. The M,
talonid in A. eximia variabilis is variably developed, but both entoconid and
hypoconid may be prominent. The fact that the talonid is variable in this taxon
suggests that it was already evolving away from the more complex condition
characteristic of primitive hyaenids and tending towards that evident in C. kani.
The significance of carnassial shortening in the hypothetical A. eximia variabilis—
C. kani lineage is not known. Possibly the development of the premolars as more
effective shearing teeth in C. kani reduced the demand on the carnassials for
this function.

Galiano & Frailey (1977) record the absence of the metaconid and presence
of a unicuspid talonid on M, as characteristics of Euryboas. This does not apply
in the case of the primitive Langebaanweg species, which has a bicuspid talonid
and sometimes has a metaconid. These features would not, however, exclude
this species as a structural ancestor of C. kani.

Relevant here is the nature of the M, in the Langebaanweg A. australis.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Although this species, like 4. eximia variabilis, exhibits appreciable variation in
certain characters, its M, typically lacks the metaconid and has a unicuspid
talonid. Thus A. australis and C. kani are similar in terms of M, morphology,
whereas the Langebaanweg Euryboas 1s, in this respect, more primitive in an
evolutionary sense. These comments also apply with respect to the anterior
accessory cusps of the anterior lower premolars. This is here interpreted as a
further indication that the relationships of Chasmaporthetes lie with Adcrocuta
rather than with Euryboas.

The M! of C. kani, like the P?, is a little larger than that of A. eximia
variabilis. As a general rule, hyaenid evolution is characterized by a reduction
in the size of upper molars. Consequently, the increase in the size of the C. kani
M! relative to that of its hypothetical A. eximia variabilis ancestor is an anomaly
which requires explanation. A similar situation exists in the case of the Euryboas
lineage and it will be discussed first.

The M! of the Langebaanweg Euryboas is relatively large and more trans-
versely elongated compared with those of other members of the Hyaena group
from Langebaanweg. The transverse elongation is emphasized by the reduction
of the metastyle and the root which supports it. Judging from a cast of the
Val d’Arno E. bielawskyi maxillary fragment described by Schaub (1941), the
M!? of this species was even more transversely elongated, while the metastyle
and supporting root were apparently absent. Indications are that the transverse
elongation of M! was a progressive character in the Euryboas lineage and must,
therefore, have been of functional advantage in the otherwise essentially
sectorial dentition of this genus.

In primitive or unspecialized hyaenids, of which the living Hyaena hyaena
is an example, the M! occludes with the posterior parts of M,, food being
crushed between the occlusal surfaces of the M1 and the M, talonid. In specialized
species such as Crocuta crocuta the shearing blades of the carnassials are highly
developed, the M, talonid is small and the M! reduced or absent. Euryboas is
clearly quite different from either of the above examples since the M?! appears
disproportionately large in relation to the size of the M, talonid.

The Mt? in carnivores is not necessarily used only for crushing food. Even
in such groups as the Felidae, in which the cheekteeth are highly sectorial and
the M, talonid is absent in all but primitive forms, an M1? is present. In such
cases the M! also has a shearing function. As the jaws are closed the apex of the
M, protoconid passes across the P* metastyle in a manner normal for carnivore
carnassial shear. Thereafter it comes into contact with the anterior edge of M!
and the posterior keel of the protoconid slides along this edge with a slicing
action, which is at right angles to the main carnassial shear. Presumably it is
occlusion of this kind which had developed to a high degree in Euryboas. In this
instance there was the added refinement provided by the M, talonid, itself
sectorial in advanced species, which slid across the occlusal surface of the M?’,
probably with a cutting, rather than crushing, action.

The Langebaanweg Euryboas was relatively primitive in having a bicuspid

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 291

M, talonid and, in addition, it sometimes had a small M,. Thus, in this species
the crushing function of the molars had not yet been completely replaced by the
shearing function evident in later Euryboas.

The situation in the hypothetical Adcrocuta—Chasmaporthetes lineage was
probably essentially similar and the relatively large M! of C. kani may be
interpreted as a specialized, rather than primitive, character.

The M? of the Langebaanweg Euryboas is comparable in size to that of
C. kani and this may be yet another indication that there was no direct phylo-
genetic link between them. There evidently was transverse elongation (i.e. size
increase) in M?* on the Langebaanweg Euryboas—E. bielawskyi lineage, but
nothing comparable in the case of C. kani if it, too, stemmed from a Euryboas
resembling that from Langebaanweg. This would be yet another example of
possible descendants of the Langebaanweg Euryboas following different
evolutionary trends. While this is, of course, not impossible, it is unlikely.

The postcranial skeleton of Chasmaporthetes has yet to be recorded but,
since the dentition of this genus deviates markedly from those of most hyaenids
and in some respects resembles that of Euryboas, it is possible, and perhaps even
likely, that the postcranial skeleton also differed from those of ‘conventional’
hyaenids, and that it shared the specializations evident in Euryboas.

To sum up, the observable similarities and differences between the taxa
discussed here combine to weight the evidence in favour of an Adcrocuta rather
than Euryboas origin for Chasmaporthetes. All those characters in which the
dentition of early Pleistocene C. kani differs from that of late Miocene A. eximia
variabilis may be interpreted as advanced and none would exclude the latter
from an ancestral role. Most of the differences reflect stages in the development
of the highly sectorial postcanine dentition characteristic of North American
Chasmaporthetes, and C. kani is, in an evolutionary sense, in an intermediate
position in the hypothetical A. eximia variabilis-North American Chasma-
porthetes lineage suggested earlier (Hendey 1975). The similarities between
A. eximia variabilis, C. kani, and North American Chasmaporthetes also point
to a close relationship between these taxa.

By contrast, the latest Miocene/early Pliocene Euryboas from Langebaan-
weg is less well suited to the role as structural ancestor of Chasmaporthetes, and
a direct phylogenetic connection between these taxa is, at the very least, highly
improbable. On the other hand, the Langebaanweg Euryboas is in every respect
ideally suited to be the ancestor of the late Pliocene/early Pleistocene
E. bielawskyi of Europe.

The earlier view that Chasmaporthetes and Euryboas evolved independently,
with the latter being an essentially African genus which also spread into southern
Europe (Hendey 1975), is here maintained.

In considering the origins and relationships of Chasmaporthetes and
Euryboas, another important species which has to be taken into account is
‘Hyaena’ borissiaki from the Pliocene of Moldavia in the Soviet Union
(Khomenko 1932). De Beaumont (1967) believed it to be an intermediate

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

between Lycyaena and various species now assigned to Chasmaporthetes and
Euryboas. More recently Galiano & Frailey (1977: 9) noted that it has characters
in common with both these genera, but concluded that ‘H.’ borissiaki ‘appears
to be referable to Chasmaporthetes senso stricto’.

‘H. borissiaki is evidently broadly contemporary with the hyaenids from
Langebaanweg, although the actual ages of the Moldavian and South African
species are not known. The relative ages of these hyaenids could be crucial to
the interpretation of their relationships. ‘H.’ borissiaki has characters in common
with both Adcrocuta australis and the Euryboas from Langebaanweg. For
example, both ‘H.’ borissiaki and the Euryboas have straight cheektooth rows
and jaws, while their cheekteeth are comparable in both size and morphology.
On the other hand, ‘H.’ borissiaki resembles A. australis in cheektooth
morphology, most significantly in having a reduced P* protocone. In this
respect, and in the occasional presence of P,, it is also similar to A. eximia
variabilis.

In terms of the interrelationships postulated here, ‘H.’ borissiaki may be
visualized either as an intermediate between A. eximia variabilis and Chasma-
porthetes, or as an early Euryboas which was not conspecific with the Langebaan-
weg species. The first of these alternatives is perhaps the more likely and,
following Galiano & Frailey (1977), ‘H.’ borissiaki is here tentatively regarded
as an early Chasmaporthetes.

Nevertheless, the similarities to Euryboas are striking and, as a further
alternative, Chasmaporthetes ? borissiaki may be interpreted as the species from
which both Euryboas and Chasmaporthetes were derived. This possibility would
be strengthened ould it transpire that C. ? borissiaki predates the Langebaan-
weg Euryboas. In the case of this alternative, the stem form might either have
been Lycyaena, as De Beaumont (1967) suggested, or Adcrocuta, as indicated
above.

While the phylogenetic position of C. ? borissiaki is uncertain, it is clearly
a species of great significance in the matter of ‘hunting hyaena’ interrelationships.

Judged on available evidence the hypothetical Adcrocuta—Chasmaporthetes
transition took place in Asia some time between the late Miocene and early
Pleistocene. Members of this lineage dispersed eastwards into North America
by way of the Bering Land Bridge, and westwards into Europe and perhaps also
Africa.

A problem which arises in connection with this theory concerns the age of
C. kani, which was given as ‘Early Pleistocene’ by Galiano & Frailey (1977: 2).
If this was indeed the case, then C. kani must be younger than at least some of
the European and North American specimens assigned to Chasmaporthetes,
even though it is more primitive in an evolutionary sense. Either C. kani is
older than Galiano & Frailey suppose, or it was a conservative species, little
different from a Pliocene form from which late Pliocene/early Pleistocene
European and North American Chasmaporthetes must have been derived.
In other words, the latter are likely to have stemmed from a C. kani-like

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 293

ancestor of Pliocene rather than early Pleistocene age. C.? borissiaki could
possibly be the species in question.

The actual dates when Chasmaporthetes first appeared in Europe and North
America have yet to be firmly established. If the Plio/Pleistocene boundary is
taken at about 2 m.y. before present, then migration to North America must
have been during the late Pliocene, since Chasmaporthetes is recorded from the
‘early Blancan’ (Repenning 1967), which is of late Pliocene age (Kurtén 1971).
The Olivola and Senéze faunas of Europe, which include C. lunensis (Galiano &
Frailey 1977), date from the latter part of the Villafranchian (Kurtén 1968),
which postdates the early Blancan. The migration to Europe may thus have
been later than that to North America.

The ranges of Chasmaporthetes and Euryboas overlap both temporally
and geographically in Europe and, in view of their shared characteristics, they
may well have competed with one another. It is not known which of the two
taxa survived longest in Europe, but that honour may go to Chasmaporthetes.

The situation in Africa is somewhat obscure, the basic problem being to
which of the two genera material from the Transvaal caves and undescribed
specimens from east Africa (e.g. Howell & Petter 1976) belongs. The best

14
\ \

Sal

Fig. 10. Lateral and occlusal views of Chasmaporthetes nitidula hemimandible from Swartkrans
(Transvaal Museum SK14005). (80% of natural size.)

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Some characteristics of the lower teeth and jaws in Adcrocuta, Chasmaporthetes and Euryboas

late Miocene latest Miocene/ | ‘Early Pleistocene’
Adcrocuta eximia early Pliocene Chasmaporthetes
variabilis Adcrocuta australis kani
(China) (Langebaanweg) (China)
Shape of jaw straight (?or straight or
(ventral view) curved) curved

Shape of cheektooth curved curved curved curved
row (occlusal view)

Canine height low low h low

Shape of P; rectangular rectangular ovate ovate

Anterior accessory sometimes sometimes sometimes present
cusps of P, and P3 absent absent absent

M, talonid bicuspid or unicuspid unicuspid unicuspid
unicuspid

represented of the later African forms is ‘Euryboas’ nitidula from Swartkrans,
which Galiano & Frailey (1977) believe may be a Chasmaporthetes. In their
view referral to this genus would be more certain if ‘the condition of P! and the
degree of curvature of the tooth rows’ were known (Galiano & Frailey 1977: 9).
The condition of P? is still not known, although it may well have been absent,
but the tooth rows, and jaws, are curved (Fig. 10), as in Chasmaporthetes. The
case for referral of ‘E.’ nitidula to Chasmaporthetes is thus strengthened and the
Swartkrans species is here recognized as Chasmaporthetes nitidula.

The date when Chasmaporthetes entered Africa is not known, but it may
have postdated its entry into Europe. That unit of the Swartkrans fauna which
includes C. nitidula dates back about 1,5 m.y. (Vrba 1976), which makes it one
of the youngest records of the genus, another being the C. ossifragus from
Inglis IA in Florida (Webb 1974). There is thus evidence that in Africa
Chasmaporthetes survived longer than Euryboas.

Present indications are that Chasmaporthetes was the more widespread,
and ultimately also the more successful of the two ‘hunting hyaena’ genera.
It was, in fact, the most widely distributed of all hyaenids, having occurred over
much of North America as well as the Old World.

The interrelationships of Adcrocuta, Chasmaporthetes and Euryboas as
interpreted here are summarized in Figure 11, while some of their more
significant characters from a phylogenetic point of view are listed in Table 4.
Chasmaporthetes ? borissiaki is omitted owing to the uncertainties surrounding it.

The discovery of additional specimens of different ages and from different
geographical locations, and accurate dating of specimens already known,
should allow testing of the opinions expressed in this paper. The recent dis-

LATE TERTIARY HYAENIDAE FROM LANGEBAANWEG, SOUTH AFRICA 295

early Pleistocene | early Pleistocene latest Miocene late Pliocene

Chasmaporthetes | Chasmaporthetes early Pliocene Euryboas
ossifragus nitidula Euryboas sp. bielawskyi
(Inglis 1A) (Swartkrans) (Langebaanweg) (Roccaneyra)

curved curved straight straight

curved curved straight straight
low high high

ovate ovate rectangular rectangular

present present absent absent

unicuspid unicuspid bicuspid unicuspid

coveries at Langebaanweg and identification of the first Asiatic Chasmaporthetes
have already clarified the situation to some extent and no doubt more relevant
material will still come to light.

AFRICA EUROPE NORTH AMERICA

Chasmaporthetes
nitidula

Euryboas Chasmaporthetes Chasmaporthetes

bielawskyi lunensis kani SIGE Tes

johnstoni and
other specimens

Ww
Z
Ww
1S)
O
=
2
wi
=i
a
>
=
[4
<
wi

wi
wi
EO
SO ‘Chasmaporthetes
= kani’-like
ae species
Sain lu
Sez
ta = ai res Euryb
FO“Y uryboas Adcrocuta
< O < g Sp. australis*
PE VS

1
!
Hyaena ?Adcrocuta - | ~-~-~- Adcrocuta eximia Adcrocuta
group eximia eximia eximid
(?Lycyaena) subsp. variabilis

LATE MIOCENE

*Langebaanweg species

Fig. 11. Tentative phylogeny of Euryboas and Chasmaporthetes.

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Some characteristics of the lower teeth and jaws in Adcrocuta, Chasmaporthetes and Euryboas,

‘Early Pleistocene’
Chasmaporthetes

latest Miocene/

late Miocene /
early Pliocene

Adcrocuta eximia

late Pliocene
C hasmaporthere

variabilis Adcrocuta australis kani liters
(China) (Langebaanweg) (China) (Saint-Valliet)
Shape of jaw Sstraight (?or straight or euned
(ventral view) curved) curved
Shape of cheektooth curved curved ater
row (occlusal view)
Canine height low low ? low
Shape of P; rectangular rectangular ovate ovate
Anterior accessory sometimes sometimes sometimes present
cusps of P, and P; absent absent absent
M, talonid bicuspid or unicuspid unicuspid unicuspid

unicuspid

represented of the later African forms is ‘Euryboas’ nitidula from Swartkrans,
which Galiano & Frailey (1977) believe may be a Chasmaporthetes. In their
view referral to this genus would be more certain if ‘the condition of P! and the
degree of curvature of the tooth rows’ were known (Galiano & Frailey 1977: 9).
The condition of P? is still not known, although it may well have been absent,
but the tooth rows, and jaws, are curved (Fig. 10), as in Chasmaporthetes. The
case for referral of ‘E.’ nitidula to Chasmaporthetes is thus strengthened and the
Swartkrans species is here recognized as Chasmaporthetes nitidula.

The date when Chasmaporthetes entered Africa is not known, but it may
have postdated its entry into Europe. That unit of the Swartkrans fauna which
includes C. nitidula dates back about 1,5 m.y. (Vrba 1976), which makes it one
of the youngest records of the genus, another being the C. ossifragus from
Inglis IA in Florida (Webb 1974). There is thus evidence that in Africa
Chasmaporthetes survived longer than Euryboas.

Present indications are that Chasmaporthetes was the more widespread,
and ultimately also the more successful of the two ‘hunting hyaena’ genera.
It was, in fact, the most widely distributed of all hyaenids, having occurred over
much of North America as well as the Old World.

The interrelationships of Adcrocuta, Chasmaporthetes and Euryboas as
interpreted here are summarized in Figure 11, while some of their more
significant characters from a phylogenetic point of view are listed in Table 4.
Chasmaporthetes ? borissiaki is omitted owing to the uncertainties surrounding it.

The discovery of additional specimens of different ages and from different
geographical locations, and accurate dating of specimens already known,
Should allow testing of the opinions expressed in this paper. The recent dis-

late Pliocene

hasmaporthetes
johnstont

LATE TERTIAR
Y HYAENIDAE FROM LANGEBAANWEG SOUTH
, AFRICA

early Pleistocene
Chasmaporthetes
ossifragus

(Cita Canyon) (Inglis 1A)

curved

curved

OW
ovate

present

nicuspid

curved

curved

?
ovate

present

unicuspid

early Pleistocene
Chasmaporthetes
nitidula
(Swartkrans)

curved
curved

low
ovate

present

unicuspid

latest Miocene

early Pliocene

Euryboas sp,
(Langebaanweg)

Straight
Straight

high
rectangular

absent

bicuspid

295

late Pliocene
Euryboas
bielawskyi

(Roccaneyra)

Straight
Straight

high
rectangular

absent

unicuspid

coveries at Langebaanweg and identification of the first Asiatic Chasmaporthetes
have already clarified the situation to some extent and no doubt more relevant
material will still come to light.

EARLY PLEISTOCENE

wi
Zz
my
O
Q
=
a

PLIOCENE

LATE MIOCENE

AFRICA

Chasmaporthetes
nitidula

Euryboas
*

Hyaena

group
(?Lycyaena)

*Langebaanweg species

EUROPE

Euryboas
bielawskyi

Adcrocuta
SP. australis*
1

'
-—--- Adcrocuta eximia
eximia

Chasmaporthetes

Chasmaporthetes
kani

*Chasmaporthetes
kani’-like
species

Adcrocuta
eximia
yoriabilis

NORTH AMERICA

Chasmaporthetes
Johnstoni and
other specimens

Fig. 11. Tentative phylogeny of Euryboas and Chasmapo wtheres.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

I am indebted to Drs G. de Beaumont (Natural History Museum, Geneva),
A. W. Gentry (British Museum (Natural History)), R. G. Klein (University of
Chicago), B. Kurtén (University of Helsinki), P. V. Rich (Monash University),
T. H. Rich (National Museum of Victoria), and R. H. Tedford (American
Museum of Natural History) for assistance in preparing the manuscript of this
paper. The opinions expressed here are not necessarily shared by them. Many
other persons have contributed directly and indirectly to my studies on fossil
carnivores and to the South African Museum’s Langebaanweg Research
Project and, although not named here, my thanks to them for their interest and
assistance.

I am also indebted to Dr. C. K. Brain (Director, Transvaal Museum) for
the loan of specimens, and Drs V. J. Maglio (formerly of the Museum of
Comparative Zoology, Harvard), C. A. Repenning (U.S. Geological Survey,
Menlo Park) and S. D. Webb (University of Florida) for casts of specimens.

I thank Miss T. Salinger for the photographs, Miss K. Scott for preparing
Figure 10 and Mrs P. Eedes for assistance with the typing of the manuscript.

The Langebaanweg Research Project is supported by Chemfos Ltd, the
South African Council for Scientific and Industrial Research and the Wenner-
Gren Foundation for Anthropological Research (grant no. 2752-1834), and the
assistance of these organizations is gratefully acknowledged.

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Pee ne Peet
ea cee

a 7

Pia

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
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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. Zen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development i 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
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(continued inside back cover)

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

Volume 76 Band
September 1978 September
Patt) 73) Deel

NESS
© VVVWIS

S ‘
Coury now 6°

A FRAGMENTARY SPECIMEN OF SAURICAHTHYS
SP. FROM THE UPPER BEAUFORT SERIES OF
SOUTH AFRICA

By

JOHN GRIFFITH

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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A FRAGMENTARY SPECIMEN OF SAURICHTHYS SP. FROM THE
UPPER BEAUFORT SERIES OF SOUTH AFRICA

By
JOHN GRIFFITH

Westfield College, University of London
(With 1 figure)
[MS. accepted 18 July 1978]

ABSTRACT

The tip of an elongate jaw, found in association with the holotype of the amphibian
Capitosaurus africanus, is identified as belonging to the actinopterygian genus Saurichthys.
This is the first record of the family Saurichthyidae from the mainland of Africa; an additional
point of interest is its almost certain freshwater provenance.

CONTENTS

PAGE
Introduction : é ; » 299
Locality and horizon : 299
Description : : ; . 300
Identification . ; : : 302
Remarks . : : , F 305
Acknowledgements . : 2. 806
References . 2 2 ; : 306

INTRODUCTION

The specimen was discovered by Mrs Jone Rudner of the South African
Museum during preparation of the holotype of the amphibian Capitosaurus
africanus Broom, 1909 (SAM-—2360). It lay in the right interpterygoid vacuity of
the amphibian skull roughly at the same level as, but not in contact with, the
ventral surface of the palatal bones. The protection so afforded is probably
responsible for its preservation, but it is not possible to determine if there is
any other significance in the association.

The specimen was subsequently freed of matrix by Mrs Rudner and
tentatively identified as belonging to the genus Saurichthys by Dr J. Cosgriff,
of Wayne State University, Detroit, U.S.A.

LOCALITY AND HORIZON

The Capitosaurus skull was collected by Broom from the Cynognathus beds
of Farm Vaalbank, near Burgersdorp (Broom 1909). The Cynognathus zone
comprises the whole of the Upper Beaufort Series (Broom 1932; Hotton 1967)
and, according to Harland et al. (1967), is of Lower Triassic (Olenekian) age.

299

Ann. S. Afr. Mus. 76 (8), 1978: 299-307, 1 fig.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

DESCRIPTION

Order SAURICHTHYIFORMES
Family Saurichthyidae

Genus Saurichthys Agassiz, 1834
Saurichthys sp. indet.

The specimen (SAM-—2360a) consists solely of the incomplete tip of an
elongate jaw. It measures 11,8 mm in length and roughly 3,6 mm in width. In
dorsal and ventral view (Fig. 1) the anterior end appears roughly rounded and
behind this the two sides are more or less parallel. In lateral view the jaw is
slightly curved and decreases in height anteriorly so that, whereas the tooth-
bearing surface is almost flat, the opposite surface shows a more pronounced
curvature that increases towards the anterior end.

The external surfaces show traces of a weak ornamentation of short,
mainly longitudinal, ridges. There is no sign of a sensory canal.

The oral surface bears a narrow median ridge that projects slightly above
the general level of the bone and, in places, seems to show an indistinct
longitudinal suture-line. On either side of this ridge lie four large teeth or
tooth-bases, arranged more or less symmetrically, and between these a number
of less regularly arranged, smaller teeth. All, except one of the smallest, were
damaged or missing before fossilization.

The larger teeth lie in pairs, one on each side of the midline. Their bases
occupy the whole width between the median ridge and the edge of the jaw. They
are conical in shape and inclined away from the sagittal plane but with a slight
medial curvature so that their apices must have lain vertically above, or below,
the jaw edge. The anterior pair, in addition, project slightly forwards. The
remains of the best preserved large tooth measure 2,8 mm in height; when
intact it must have been at least 0,5 mm taller. The apical part of the tooth is
more transparent than the rest and is ornamented with fine, closely-spaced,
parallel striations running in an apical-basal direction. This part is marked off
from the remainder of the tooth by a barely visible groove and is estimated to
have accounted for slightly more than one-third of the total height. The surface
of the rest of the tooth is practically smooth except for moderate plication near
the region of attachment to the underlying bdne.

The small teeth are less than half the height of the large ones, about twice
as numerous, and occur either singly or in groups of two or three along the edge
of the jaw in the gaps between the large teeth. They appear to have been similar
in shape and structure to the larger teeth but all except one are broken and
abraded. The smallest tooth of all (arrowed in Fig. 1) is intact and, though no
surface detail can be made out in this tooth either, examination by transmitted
light with the specimen immersed in cedar-wood oil reveals some details of its
internal structure. There is a distinct apical cap of clear material which is
moderately birefringent and contains few tubules and is therefore identified as
enameloid (Schaeffer 1977) or modified dentine (Peyer 1968). The remainder of

A FRAGMENTARY SPECIMEN OF SAURICHTHYS SP. FROM SOUTH AFRICA 301

the tooth consists of orthodentine with the characteristic, numerous, closely-
spaced dentine tubules radiating from a central pulp cavity. This part shows only
very slight optical activity between crossed polarizers. The pulp cavity, in its
distal half at least, is relatively narrow and ends just short of the amelodentinal
junction. The proximal part of the tooth is too opaque for much detail to be
distinguished.

The broken end of the specimen is roughly D-shaped with a single, small,

Fig. 1. Stereoscopic pair of scanning electron microscope photographs of specimen SAM-—2360a
in oral view. The arrow points to the small tooth referred to in the text.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

nearly circular central cavity infilled with calcite. The bone is relatively thick
but compact and shows no sign of endochondrial origin.

IDENTIFICATION

The presence of a distinct apical cap of enameloid or modified dentine
clearly shows that the specimen belongs to an actinopterygian fish (Peyer 1968).
With this and the conspicuous elongation of the jaw in mind there appear to be
only four reasonable possibilities, that it is from: (i) a saurichthyid; (ii) an
aspidorhynchid; (iii) an early representative of one of the several groups of
long-jawed euteleosts; or (iv) a species unrelated to any of the known fishes with
elongate jaws. These alternatives will be considered in order.

(i) In the family Saurichthyidae the two jaws are more or less equal in
length. The anterior end of the upper jaw appears to be composed almost
entirely of the premaxillae (rostalo-premaxillaries in Stensi6’s (1925) nomen-
clature). These are curved transversely so as to form a tapering, hollow half-
cylinder that is completed ventrally by the horizontal dental lamellae which
extend medially to meet the lateral edges of the slender vomers. The resulting
structure has a single cavity, as in specimen SAM-—2360a. It seems unlikely that
the vomers extended right to the tip of the jaw, and here the two premaxillae
must have met ventrally as well as dorsally. Stensi6 (1925) suggests that separate
rostral and, or, postrostral elements were also involved but, whatever the
ontogeny of this region, none of the numerous specimens belonging to several
species examined by the author shows any sign of sutures dorsally though
suggestions of a midventral suture are occasionally visible. The anterior end of
the lower jaw appears to have been formed in a closely analogous manner from
the dentaries (dentalosplenials) with a minor contribution from the coronoids
(mixicoronoids) paralleling that of the vomers to the upper jaw. According to
Stensi6 (1925) in Saurichthys hamiltoni the two mandibular rami meet in a very
long, rigid symphysis but do not coalesce and a distinct median ventral suture
persists. However, in most species in which this region is known no sutures are
visible and the tip of the mandible is so similar, apart from its inversion, to the
corresponding part of the upper jaw that it is impossible to distinguish them on
morphological grounds alone.

The ethmoidal and mandibular sensory canals have never been traced
right to the tips of the jaws in any species and may have ended some distance
short of this.

In nearly all species in which the dentition is known each half of each jaw
carries a single series of fairly evenly spaced, large teeth interspersed with less
regularly arranged, smaller teeth and these tooth-rows extend right to the tips
of the jaws. Again there is no observable difference between upper and lower
jaws. The teeth themselves do show some interspecific differences but in general
are perfectly compatible in shape, proportions, ornamentation and structure
with those of the specimen described above.

There are, at present, four valid genera in the family Saurichthyidae:

A FRAGMENTARY SPECIMEN OF SAURICHTHYS SP. FROM SOUTH AFRICA 303

Saurichthys; Saurorhynchus Reis, 1892; Brevisaurichthys Beltan, 1972; and
Systolichthys Beltan, 1972.

The genus Saurichthys, which contains at least thirty-five species ranging
throughout the Triassic, has never been adequately defined because of the
fragmentary nature of the material of the type species, Saurichthys apicalis
Agassiz, 1834. However, several of the species ascribed to this genus are known
in some detail and nothing that the author has been able to discover suggests
that the specimen SAM-—2360a does not belong to this genus.

The genus Saurorhynchus contains two Lower Jurassic species. Though
not a diagnostic character, as it is shared with at least one species of Saurichthys,
both species of Saurorhynchus show the presence of ‘incissivlucken’, distinct
depressions in the bone of the jaw into which the apices of the large teeth of the
opposite jaw fitted in occlusion. The absence of this feature in the specimen
under discussion argues against it belonging to this genus.

The genera Brevisaurichthys and Systolichthys were erected by Beltan (1972)
each to contain a single new species from the Middle Triassic (? Ladinian)
of Spain. In both of these genera the elongation of the head is noticeably less
than in Saurichthys and Saurorhynchus and the relatively obtuse ends to the
jaws make it unlikely that SAM~—2360a belongs to either.

(ii) In the family Aspidorhynchidae the upper and lower jaws are dissimilar
in structure. In most species the head is continued beyond the anterior limit of
the mouth as a slender, tapering rostrum which is roughly circular in cross-
section and is toothless for all, or nearly all, of its length. In a few species,
e.g. Aspidorhynchus tenuirostris Agassiz, 1833, there is no rostrum, strictly
speaking, and both jaws extend to the anterior end of the head. In these species
the major part of the upper jaw is formed, as in the Saurichthyidae, from the
elongated premaxillae but each premaxilla is rolled into a complete, hollow
cylinder and, though these meet and fuse dorsally, they remain separated
ventrally for most, if not all, of their length, so forming a median groove for the
reception of the presymphysial teeth. The detailed structure of the extreme tip
of the upper jaw of these species is poorly known but appears to consist of a
half-cylinder grooved ventrally and with its interior filled with spongy
(? endochondrial) bone; a short distance behind the tip the paired, cylindrical
premaxillae can clearly be recognized, so that in cross-section the upper jaw
shows either two cavities or none at all. The dentition of the upper jaw differs
from that of the saurichthyids and of specimen SAM~—2360a in the segregation
of the teeth into rows or groups within which all neighbouring teeth are of
roughly similar size. Thus the fused premaxillae bear two principal tooth-rows
composed of relatively large teeth posteriorly and these decrease in size gradually
and evenly towards the tip of the jaw; smaller teeth are present on the posterior
part of the premaxillae in some species but these are confined to separate rows
flanking the larger teeth and never extend into the anterior part of the bone.

The anterior end of the lower jaw is formed by a single, median pre-
symphysial bone which carries a highly distinctive dentition consisting of a

304. ANNALS OF THE SOUTH AFRICAN MUSEUM

single, median row of large teeth flanked posteriorly, in some species only, by
one or two separate rows of much smaller teeth on either side.

The earliest known aspidorhynchid comes from the Bathonian Stage of the
Middle Jurassic which does leave an appreciable time-gap, but quite apart from
this it seems fairly obvious on anatomical grounds that specimen SAM-—2360a
cannot possibly belong to an aspidorhynchid.

(iii) The Euteleoste1 (Division III of Greenwood et al. 1967) include a
number of fishes in which the jaws show conspicuous elongation.

In the living Belonidae (garfishes) the jaws are roughly equal in length and
toothed throughout. The anterior end of the upper jaw is narrower than the
corresponding part of the lower jaw and fits partly within it during occlusion.
The upper jaw is formed, as in the other families described above, mainly from
the elongate premaxillae, each of which is completely rolled upon itself prior to
joining with its fellow so that in cross-section this jaw displays two cavities, one
on each side of the midline, and, moreover, is roughly oval in shape. The lower
jaw is roughly W-shaped in section with a well-developed median ridge separating
two deep channels into which the teeth of the upper jaw fit. The teeth, though
sharply pointed, are all relatively small and confined to the edges of the jaws.
As in the Aspidorhynchidae, larger and smaller teeth are segregated and do not
intermingle in the same row, and the smaller teeth do not extend to the anterior
ends of the jaws. In the closely related Hemirhamphidae (half-beaks) only the
mandible is elongated and this projects beyond the upper jaw as a toothless
pseudorostrum. The geological record of the suborder Exocoetoidei, to which
these fishes belong, is only known with certainty to extend back to the Middle
Eocene though there is a doubtful record from the Cretaceous (Maestrichtian)
(Patterson 1967).

The extinct suborder Alepisauroidei Rosen, 1973, contains several Upper
Cretaceous genera with greatly elongated jaws. In Ichthyotringa Cope, 1878,
Apateopholis Woodward, 1891, and Dercetis Agassiz, 1834, the two jaws are of
roughly equal length; in Rhynodercetis Arambourg, 1944, the upper jaw is
considerably longer than the lower and so forms a true rostrum. In all, the
anterior part of the upper jaw is formed largely from the premaxillae with
contributions from the mesethmoid, vomer and palatines, and the anterior part
of the lower jaw from the dentaries (Goody 1969). The dentition in all four
genera is unlike that of specimen SAM~—2360a. In Ichthyotringa each half-jaw
has a row of modestly-sized conical teeth and lateral to this a completely separate
marginal row of minute teeth. In Apateopholis the premaxillary and dentary
teeth are all extremely small. Dercetis has small, needle-like hollow teeth with
peculiar apecies, and on the dentary these teeth are arranged in clusters not rows.
Rhynodercetis has no teeth at all on the premaxillae and only small teeth on the
dentary.

The Xiphiidae (swordfishes) and Istiophoridae (marlins and sailfishes) have
conical rostra which project for some distance beyond the mandibles. Their
lower jaws are slightly elongated but remain more or less normal in appearance

A FRAGMENTARY SPECIMEN OF SAURICHTHYS SP. FROM SOUTH AFRICA 305

and the anterior ends do not resemble SAM-—2360a. The premaxillae are
toothless for all or most of their length. Dentary teeth are either minute or
absent.

The extinct family Palaeorhynchidae, grouped with the xiphiids and
istiophorids in the suborder Scombroidei, contains two Lower Tertiary genera:
Palaeorhynchus Blainville, 1818, with jaws of roughly equal length; and
Hemirhynchus Agassiz, 1844, in which the lower jaw is shorter than the upper.
In both genera the anterior parts of upper and lower jaws are circular in cross-
section, solid and completely devoid of teeth.

The Upper Cretaceous and Eocene Blocidae are poorly known and,
though they, too, were at one time included in the suborder Scombroidei,
Patterson (1973) regards their affinities as uncertain. The upper jaw is extended
into a rostrum which, according to Woodward (1901), ‘resembles a pair of tubes
pressed together’. The mandible is only about two-thirds of the length of the
upper jaw and rostrum; very little is known of its structure. Teeth are either
very minute or absent altogether.

None of the above shows sufficient resemblance to specimen SAM-—2360a
to indicate close affinity and, in any case, none can be traced backwards in time
beyond the Upper Cretaceous, and it seems extremely unlikely that a repre-
sentative of one of these groups, or indeed any euteleost, should have been in
existence in Lower Triassic times.

(iv) The specimen may not be closely related to any of the groups discussed
above. It could belong to a specialized offshoot of a group of more normal-
shaped fishes, or may represent a hitherto unknown group of long-jawed fishes.
As the only derived character shown by the specimen, namely the great elonga-
tion of the jaw, appears to have been independently acquired in a number of
phyletic lines such speculations are impossible to prove or disprove. On the
other hand, as all the observed features can be accounted for by the assumption
that the specimen belongs to a species of the genus Saurichthys (see (i) above),
these hypotheses are unnecessary and must be rejected in the interests of
parsimony.

In conclusion, in all of the observed features specimen SAM—2360a agrees
with, or falls within the range of structure shown by, the genus Saurichthys.
There is not sufficient information to determine if it is part of an upper or of a
lower jaw, to identify it as belonging to a known species of Saurichthys, or to
warrant its description as a new species of that genus. It differs from each of the
other taxa considered above in at least one character. It is unnecessary to
postulate that it belongs to some previously unknown genus. The specimen is
therefore determined as Saurichthys sp. indet.

REMARKS
There are two points which are perhaps worthy of brief comment:

(i) This is the first record of the Saurichthyidae from the mainland of Africa.
According to the palaeocontinental maps produced by Smith & Briden (1977)

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

the geographically nearest occurrences (in Triassic times) so far known are, in
order: Saurichthys madagascariensis Piveteau, 1944, and S. stensioi Lehman,
1952, from the Lower Triassic (Induan) of Madagascar; S. gigas and S. gracilis
(both first described by Woodward in 1890) from the Lower or Middle Triassic,
and S. parvidens Wade, 1935, from the Middle Triassic of New South Wales,
Australia; numerous Lower, Middle and Upper Triassic and Lower Jurassic
species from Europe. The saurichthyids appear to have had an almost world-wide
distribution and their previous absence from records for South Africa probably
merely reflects the paucity of fossiliferous marine deposits of suitable age.

(ii) The Upper Beaufort Series are generally accepted as being continental fresh-
water fluviatile and lacustrine deposits and, though Hotton (1967) suggests that
these conditions alternated with persistent deltaic ones, Rayner (1971) regards
this hypothesis as untenable. The saurichthyids appear to have been pre-
dominantly marine though a few species have been recorded from deposits
thought to have been laid down under fresh or ‘brackish’ water conditions.
Specimen SAM-2360a could have been reworked from earlier, marine beds
such as the marine phases of the Ecca or Dywka Series, but as no saurichthyid
has ever been recorded there this seems improbable. It therefore appears that
some species of saurichthyid were at least capable of temporary excursions into
fresh water and may even have lived there permanently.

ACKNOWLEDGEMENTS

I wish to record my gratitude to the Director of the South African Museum
for the loan of the specimen and to Dr M. A. Cluver for bringing it to my
attention and for a helpful correspondence. I am also indebted to Mrs M. Petri
for taking the scanning electron microscope photographs reproduced here as
Figure 1.

REFERENCES

Acassiz, L. 1833. Recherches sur les poissons fossiles 2 (1). Neuchatel.

Acassiz, L. 1834. Aberissene Bemerkungen uber fossile Fische. Neues Jb. Miner. Geol.
Paldont. 1834: 379-390.

Acassiz, L. 1844. Recherches sur les poissons fossiles 5 (1). Neuchatel.

ARAMBOURG, C. 1944. Note préliminaire sur quelques poissons fossiles nouveaux. Bull. Soc.
géol. Fr. (5) 8: 281-285.

BELTAN, L. 1972. La faune ichthyologique du Muschelkalk de la Catalogne. Mems. R. Acad.
Cienc. Artes Barcelona 41: 281-325.

BLAINVILLE, H. D. 1818. Poissons fossiles. In; Nouveau Dictionnaire d’Histoire Naturelle,
Paris 27: 310-395.

Broom, R. 1909. Notice of some new South African fossil amphibians and reptiles. Ann. S. Afr.
Mus. 7: 270-278.

Broom, R. 1932. The mammal-like reptiles of South Africa and the origin of mammals. London:
Witherby.

Cope, E. D. 1878. Description of fishes from the Cretaceous and Tertiary deposits west of the
Mississippi River. Bull. U.S. geol. geogr. Sury. Territ. 4 (1): 67-77.

Goopy, P. 1969. The relationship of certain Upper Cretaceous teleosts with special reference
to the myctophoids. Bull. Br. Mus. nat. Hist. (Geol.) Suppl. 7: 1-155.

A FRAGMENTARY SPECIMEN OF SAURICATHYS SP. FROM SOUTH AFRICA 307

GREENWOOD, P. H., ROSEN, D. E., WEITZMAN, S. H. & Myers, G. S. 1966. Phyletic studies of
teleostean fishes, with a provisional classification of living forms. Bull. Am. Mus. nat.
Hist. 131: 339-456.

HARLAND, W. B. ef al., eds. 1967. The fossil record. London: Geological Society.

Hotton, N. III. 1967. Stratigraphy and sedimentation in the Beaufort series (Permian-
Triassic), South Africa. In: TEICHERT C. & YOCHELSON E. L. eds. Essays in paleontology
and stratigraphy: 390-428. Lawrence, London: University of Kansas Press. (Special
publication 2.)

LEHMAN, J-P. 1952. Etude complémentaire des poissons de |’Eotrias de Madagascar. K. svenska
Vetensk Akad. Hand. (4)2(6): 1-201.

PATTERSON, C. 1967. Subclass Teleostei. Jn: HARLAND, W. B. et al., eds. The fossil record:
654-666. London: Geological Society.

PATTERSON, C. 1973. Interrelationships of holosteans. Jn: GREENWOOD, P. H. ef al., eds.
Interrelationships of fishes: 233-305. London: Academic Press.

PEYER, B. 1968. Comparative odontology. xiv + 347 pp. Chicago, London: University of
Chicago Press.

PIVETEAU, J. 1944-45. Paléontologie de Madagascar XXV: les poissons du Trias inferieur.
La famille des saurichthyides. Ann/s Paléont. 31: 77-87.

RAyner, D. H. 1971. Data on the environment and preservation of late Palaeozoic tetrapods.
Proc. Yorks. geol. Soc. 38: 437-495.

Reis, O. M. 1892. Zur Osteologie und Systematik der Belonorhynchiden und Tetragono-
lepiden. Geogn. Jh. 4: 143-170.

Rosen, D. E. 1973. Interrelationships of higher euteleosteans. Jn: GREENWOOD, P. H. ef al., eds.
Interrelationships of fishes: 397-513. London: Academic Press.

SCHAEFFER, B. 1977. The dermal skeleton in fishes. In: ANDREWS, S. M. et al., eds. Problems in
vertebrate evolution. London: Academic Press.

SmiTH, A. G. & BRIDEN, J. C. 1977. Mesozoic and Cenozoic palaeocontinental maps. Cambridge:
Cambridge University Press.

STENSIO, E. A. 1925. Triassic fishes from Spitzbergen. Part 2. K. svenska Vetensk Akad. Hand.
(3)2(1): 1-261.

WADE, R. T. 1935. The Triassic fishes of Brookvale, New South Wales. London: British Museum
(Natural History).

Woopwaprb, A. S. 1890. The fossil fishes of the Hawkesbury series at Gosford. Mem. geol.
Sury. N.S.W. (Palaeont.) 4: 1-5.

Woopwaprp, A. S. 1891. On some Upper Cretaceous fishes of the family Aspidorhynchidae.
Proc. zool. Soc. Lond. 1890: 629-637.

WooDwaprbD, A. S. 1901. Catalogue of the fossil fishes in the British Museum (Natural History) 4.
London: British Museum (Natural History).

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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
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An author’s name when cited must follow the name of the taxon without intervening
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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
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Synonymy arrangement should be according to chronology of names, i.e. all published
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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, ee 37.
Leda plicifera A. Adams, 1856: 5
Laeda bicuspidata Hanley, 1859: is, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861:
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
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In describing new species, one specimen must be designated as the holotype; other speci-
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Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
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’

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Name of new genus or species is not to be included in the title: it should be included in the
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Biological Abstracts.

JOHN GRIFFITH

A FRAGMENTARY SPECIMEN OF SAURICHTHYS
SP. FROM THE UPPER BEAUFORT SERIES OF
SOUTH AFRICA

VOLUME 76 PART 9 SEPTEMBER 1978 ISSN 0303-2515

ANNALS

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

FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100—140.

FiscHER, P.-H., DuvAL, M. & RaArry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. 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.

(continued inside back cover)

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

Volume 76 + #4Band
September 1978 September
Part 9 Deel

TWO NEW SPECIES OF GASTROSACCUS
(CRUSTACEA, MYSIDACEA) FROM SANDY BEACHES
IN TRANSKEI

By

T. WOOLDRIDGE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

TWO NEW SPECIES OF GASTROSACCUS (CRUSTACEA,
MYSIDACEA) FROM SANDY BEACHES IN TRANSKEI

By

T. WOOLDRIDGE
Department of Zoology, University of Port Elizabeth

(With 8 figures)
[MS accepted 2 August 1978]

ABSTRACT

Sampling by means of a sledge from the low-water mark out to a depth of 1,5 m showed
clear peaks of maximum distribution for Gastrosaccus bispinosa sp. nov. which was more
abundant at the upper limit of the low-water mark. Gastrosaccus longifissura sp. nov. was
more common in the breaker zone. Both species were also collected in the more sheltered
waters along the edge of the sandbank inside the mouth of the Mgazana estuary.

The known distribution of Gastrosaccus psammodytes Tattersall, 1958, is extended further
eastwards as far as Kei Mouth. The appearance of G. psammodytes in off-shore plankton
samples is discussed. A note on the distribution and a key to the species of Gastrosaccus
recorded in southern Africa are given.

CONTENTS

PAGE
Introduction . : ? . : : : ; : ; <2 309
Description of material . : : ; : : : : 5 aot
Distribution of Gastrosaccus recorded in southern Africa eS)
Key to the species of Gastrosaccus recorded in southern Africa 326
Acknowledgements . . : ; : : ‘ : : 3 4326
References : : ; : : ; : : : : 26

INTRODUCTION

Two new species of Gastrosaccus (Crustacea, Mysidacea) from sandy
beaches in Transkei are described and illustrated. G. psammodytes is described
from South Africa (Tattersall, O. S. 1958) while other sandy beach mysids are
known from various parts of the world. G. vulgaris is reported from Japan
(Nakazawa 1910) while Gauld & Buchanan (1956) have recorded G. spinifer
on sandy beaches in Ghana. Macquart-Moulin (1977) notes the distribution of
G. mediterraneus and G. spinifer along Mediterranean sandy shores. G. sanctus
is known from sandy beaches in the northern hemisphere (Bacescu 1934;
Tattersall, W. M. 1927; Tattersall & Tattersall 1951; Moran 1972). In southern
Africa G. sanctus is recorded from inshore waters (Lazarus 1975) and off-shore
waters only (Tattersall, O. S. 1957). Other reports of mysids from sandy beaches
are noted by Brown & Talbot (1972).

The two new species reported here are recorded from sandy beaches at
Mbotyi (31°28’S) and at Mgazana (31°41’S) on the Pondoland coast, Transkei

309

Ann. S. Afr. Mus. 76 (9), 1978: 309-327, 8 figs.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Fig. 1). G. longifissura sp. nov. is also recorded from Kei Mouth (32°41’S),
South Africa (Fig. 1), where it occurred with G. psammodytes.

The present species overlapped along a 70 m transect into the breaker zone.
Sampling was done with a sledge from the low-water mark out to a depth of
1,5 m. Beyond the 1,5 m depth contour, sampling with the sledge apparatus was
not possible due to excessive turbulence. Along the transect clear peaks of
maximum distribution were evident for each of the two species. G. bispinosa
sp. nov. was more abundant at the upper limit of the low-water mark, while
G. longifissura sp. nov. was more common in the breaker zone.

Both species were also collected along the edge of the sandbank inside the
mouth of the Mgazana estuary. At night they were taken in the breaker zone
with a plankton net (WP 2 net of 190 micron aperture towed by hand in water
of 1 to 1,5 m depth). They were absent in surface plankton samples taken at

Indian

Fig. 1. Map of the coastline indicating the position of the localities mentioned in the text.

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 311

night in the mouth channel of the estuary (samples collected on ten occasions
between 1971 and 1973), except in September 1972 when two specimens of
G. bispinosa sp. nov. were captured (Wooldridge 1977).

DESCRIPTION OF MATERIAL
Gastrosaccus bispinosa sp. nov.
Figs 2-5
Holotype

SAM-A15749 lodged in the South African Museum, Cape Town. Adult
female from Mgazana beach (31°42’S), collected by T. Wooldridge, 30 September
w97 7.

Paratypes

SAM-—A15750 lodged in the South African Museum, Cape Town. Numerous
adult males and adult females from Mgazana beach (31°42’S), collected by
T. Wooldridge, 30 September 1977.

Description

Carapace long, anterior margin produced into a triangular rostrum not
covering the eyestalks (Fig. 2A). Apex smoothly rounded. Posterior margin of
carapace emarginate, exposing the last thoracic somite. Posterior half of this
emargination forming a forwardly directed lobe which overlaps the more
proximal portion of the emargination on each side. In lateral view carapace
extends posteriorly to cover the whole of the thorax and the first abdominal
somite.

Antennule (Fig. 4A), first segment of peduncle slightly shorter than second
and third combined. Five or six small setae on outer distal angle. Second
segment of peduncle short and armed with three strong spines set obliquely
along its outer margin. Inner distal corner with a fine seta. Third segment
relatively slender, bearing a curved finger-shaped process on dorsal side at the
anterior end. A small spine present just posterior to the process. Outer flagellum
swollen at the base and bearing relatively long, flattened setae. In the male the
usual hirsute lobe present.

Antennal peduncle extending slightly beyond the distal end of the second
segment of the antennular peduncle (Fig. 2A). The second segment of antennal
peduncle about two and a half times as long as the third, bearing five plumose
setae on the inner margin and two setae on the inner distal angle (Fig. 2B).
Third segment with four plumose setae on inner margin and four and two
smaller plumose setae on the inner and outer distal corners respectively.
Antennal scale as long as peduncle and about three times as long as broad.
Outer margin straight, naked, terminating in a strong spine which does not
extend beyond the rounded apex of the scale. Inner margin convex and setose
(Fig. 2B).

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 2. Gastrosaccus bispinosa sp. nov.

A. Carapace in dorsal view. B. Antennae. C. First thoracic appendage.
D. Eighth thoracic appendage.

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 313

Mandible (Fig. 3A) well developed without spine row. Palp long and
slender, first segment unarmed. Second and third segments extremely setose,
the third segment with a terminal comb-like process and two strong apical
spines.

Maxillule (Fig. 3B) three segmented, first and second segments appear to be
fused. Lobe from first segment with three short serrated spines and three
longer spines armed with a row of fine setae and four to six teeth distally. Third
segment drawn out into a well-developed lobe which is armed with a close
group of short, strong, spinous and slightly curved spines. Row of six serrated
spines along the inner, sub-terminal margin which are setose proximally.

Maxilla (Fig. 3C) typical of the genus.

First thoracic limb with well-developed endite on basal segment (Fig. 2C).
Endopod short and robust, densely setose along inner margin. A well-developed
seta present on outer distal angle of carpus segment. Dactylus well developed,
without claw. First segment of exopod large and expanded, the outer distal
angle smooth (exopod twisted in figure). Flagellum fourteen-segmented, each
segment with one or two long plumose setae.

Second thoracic limb similar in form to first, dactylus of endoped with
claw. First exopod segment with a small tooth on outer distal angle.

Third to eight thoracic limbs similar in form, but becoming progressively
stronger and longer posteriorly. Carpus and propodus of endopod fused and
divided into many short subsegments. The number of these subsegments
increases posteriorly as follows: in the third limb, ten; in the fourth, eleven;
in the fifth, twelve; in the sixth, thirteen; in the seventh, sixteen; in the eighth,
eighteen (Fig. 2D). Each subsegment bears a small brush of setae and a small
spine on the inner distal angle, and a small spine on the outer distal angle. The
first segment of exopod in third to seventh pairs of thoracic limbs large and
armed with a strong tooth on outer distal corner. In the eighth pair of appen-
dages this angle is smoothly rounded (Fig. 2D). Flagellum sixteen- to eighteen-
segmented, each segment with one or two long plumose setae.

First pleoped of female (Fig. 4B) with long, slender sympod armed
proximally with three, and distally with nine, long plumose setae. Exopod about
twice as long as wide, armed with one spine-like seta, nine short plumose setae
and two relatively long plumose setae at the distal end. Endopod almost four
times as long as wide, armed with nine plumose setae distally. Remaining
pleopods in the female in the form of simple, unjointed plates, becoming
progressively longer on the posterior somites.

First pleopod of the male (Fig: 4D) with swollen sympod, outer margin
armed with thirteen long plumose setae. Endopod short, unsegmented, with two
terminal plumose setae and a single subterminal spine-like seta. Seven short
plumose setae also present, their relative position illustrated in Figure 4D.
Exopod composed of thirteen or fourteen segments, each segment armed with
two unequal plumose setae.

Second pleopod of the male (Fig. 5A) with large rectangular sympod which

314

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——

ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 3. Gastrosaccus bispinosa sp. nov.
A. Mandible. B. Maxillule. C. Maxilla.

pg

PSS

315

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI

Ae

Ie
ales
EIO

5

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Fig. 4. Gastrosaccus bispinosa sp. nov.
A. Antennule. B. First pleopod of female. C. Second pleopod of female.

D. First pleopod of male. E. Uropod.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

is indented on distal margin. Endopod eight-segmented and slender, subequal
in length to sympod. A well-developed pseudobranchial lobe at base of first
endopod segment, armed with a number of small plumose setae and one spine-
like seta. Exopod robust, almost twice as long as endopod, setae along inner
margin considerably shorter than those on outer margin.

Third pleopod of the male (Fig. 5B) with four-segmented endopod, the
first segment large and bulbous, bearing a well-developed pseudobranchial lobe
which is armed with a number of small plumose setae and a single spine-
like seta. Second and third endopod segments small, each armed with two short
plumose setae. The terminal segment of endopod reaching midpoint of first
exopod segment, bearing two relatively long plumose setae. Exopod four-
segmented, extending backwards to proximal end of the telson. First segment
subequal in length to the second, which is about twice the length of the third.
Fourth segment equal in length to the first segment. In some specimens a
prominent protrusion along concave margin. Apex armed with two strong,
barbed setae, the barbs of the one seta thin and spine-like, those on the other
seta robust and considerably smaller in the distal half (Fig. 5C). Remaining
pleopods in the male small, endopod reduced to a single segment.

Uropods (Fig. 4E) extending a short distance beyond telson. Exopod sub-
equal in length to endopod, armed along outer margin with seventeen strong,
regular spines which are finely plumose along the posterior margins. Apex of
these spines with a short curved tip. Endopod more slender than exopod,
tapering distally with seven long, curved irregularly-spaced spines amongst setae
on inner margin. Two posteriorly directed spines present on inner side of
statocyst. Anterior to the statocyst a row of small, graduated, closely set setae
and a row of three small setae present. Outer margin of endopod with a row of
plumose setae which increase in length posteriorly. Along the outer margin and
set irregularly amongst the longer setae are a number of extremely short plumose
setae.

Telson (Fig. 5D) about three times longer than its width at the base.
Lateral margins armed with six strong spines of which the distal two on each
side are longer than the others. Apical spines long and strong. Spaces between
the last three lateral and terminal spines occupied with one to six small spinules
which become more numerous distally. A strong spine present on each side of
the cleft on the dorsal side. Cleft one-quarter of the length of the telson and
armed with twenty to twenty-five spinules on either side. Spinules increase in
length and robustness posteriorly.

Length
Adult female 13,3—-17,5 mm
Adult male 11,0-14,5 mm

Remarks
Gastrosaccus bispinosa shows small affinities to Gastrosaccus gordonae
Tattersall, 1952. It is readily distinguished, however, by the prominent spine on

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 317

f5mm_—___. B,D
05 Vi A

05mm

@

Fig. 5. Gastrosaccus bispinosa sp. nov.

A. Second pleopod of male. B. Third pleopod of male. C. Terminal setae of third
pleopod of male. D. Telson.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

each side of the cleft and the armature along the lateral margins of the telson.
There is some diversity in the armament of the small spinules between the last
three pairs of lateral and terminal spines. In some specimens the spinule in the
space between the anti-penultimate and penultimate lateral spine was absent,
while in others it was present on one side only. The number of spinules distal to
the last pair of lateral spines also varied slightly and numbered either five or six
on each side.

The species may also be identified by the form of the third pleopod of the
male. The endopod is composed of only four segments, while the armature of
the barbed setae at the base of the fourth exopod segment is characteristic.

Gastrosaccus longifissura sp. Nov.
Figs 6-8

Holotype

SAM-—A15751 lodged in the South African Museum, Cape Town. Adult
female from Mgazana beach (31°42’S), collected by T. Wooldridge, 30 September
LOFT.

Paratypes

SAM-A15752 lodged in the South African Museum, Cape Town.
Numerous adult males and adult females from Mgazana beach (31°42’S),
collected by T. Wooldridge, 30 September 1977.

Description

Carapace (Fig. 6A) with anterior margin produced to form a triangular
rostrum, acutely rounded. Posterior margin deeply and narrowly emarginate,
exposing the last thoracic somite. The emargination is notched proximally,
forming a forwardly directed lobe which overlaps the anterior portion of the
emargination laterally. In lateral view the carapace extends posteriorly to cover
the thorax entirely as well as the first abdominal somite.

Antennule (Fig. 6B), first segment of peduncle slightly shorter than second
and third peduncular segments combined, outer distal angle with four or five
plumose setae. Second segment of peduncle short, armed with three slender
spines set obliquely along the outer margin. Inner distal corner with a fine
setae. Third segment almost twice as long as broad, with a slender, curved,
finger-shaped process dorsally on the outer distal corner. Outer flagellum
swollen at the base and bearing long flattened setae. In the male a well-developed
hirsute lobe present.

Antennal peduncle relatively long and extending forward almost to mid-
point of the third segment of antennular peduncle (Fig. 6C). Second segment of
antennal peduncle about two and a half times as long as broad, with four
plumose setae on the inner lateral margin and two or three plumose setae at the
distal corner. Third segment about one-third the length of second segment, with

319

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI

Fig. 6. Gastrosaccus longifissura sp. nov.
A. Carapace in dorsal view. B. Antennule. C. Antennae. D. Mandible. E. Maxilla.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

eight plumose setae at the distal end. Antennal scale as long as peduncle and
about four times as long as broad. Outer margin naked, terminating in a strong
spine which does not extend beyond rounded apex of scale. Inner margin
setose.

Mandible (Fig. 6D) without spine row. Palp long and slender, first segment
unarmed. Setation of second and third segments as in Figure 6D. Third segment
with a terminal comb-like process and two apical spines.

Maxillule (Fig. 7A), lobe from first segment with five relatively short
spines and three long terminal spines, the armature of the shorter spines as in
Figure 7A. Three terminal spines about twice as long as the shorter spines, each
armed distally with about four strong teeth. Lobe from third segment armed
with a close group of short, strong, spinous, slightly curved spines. A row of
six spines along inner, subterminal margin, serrated distally and plumose
proximally.

Maxilla (Fig. 6E), lobe from coxal segment with many well-developed spine-
like setae, those at the apex shorter and more closely set. Lobe from basis
incised to base, armed with many short setae along inner margin in the distal
half and at the apex. Exopod with outer margin markedly convex, armed with
about fourteen plumose setae.

First thoracic limb with well-developed endite on basal segment (Fig. 7B).
Endopod short, robust, densely setose along inner margin of first segment.
A single seta on outer distal angle of carpus. Propodus armed on outer margin
only. Dactylus with many spine-like setae, without claw. First exopod segment
large and expanded, the outer distal angle smooth (exopod twisted in figure).
Flagellum fourteen-segmented, each segment with one or two plumose
setae.

Second thoracic limb similar in form to first, dactylus of endopod with
claw. First exopod segment with a well-developed tooth on outer distal angle.

Third to eighth thoracic limbs similar in form, but becoming progressively
stronger and longer posteriorly. Carpus and propodus of endopod fused and
divided into many short, subsegments. The number of these subsegments
increases posteriorly as follows: in the third limb, eight; in the fourth, nine;
in the fifth, ten; in the sixth, eleven; in the seventh, fourteen; in the eighth,
sixteen. Each subsegment bears a brush of small setae and a single spine on the
inner distal angle and a small spine on the outer distal angle. The first exopod
segment in the third to seventh pairs of thoracic limbs armed with a strong
tooth on the outer distal angle. In the eighth pair of appendages (Fig. 7C) this
angle is smoothly rounded (exopod twisted in figure). Flagellum of exopod
fourteen- to sixteen-segmented, each segment with one or two long plumose
setae.

First pleopod of female (Fig. 7D) with long, slender sympod armed
proximally with three, and distally with seven long plumose setae. Exopod
about three times as long as wide, armed with one spine-like seta, seven short
plumose setae, and two relatively long plumose setae at the distal end. Endopod

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 321

(A = x
SSS
===

— SS

Fig. 7. Gastrosaccus longifissura sp. nov.

A. Maxillule. B. First thoracic appendage. C. Eighth thoracic appendage.
D. First pleopod of female.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

almost four times as long as wide, armed with five plumose setae in the distal
half. Remaining pleopods in the female becoming progressively longer on the
posterior somites, each armed with many plumose setae and a single spine-like
seta.

First pleopod of the male (Fig. 8B) with swollen sympod, the outer margin
armed with ten long plumose setae. Endopod short, unsegmented, with two
terminal plumose setae and a subterminal spine-like seta. Seven short plumose
setae present, their relative positions illustrated in Figure 8B. Exopod with
eight segments, the first almost twice as long as endopod. Each segment armed
with two unequal setae except the last which has two equal terminal setae and a
single sub-equal lateral seta.

Second pleopod of the male with sympod twice as long as wide (Fig. 8C).
Endopod slender and five-segmented, almost three-quarters as long as sympod.
A well-developed pseudobranchial lobe on first endopod segment, armed with
a number of small plumose setae and one spine-like seta. Exopod robust,
almost twice as long as endopod, setae along inner margin considerably shorter
than those on the outer margin.

Third pleopod of the male with three-segmented endopod (Fig. 8D). The
first segment large and bulbous, bearing a well-developed pseudobranchial lobe
which is armed with a number of plumose setae and a single spine-like seta.
Second and third endopod segments small, the terminal segment reaching
almost one-third along the length of the first exopod segment and armed with
two relatively long plumose setae. Exopod four-segmented, extending back-
wards to proximal end of telson. Exopod segments become progressively shorter
distally. First segment more than twice as long as the fourth, which bears two
non-plumose setae at the proximal end. Apex armed with two curved, barbed
setae, one of which is two and a half to three times the length of the other. The
longer seta is armed in the proximal half only. Remaining pleopods in the male
small, endopod reduced to a single segment.

Uropod (Fig. 8E) extending a short distance beyond telson. Exopod as long
as endopod and armed on the outer margin with sixteen strong, regular spines
which are finely plumose along the posterior margins. Apex of these spines with
a short, curved tip. Endopod more slender than exopod, tapering distally, with
seven long, curved, irregularly spaced spines amongst setae on inner margin.
A single posteriorly directed spine present on inner side of statocyst. Two rows
of small, closely set setae present anterior to statocyst on dorsal side. Outer
margin of endopod with a row of eight short plumose setae opposite statocyst.
Outer margin of endopod armed posteriorly with a row of graduated plumose
setae, which are interspersed with a number of short plumose setae.

Telson (Fig. 8F) about three times as long as broad at the base. Lateral
margins armed with seven strong spines of which the distal two on each side are
longer than the others. The spaces between the last three pairs of lateral and
terminal spines occupied with two, three or four small spinules which become
more numerous distally. Apical spines long and strong, cleft slightly less than

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 323

Fig. 8. Gastrosaccus longifissura sp. nov.

A. Second pleopod of female. B. First pleopod of male. C. Second pleopod of male.
D. Third pleopod of male. E. Uropod. F. Telson.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

one-quarter of the length of the telson. Cleft armed with twenty to twenty-five
small spinules on either side.

Length

Adult female, 8,5-10,5 mm
Adult male, 8,2—9,8 mm

Remarks

G. longifissura shows affinities to G. bispinosa sp. nov. and G. gordonae
Tattersall, 1952. It is distinguished from G. bispinosa by the absence of a spine
on each side of the cleft on the dorsal side of the telson. The third pleopod of
the male is also characteristic for each species. The endopod is four-segmented
in G. bispinosa and three-segmented in G. /ongifissura. The terminal setae on the
fourth exopod segment are subequal in length in G. bispinosa. In G. longifissura
one of the seta is two and a half to three times the length of the other, while the
fourth exopod segment also bears two non-plumose setae at the proximal end.
These setae are absent in G. bispinosa.

G. longifissura is distinguished from G. gordonae mainly in the form of the
telson. In G. gordonae, there are eight large spines on the lateral margin of the
telson (the terminal spine is not grouped with the lateral spines in the present
work), with the spaces between the third to the terminal spine occupied with
three to six small spinules on each side. In G. /ongifissura the lateral margin of
the telson is armed with seven large spines. The spaces between the fifth to the
terminal spines are occupied with two to four spinules.

Important differences between the two species are also found on the
uropods. In G. gordonae, the endopod is armed on the inner side of the statocyst
with two unequal, posteriorly directed spines. The inner margin is armed with
a row of ten small, unequal spines extending from the statocyst to half-way
along the margin. In the distal half there are three long, widely-spaced spines.
In G. longifissura there is only one posteriorly directed spine on the inner side
of the statocyst and seven long, irregularly spaced spines among the setae on the
inner margin.

Differences are also apparent on the third pleopod of the male. In
G. gordonae the endopod is well developed and seven-segmented, extending well
beyond the midpoint of the first exopod segment. The fourth exopod segment is
armed midway along its outer margin with one simple seta. In G. /ongifissura
the endopod is three-segmented and extending about one-third along the length
of the first exopod segment. The fourth exopod segment bears two simple setae
at the proximal end.

The second male pleopod in G. gordonae has an eight-segmented endopod,
while the setae on the outer distal corners of the second to the fourth exopod
segments are modified, being thickened and having their proximal margins
serrated. In G. Jongifissura the endopod is five-segmented, with no modifications
to the outer setae on the exopod segments.

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 325

DISTRIBUTION OF GASTROSACCUS RECORDED IN
SOUTHERN AFRICA

Five species of Gastrosaccus are so far recorded from southern Africa.
G. dunckeri is recorded from the Morrumbene estuary in Mozambique
(Tattersall, O. S. 1958; Day 1974). G. gordonae is also reported from estuaries
on the east coast (Tattersall, O. S. 1952; Scott et al. 1952; Day et al. 1954;
Grindley & Wooldridge 1974), and from Saldanha Bay and Stompneus Bay
on the West Coast (Lazarus 1975). G. brevifissura is recorded from estuarine
and coastal waters (Tattersall, O. S. 1952; Day et a/. 1954; Milland & Harrison
1954; Day 1958; Tattersall, O. S. 1962; Connell 1974; Lazarus 1975; Grindley
1977; Wooldridge 1976, 1977) as well as the intertidal zone (Tattersall, O. S.
1962). G. sanctus has been collected on several occasions off the coast of South
Africa (Tattersall, O. S. 1957; Brown & Talbot 1972; Lazarus 1975).

G. psammodytes is the only species so far reported from sandy beaches in
southern Africa and was until recently collected only in the intertidal and surf
zone (Tattersall, O. S. 1958; Day 1958; Brown & Talbot 1972). McLachlan et al.
(1978) have shown that numbers of G. psammodytes collected at night in the
upper surf zone were significantly lower than numbers collected during the day
and have suggested a general emergence from the sand after dark when the
animals become planktonic in deeper waters. Samples collected at night
(unpublished data) on a number of occasions in Algoa Bay with a WP-2
plankton net have shown them to be present in surface waters 250-300 m
off shore, while Lazarus (1975) collected a number of specimens at night in
vertical plankton samples off the west coast of South Africa. Similar activity
patterns of diurnal burrowing and a nocturnal pelagic life have been shown for
G. sanctus (Bacescu 1934; Moran 1972) and for G. mediterraneus and G. spinifer
(Macquart-Moulin 1977). The most easterly extension of G. psammodytes is
given as Kleinmond (33°33’S) in the eastern Cape Province (Brown & Talbot
1972). During the course of the present study it was collected at Gulu (33°08’S)
and Nahoon (32°59’S) near East London, and at Kei Mouth (32°41’S).

Past workers divided the genus Gastrosaccus into two groups based on the
form of the endopod of the third pleopod of the male. In the Spinifer group the
endopod is multiarticulate, while in the Normani group it is reduced to a single
segment or lacking. All species of Gastrosaccus from southern Africa are
members of the Spinifer group with the exception of G. dunckeri. In this species
the endopod of the third male pleopod is completely lacking.

A further broad separation of the species can be found in the form of the
posterior margin of the carapace. Members of the genus from the southern
African region belong to that group in which the posterior margin of the
carapace is notched, forming an overlapping lobe on either side, or where the
posterior margin of the carapace is produced to form a pair of lappets which
are reflexed forward.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

KEY TO THE SPECIES OF GASTROSACCUS RECORDED IN
SOUTHERN AFRICA

Characteristics common to both sexes are used where possible.

1. Posterior margin of carapace cleft in the median line. Margins on each side of the cleft

produced into two lappets which are reflexed forward . pe

— Posterior margin of carapace forming an overlapping lobe on ‘either side of the emargina-

tion, not reflexed . 3

2. Endopod of third pleopod of the male eight- eeeenicds Se eroup. Telson with five

lateral spines and no spinules between them G. sanctus (van Beneden), 1861

— Endopod of third pleopod of the male lacking; Nocment group. Telson with about ten to

twelve lateral spines. No spinules between them ; G. dunckeri Zimmer, 1915

3. Cleft in telson deep, one-quarter to one-sixth the length of the telson . ‘ . 4
— Cleft in telson shallow, in some specimens little more than an emargination

G. brevifissura Tattersall, 1952

4. Telson with six lateral spines not interspersed with small spinules, or at most one or two

spinules between the larger spines on each side . : G. psammodytes Tattersall, 1958
— Telson with six, seven or eight lateral spines interspersed with at least six to eight small
spinules between the larger spines on each side . ; 5)

5. Telson with eight lateral spines. Spaces between last six pairs of lateral and the terminal
spines occupied with three to six mia Endopod of uropod with thirteen spines among
setae along inner margin 3 t G; gordonae Tattersall, 1952

— Telson with six or seven lateral spines. Spaces between last three pairs of lateral and the
terminal spines occupied with one to five spinules. Endopod of uropod with seven spines
among the setae along the inner margin. : : : 6

6. Telson with a strong spine on each side of the cleft on ‘the dorsal side G. bispinosa sp. nov.
Telson without a spine on each side of the cleft on the dorsal side __G. Jongifissura sp. nov.

ACKNOWLEDGEMENTS

Financial support from the Department of Planning and the Environment
is acknowledged. I also thank Ken McLeod, Ian Davidson, Dan Baird and
Nicky Hanekom for help in the field, and Anton McLachlan who critically read
parts of the manuscript.

REFERENCES

BAcescu, M. 1934. Contribution a’ l’etude des Mysidés de la Mer Noise ainsi que des limans
et des lacs en relation avec la mer avec le Danube. Annis scient. Univ. Jassy 19: 331-338.

Brown, A. C. & TALBOT, M. S. 1972. The biology of the sandy beaches of the Cape Peninsula,
South Africa. Part 3: A study of Gastrosaccus psammodytes Tattersall (Crustacea:
Mysidacea). Trans. R. Soc. S. Afr. 40: 309-333.

CONNELL, A. D. 1974. Mysidacea of the Mtentu River estuary, Transkei, South Africa.
Zoologica afr. 9: 147-159.

Day, J. H. 1958. The Biology of Langebaan Lagoon: a study of the effect of shelter from wave
action. Trans. R. Soc. S. Afr. 35: 475-547.

Day, J. H. 1974. The ecology of Morrumbene estuary, Mocambique. Trans. R. Soc. S. Afr.
41: 43-96.

Day, J. H., MILLARD, N. A. H. & BROEKHUYSEN, G. J. 1954. The ecology of South African
estuaries. Part 4. The St. Lucia system. Trans. R. Soc. S. Afr. 34: 129-156.

GAULD, D. T. & BUCHANAN, J. B. 1956. The Fauna of sandy beaches in the Gold Coast.
Oikos 7: 293-301.

GRINDLEY, J. R. & WOOLDRIDGE, T. 1974. The plankton of Richards Bay. Hydrobiol. Bull. 8:
201-212.

TWO NEW SPECIES OF GASTROSACCUS FROM SANDY BEACHES IN TRANSKEI 327

GRINDLEY, J. R. 1977. The zooplankton of Langebaan Lagoon and Saldanha Bay. Trans. R.
Soc. S. Afr. 42: 341-370.

LAZARUS, B. I. 1975. The inshore zooplankton of the Western Cape. Unpublished Ph.D. thesis,
University of Stellenbosch.

MACQUART-MOULIN, C. 1977. Le contrdéle de émergence et des nages nocturnes chez les
Péracarides des plages de Méditerranée. Eurydice affinis Hansen (Isopoda), Gastrosaccus
mediterraneus Bacescu, Gastrosaccus spinifer (Goés) (Mysidacea). J. exp. mar. Biol. Ecol.
27: 61-81.

McLACHLAN, A., WOOLDRIDGE, T. & VAN DER Horst, G. 1978. Tidal movements of the
macrofauna on a high energy sandy beach in South Africa. J. Zool., Lond. (In press.)
MILLARD, N. A. H. & HARRISON, A. D. 1954. The ecology of South African estuaries. Part 5.

Richards Bay. Trans. R. Soc. S. Afr. 34: 157-179.

Moran, S. 1972. Ecology and distribution of the sand-dwelling mysid Gastrosaccus sanctus
(van Beneden 1961) along the Mediterranean sandy shore of Israel. Crustaceana Suppl. 3:
357-361.

NAKAZAWA, K. 1910. Notes on Japanese Schizopoda. Annotnes zool. jap. 7: 247-261.

Scott, K. M. F., Harrison, A. D. & MAcnéAE, W. 1952. The ecology of South African
estuaries. Part 2. The Klein River estuary, Hermanus, Cape. Trans. R. Soc. S. Afr. 33:
283-332.

TATTERSALL, O. S. 1952. Report on a small collection of Mysidacea from estuarine waters of
South Africa. Trans. R. Soc. S. Afr. 33: 153-188.

TATTERSALL, O. S. 1957. Report on a small collection of Mysidacea from the Sierra Leone
estuary together with a survey of the genus Rhopalophthalamus terrantalis Ulig and a
description of a new species of Tenagomysis from Lagos, Nigeria. Proc. zool. Soc. Lond.
129: 81-128.

TATTERSALL, O. S. 1958. Further notes on the Mysidacea from South African waters. Trans.
esoe. Ss. Afr. 35: 373-383.

TATTERSALL, O. S. 1962. Report on a collection of Mysidacea from South African off-shore
and coastal waters (1957-59) and from Zanzibar. Proc. zool. Soc. Lond. 139: 221-247.

TATTERSALL, W. M. 1927. Report on the Crustacea Mysidacea. Trans. zool. Soc. Lond. 22:
185-199.

TATTERSALL, W. M. & TATTERSALL, O. S. 1951. The British Mysidacea. London: Ray Society.

VAN BENEDEN, P. J. 1861. Recherches sur les Crustacés du littoral de Belgique. Les Mysidés.
Mem. Acad. r. Belg. Cl. Sci. 33: 1-77.

WOOLDRIDGE, T. H. 1976. The zooplankton of Msikaba estuary. Zoologica afr. 11: 23-44.

WOOLDRIDGE, T. H. 1977. The zooplankton of Mgazana, a Mangrove estuary in Transkei,
southern Africa. Zoologica afr. 12: 307-322.

ZIMMER, C. 1915. Die Systematik der Tribus Mysini H. J. Hansen. Zool. Anz. 46: 202-216.

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus ...’; ‘*.. . in C. namacolus (Fig. 10)...’

(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 but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

T. WOOLDRIDGE
TWO NEW SPECIES OF GASTROSACCUS

(CRUSTACEA, MYSIDACEA) FROM SANDY BEACHES
IN TRANSKEI

| OLUME 76 PART 10 SEPTEMBER 1978 ISSN 0303-2515

q

7 e
lh
oA ’
F ANNALS

YF THE SOUTH AFRICAN
MUSEUM

JAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

(a) Centred masthead to consist of
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the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

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‘Smith (1969: 36, fig. 16) describes .

“As described (Smith 1969a, 19695; renee i)
‘As described (Haughton & Broom Ones

‘As described (Haughton et al. 1927) .

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et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).

Examples (note capitalization and punctuation)

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

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. Zen. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19605. 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

(continued inside back cover)

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

Volume 76 Band
September 1978 September
Part “10>” Deel

Saree TERTIARY, MUSTELIDAE
(MAMMALIA, CARNIVORA)
FROM LANGEBAANWEG, SOUTH AFRICA

By

Q. B. HENDEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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

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van stof

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

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

LATE TERTIARY MUSTELIDAE (MAMMALIA, CARNIVORA) FROM
LANGEBAANWEG, SOUTH AFRICA

By
Q. B. HENDEY
South African Museum, Cape Town

(With 11 figures and 10 tables)

[MS. accepted 2 August 1978]

ABSTRACT

The Mustelidae of the latest Miocene/early Pliocene Varswater Formation in ‘E’ Quarry,
Langebaanweg, are described. They are identified as Plesiogulo monspessulanus Viret, 1939,
which is the southernmost record of a wolverine, Mellivora benfieldi sp. noy., which is a likely
ancestor of the living M. capensis, and Enhydriodon africanus Stromer, 1931, which is a
structural and temporal intermediate between E. /luecai and E. sivalensis.

CONTENTS

PAGE
Introduction . , ; ; 2 829
Systematic discussion . : mm ~330
General discussion : : ie Po54
Acknowledgements : : 50
References ; 4 : : ee Se)

INTRODUCTION

Since publication of an account of the Carnivora from the Varswater
Formation in ‘E’ Quarry, Langebaanweg, Cape Province (Hendey 19745), a
great deal of additional material belonging to this order has been discovered
(Hendey 1976b, 1977, 1978c). Although the original species list has not been
substantially altered, the additional material does include specimens belonging
to species not previously known from this locality and some which have led to
revision of earlier identifications. Others have served simply to confirm original
identifications.

Each of these situations applies in the case of the Mustelidae, a family
which is comparatively poorly represented in the ‘E’ Quarry fauna, both in
terms of numbers of species and numbers of specimens. Three species are
recognized. The undescribed mustelid is the first African record of the extinct
wolverine, Plesiogulo (Hendey 1976b: 239), the species previously incorrectly
identified is a honey badger, Mellivora (Hendey 19746: 68), while the species
whose identity is confirmed is an otter, Enhydriodon (Hendey 19746: 72).

The ‘E’ Quarry fauna, of which these three mustelids are a part, has
generally been regarded as early Pliocene (4-5 Ma) in age. This dating is,
however, insecure and present indications are that the outside age limits are
3,5 and 7 Ma, that is, the fauna dates from the very late Miocene and/or early
Pliocene (Hendey 1978c).

229

Ann. S. Afr. Mus. 76 (10), 1978: 329-357, 11 figs, 10 tables.

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

The fossils dealt with in this report are from three distinct stratigraphic
units in the Varswater Formation. They are, in descending order of age, the
Quartzose Sand Member and beds 3aS and 3aN of the Pelletal Phosphorite
Member (Hendey 19765, 1978a). The durations of the intervals which elapsed
between deposition of these units are not known, but they may have been
appreciable. There is evidence, some of which will be presented below, that
certain taxa show evolutionary advances over counterparts from lower units in
the succession, although the differences are relatively slight and generally would
not warrant taxonomic distinction at species level.

The material described is housed in the South African Museum, and
catalogue numbers are prefixed SAM-—PQ-, which identifies the institution and
department concerned. This lettering is omitted from the text.

SYSTEMATIC DISCUSSION

Family Mustelidae
Subfamily Mellivorinae
Plesiogulo monspessulanus Viret, 1939
Material

L21570. Remains of an adult individual including: incomplete right
mandible with P; to M,; right I, and I’; left I, and I?. Part of right pes including
most tarsal bones, and metatarsals I to V lacking distal ends.

L40042. Remains of an adult male including: fragmented and incomplete
skull and mandible with right P! to P*, right P, to P,, several worn incisors and
fragments of other teeth. Elements of the vertebral column, limb girdles and all
four limbs, most larger bones being incomplete. ss

L28394. Left mandible fragment with M, and M,.

Locality and horizon

Varswater Formation, “E’ Quarry, Langebaanweg. L21570 and L28394 are
from the Quartzose Sand Member; L40042 is either from the Quartzose Sand
Member or the lowermost level of bed 3aS of the Pelletal Phosphorite Member.

Age
Langebaanian (latest Miocene/early Pliocene), between 3,5 and 7 Ma.

Description

The most striking characteristic of this species is its large size. It was
apparently only a little smaller than Megalictis ferox of the North American
Miocene (Matthew 1907), which was the ‘largest of all mustelids [reaching] the
size of a black bear’ (Kurtén 1971: 119). The Langebaanweg species is generally
similar to the living wolverine, Gulo gulo, in terms of its dental and osteological
characters.

The skull of L40042 was badly damaged by a mechanical excavator and
useful observations can be made only on the upper and lower premolars, and

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 331

parts of the braincase, right maxilla and left mandible.

The maxillary fragment is comprised only of the bone immediately adjacent
to the four premolars (Fig. |). Parts of the alveoli of the C and M’ are preserved.
The premolars are large and relatively broad compared with those of living

FULT ULLS LALLA ILL HLTH LAL LLL

Fig. 1. Buccal and occlusal views of Plesiogulo monspessulanus maxilla
(40042) from Langebaanweg.

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1
Dimensions of the teeth of Gulo gulo and Plesiogulo species.

Pp P EE

Gulo gulo—Scandanavia (n = 11—15)*

Plesiogulo ‘major’ —Chinat

Plesiogulo monspessulanus — Europet

Plesiogulo monspessulanus—Langebaanweg L40042
L21570
L28394

Plesiogulo crassa—China (means)} .

* Data provided by E. Anderson (Denver, Colorado).
+ Kurtén 1970.

Gulo gulo (Table 1). They are positioned as in the living species, except that P?
and P® overlap, which is unlike the fore-aft arrangement in the two available
G. gulo comparative specimens and some illustrated examples (e.g. Anderson
1977, fig. 4; Novikov 1962, fig. 128; Kurtén & Rausch 1959, figs 3-4).
Morphologically the teeth are similar to those of G. gu/o in all observable
respects. The infraorbital foramen of L40042 is more posteriorly situated than
that of G. gu/o. It opens above the apex of the P* paracone, whereas in the living
species the opening is above the anterior end of P*.

The features of the posterior parts of the skull of L40042 (Fig. 2) are
essentially similar to their counterparts in G. gulo. The fossil skull is, however,
very large (Table 2), with processes and crests greatly exaggerated. The sagittal
crest is particularly prominent, being about 20 mm high along that part which is
preserved. This crest terminates posteriorly above the level of the occipital
condyles, whereas in G. gu/o it usually, or always, projects further back. By
contrast, the paroccipital process of the fossil projects further back than that of
G. gulo.

The less posteriorly protruding sagittal crest is one manifestation of the
relatively shorter braincase of the fossil. The distance between the external
auditory meatus and most posterior part of the sagittal crest is about
30 mm in L40042, whereas in two G. gulo specimens it is 36 mm
(SAM-ZM36095 3) and 40 mm (SAM-ZM38641 9). This difference is also
indicated by the orientation of the nuchal crest in lateral view. In L40042 the
lower part of this crest is at an angle of about 70° above the horizontal, whereas
in the G. gulo comparative specimens the angle is about 50°.

In G. gulo the zygomatic process of the squamosal rises vertically above the
level of the glenoid fossa so that the inferior margin of this process is elevated

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 333

Pt P, EE P, M, M;

b l b l b l b l b l b

s =6s - 11,0- | 5,7- _ 7,7— 5,3— | 10,5- 6,2— | 19,7- 8,4— 5,0- 4,3-
) 12,8 6,8 9,6 6,6 12,4 7,8 ss) 10,0 6,5 355
—— 8,9 — [255 — 14,3 —- 30,5 ie — —

— _ 10,0 — 14,0 — 28,0 10,5 — —
eei5,6 | 9,1 Osi \wl256 8,1 16,0 Oe — — — =

— — — 11,4 Tone liss2 oF 28,3 CHRO | 11,4 c8,5

— — — — — ay — — (Os WETS) 11,4 Gale COs

12,9 7,4 59 9,8 a 1350 7,8 2359 9,6 Ted 6,6

well above the glenoid fossa and external auditory meatus. In L40042 the
process is much less elevated and its inferior margin is actually below the level
of the auditory meatus.

Lower teeth of all three fossil individuals are preserved and the only
elements not represented are I, to P,. The lower teeth, like the uppers, are
generally similar to their counterparts in G. gulo, although they are larger
(Table 1, Fig. 3). The P, of L40042 differs from its counterpart in G. gulo in
being single-rooted, a possibly significant character from a phylogenetic point
of view and one which will be discussed later.

The M,’s of both L21570 and L28394 are damaged and it cannot be
established for certain whether or not a metaconid was present. Judging from
the less damaged M, of L28394, this cusp was either very small or absent. The
M, of the fossil does differ from that of G. gulo in having a relatively longer
talonid.

The M,’s of L21570 and L28394 are strikingly different in both size and
morphology. The M, of the latter is slightly damaged, but it was evidently
similar to that of G. gu/o in proportions. By contrast, the M, of L21570 is
anteroposteriorly elongated, with the trigonid and talonid distinguishable,
although none of the individual cusps is prominently developed. The elongated
M, of L21570 may be an individual peculiarity, rather than being typical of the
Langebaanweg species. The M, : M, length ratio in L28394 (1 : 0,27) is com-
parable to the ratio in both living G. gulo (1 : 0,28 — n = 2) and the relatively
primitive late Miocene Plesiogulo crassa (1 : 0,33—Kurtén 1970), whereas the
ratio in L21570 is quite distinct from either (1 : 0,40).

The mandible of L21570 lacks much of the coronoid process and symphyseal
region (Fig. 3). The latter region was affected by a pathological condition

5

a ETT TOT a Bt Raat

ye
Avy

332 ANNALS OF THE SOUTH AFRICAN MUSEUM LATE TERTIARY MUSTELIDAE FRO
4 M LANGEBAANWEG
, SO
TABLE 1 UTH AFRICA 333
Pt Pp Pp pt P, P
—=1 4 oa bh I b M,
Gulo gulo—Scandanavia (n = 11—15)* pee Serge wes 9,6- 57a) 11,0- | 5,7- = ii Bsa \ AOS Gai alge :
4, 7,2 11,2 6,9 2 12,8 6,8 9,6 66 124 = 19,7- 84 ara =
: - =I A > 7,8 DRM, 10.0 65 43-
Plesiogulo aor Ghats Ne — — == ae as Es a, 8,9 a 12,5 = me 0 | 6 55
—t— Sate = — Sa dL ? — 30,5 11,3 a
Plesiogulo monspessulanus— Europet = = ak = — = = ee — | 10,0 = 14.0 5 ve
1, : ele ee ho es
Plesiogulo monspessulanus —Langebaanweg 140042 Grose 6,0 9,7 Te) 13,9 9,082 15,6 | 9,1 Bu Mes a1 ia =e a i
if > , = = =
(= : = =
1.21570 — — — = = . 2 = aS os _
| A 11,4 7,3 | 15,2 92 283 ae
128394 = oe = | ae | az =e a ; cll,0 | 11,4 85
— a = = i Gc DTS: 11,4 5 —
Plesiogulo crassa—China (means)t . 7] { Cota COT

* Data provided by E. Anderson (Denver, Colorado).
+ Kurtén 1970.

Gulo gulo (Table 1). They are positioned as in the living species, except that i
and P? overlap, which is unlike the fore-aft arrangement in the two available
G. gulo comparative specimens and some illustrated examples (e.g. Anderson
1977, fig. 4; Novikov 1962, fig. 128; Kurtén & Rausch 1959, figs 3-4).
Morphologically the teeth are similar to those of G. gulo in all observable
respects. The infraorbital foramen of L40042 is more posteriorly situated than
that of G. gulo. It opens above the apex of the P* paracone, whereas in the living
species the opening is above the anterior end of P*.

The features of the posterior parts of the skull of L40042 (Fig. 2) are
essentially similar to their counterparts in G. gulo. The fossil skull is, however,
very large (Table 2), with processes and crests greatly exaggerated. The sagittal
crest is particularly prominent, being about 20 mm high along that part which is
preserved. This crest terminates posteriorly above the level of the occipital
condyles, whereas in G. gulo it usually, or always, projects further back. By
oe the paroccipital process of the fossit projects further back than that of

. gulo.

The less posteriorly protruding sagittal crest is one manifestation of the
relatively shorter braincase of the fossil. The distance between the external
auditory meatus and most posterior part of the sagittal crest is about
30 mm in 140042, whereas in two G. gulo specimens it is 36 mm
(SAM-ZM36095 3) and 40 mm (SAM-ZM38641 9). This difference is also
indicated by the orientation of the nuchal crest in lateral view. In L40042 the
lower part of this crest is at an angle of about 70° above the horizontal, whereas
in the G. gulo comparative specimens the angle is about 50°.

In G. gulo the zygomatic process of the squamosal rises vertically above the
level of the glenoid fossa so that the inferior margin of this process is elevated

r)
5

well above the glenoid fossa i
process is much ie elevated aad cae eee seein
ce eee margin is actually below the level

Lower teeth of all three fossil individuals é
elements not represented are I, to P,. The inet ea ae ee oe
generally similar to their counterparts in G. gulo, Sioueh they 2 large
(Table 1, Fig. 3). The P, of L40042 differs from its counterpart in G. gulo in
being single-rooted, a possibly significant character from a phylogenetic point
of view and one which will be discussed later.

The M,’s of both L21570 and L28394 are damaged and it cannot be
established for certain whether or not a metaconid was present. Judging from
the less damaged M, of L28394, this cusp was either very small or absent. The
M, of the fossil does differ from that of G. gulo in having a relatively longer
talonid.

The M,’s of L21570 and 128394 are strikingly different in both size and
morphology. The M, of the latter is slightly damaged, but it was evidently
similar to that of G. gulo in proportions. By contrast, the M, of L21570 is
anteroposteriorly elongated, with the trigonid and talonid distinguishable,
although none of the individual cusps is prominently developed. The elongated
M, of L21570 may be an individual peculiarity, rather than being typical of the
Langebaanweg species. The M, : M, length ratio in L28394 (1: 0,27) is com-
parable to the ratio in both living G. gulo (1 : 0,28 — n= 2) and the relatively
Primitive late Miocene Plesiogulo crassa (1 - 0,33—Kurtén 1970), whereas the
ratio in L21570 is quite distinct from either (1 : 0,40).

The mandible of L21570 lacks much of the coronoid process and symphyseal
region (Fig. 3). The latter region was affected by 4 pathological sant

ANNALS OF THE SOUTH AFRICAN MUSEUM

334

1

oir s}t et zr ut oT j
hurd uit

6

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 335

TABLE 2

Dimensions of the skull and mandible of the Langebaanweg Plesiogulo monspessulanus and
living Gulo gulo.

\
Plesiogulo monspessulanus Gulo gulo
L40042 L21570 L28394 ZM36095 ZM38641
Braincase length (postorbital con-
striction to occipital condyles) 81,0 — — 69,5 65,5
Width across occipital condyles . 50,8 — — 37,8 S557
Mastoid width SE Me eetiNE Ry Ake 108,2 — — 88,8 81,4
Zygomatic width EME ails: 158,5 = = 103,7 95,0 :
Occiput height (occipital condyles
to top of sagittal crest) . : 80,5 — — 50,9 52,6
Depth of mandible below M, C2590 39,0 37,5 23,3 223
Height of ascending ramus (angle i
to top of coronoid process) . 68,7 — — 50,9 47,1
Transverse diameter of condyle . 44,1 41,4 — SUG 24,7
d

\
TA LULU LL nan LUI a
Fig. 3. Occlusal and buccal views of Plesiogulo monspessulanus mandible
(L21570) from Langebaanweg.
N

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

which has left the remaining bone spongy in texture. The abnormality extends
along the alveolar margins of the cheekteeth, becoming less pronounced
posteriorly. The area of insertion of the occipito-mandibularis muscle in the
subangular region is marked by V-shaped ridges of bone which diverge
posteriorly, that on the buccal side being more prominent and irregular in
outline. The counterparts of these ridges in L28394 and L40042 are only
slightly developed, although the latter apparently belonged to a more aged
individual, judging from wear on the cheekteeth. The ridges in L21570 may be
a further manifestation of the mandibular pathology of the individual
concerned.

The fossil mandibles are generally similar to corresponding parts of the
mandibles of the G. gu/o comparative specimens, except that the condyles are
remarkably long and tubular in shape (Table 2, Fig. 4).

A feature of the dentition of L21570 which is evidently related to the
pathological condition of the mandible is that, although the cheekteeth are only
slightly worn, the preserved incisors are well worn. Similar pathology in one of
the G. gulo comparative specimens (SAM-—ZM38641) is accompanied by broken
canines and heavily worn incisors. According to E. Granqvist of the Zoological
Museum of the University, Helsinki (letter to R. Rau), damaged symphyseal
teeth, and pathology of the adjacent parts of the jaws, are not uncommon in
wolverines. Presumably this results from the aggressive behaviour and indis-

HUH

mm

mm

qn

Fig. 4. Posterior view of Plesiogulo
monspessulanus mandible (L40042)
from Langebaanweg.

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA Sai)

criminate feeding habits of the species. The condition of L21570 suggests that
the late Tertiary wolverine from South Africa had habits similar to its living
relative.

Ten of the vertebrae of L40042 are reasonably intact and all are distinguished
from their modern counterparts only by their very much larger size. In general
this also applies to other known postcranial bones of the fossil (Figs 5-6,
Tables 3-6). The limb bones, and particularly the metapodials, are relatively less
elongated, but much more stoutly proportioned than the corresponding bones
in G. gulo. These differences, and possibly all others, are presumably due to the
size difference between the species concerned. For example, the humerus of
L40042 has a much deeper supratrochlea fossa and longer lateral condyloid
crest, which reflect the stoutness and more heavily-muscled state of this bone.
One of the differences for which no explanation can be offered is the marked
curvature of the radius and ulna of L40042 (Fig. 6).

The relatively short and stout metapodials of the fossil go together with
differences in proportions of certain carpal and tarsal bones. For example, the
fossil caleanea and astragali are also relatively short and stout. Possibly the
fossil species was more perfectly plantigrade than the living wolverine.

Postcranial bones of both L40042 and L21570 show signs of an arthritic
condition, something which is not uncommon amongst the fossil carnivores
from Langebaanweg (Hendey 19745).

Fragments of the baculum of L40042 are preserved. They are similar in
shape to corresponding parts of the baculum of G. gulo.

Discussion

The Mellivorinae are known from late Tertiary and Quaternary contexts
through much of the Old World and North America. They are a heterogeneous
group for which a subdivision into tribes has been suggested, and Webb (1969)
visualized the subfamily as follows:

Mellivorini Gulonini Brachypsalini
Aelurocyon Peterson, Ischyrictis Helbing, Paroligobunis
1906 1930 Peterson, 1906
Megalictis Matthew, Hadrictis Pia, 1939 Brachypsalis Cope,
1907 Plesiogulo Zdansky, 1890
Perunium Orlov, 1947 1924 Brachypsaloides Webb,
Eomellivora Zdansky, Gulo Frisch, 1775 1969
1924

Promellivora Pilgrim, 1932
Mellivora Storr, 1780

At least one additional genus belonging to this subfamily (Ferinestrix
Bjork, 1970) has since been described, while Promellivora has been regarded as a
synonym of Mellivora (Hendey 19746). The latter opinion may not have been
warranted (see below). 4

ANNALS OF THE SOUTH AFRICAN MUSEUM

int

4

|

2

iii

:

il
2

wn

Fig. 5. Plesiogulo monspessulanus from Langebaanweg and Gulo gulo (ZM38641).
A. Humeri. B. Scapholunars. C. Pisiforms. D. Astragali. All specimens of P. monspessulanus
belong to L40042, except the left astagalus which is of L21570.

339

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA

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340 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Dimensions of Langebaanweg Plesiogulo monspessulanus and living Gulo gulo carpal and
tarsal bones.

Plesiogulo
monspessulanus
ZM38641 | L40042 L21570

Max. transverse diameter. ; ; 2355 =

Max. diameter proximal end to distal
end : : A Gee. ee : : 11,0 14,4
Max. ant.-post. diameter, articular end a

Max. transverse diameter, articular end 11,9 10,5 =

SCAPIO-
LUNAR

PISIFORM

. Maxolengih’ 02) 0) eae 55,8
(=)

; Max. transverse diameter. ‘ ; [a [a 33,2
z

° Max. dorsoventral diameter . : ; 22,4 —

va Mam tenet ei eee 26,8 36,1 36,4
&

26 Max. transverse diameter, tibial facet 14,9 | 331 ea 2 20,0
& Max. length (dorsoventral diameter) . = 22,3
¢ Max. transverse diameter . . . a) 18,9

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LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 343

With the exception of Mellivora and Gulo, all the recorded genera are late
Tertiary in age and they are generally not well represented. As a result their
inter-relationships are somewhat obscure. Webb (1969: 66) concluded that there
was apparently ‘an early Miocene radiation of mellivorine stock which gave
rise to the Aelurocyon—Megalictis and Paroligobunis—Brachypsalis—Brachyp-
saloides lineages in North America and the Jschyrictis—Hadrictis—Plesiogulo
lineage in Europe’. Later Mellivorini were confined to the Old World. .

Plesiogulo, the genus with which the Langebaanweg wolverine is identified,
was widespread in Eurasia during the late Miocene and Pliocene (Kurtén 1970),
and was also present in North America (Kurtén 1970; Bjork 1970). Its presence
at Langebaanweg near the southern tip of Africa means that it must have been
distributed over much of this continent as well. Both Kurtén (1970) and
Anderson (1977) regard Plesiogulo as the ancestor of Gulo.

Until recently large mellivorines were not known from Africa, but now, in
addition to the Langebaanweg species, there is an as yet unidentified species
recorded from the Omo Group deposits in Ethiopia (Howell & Petter 1976).
The relationships between the Langebaanweg and Omo species have yet to be
determined.

Of the described species of Plesiogulo, the one from Langebaanweg most
closely resembles the broadly contemporaneous P. monspessulanus from Europe
and P. major from China. These Pliocene species are the largest and most
recent representatives of the genus. They are distinguished from one another
only by the fact that P. major has an M, metaconid and Kurtén (1970: 12)
concluded that they ‘were obviously closely related’. They should, perhaps,
be regarded as conspecific, with P. monspessulanus being the senior synonym.
If this step is taken, then it becomes immaterial whether or not the Langebaan-
weg species had an M, metaconid, since there are no obvious grounds for
separating it from its European and Chinese counterparts and all three may be
identified with P. monspessulanus.

Geographical factors alone are sufficient to preclude the Langebaanweg
P. monspessulanus from the role as direct ancestor of Gulo. In addition, it was
noted above that the only preserved P, of this species is single-rooted and if this
was characteristic of the Langebaanweg population, rather than just an
individual anomaly, then it, too, indicates the lack of a direct phylogenetic
connection with Gu/o, in which P, is double-rooted. The P, of the Eurasian
populations of P. monspessulanus has yet to be recorded.

Mellivora benfieldi sp. nov.
Holotype

L42838. Right mandible fragment with C and P, to M,.

Referred material

L6385. Left mandible fragment with P, and M, (Hendey 19746: 68-72,
fig. 6).

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

L31273. Right mandible fragment with M,.

L50443. Right mandible fragment with P, and M,.

L50541. Right Mt?.

A left ulna (L40080) and right radius (L45384) are tentatively assigned to
this species.

Other poorly preserved and/or fragmentary specimens were excluded from
the present study.

Locality and horizon

Varswater Formation, ‘E’ Quarry, Langebaanweg. The holotype (L42838),
L6385 and L40080 are from bed 3aS, and the remaining specimens are from
bed 3aN, both units of the Pelletal Phosphorite Member.

Etymology

Named for Graham Benfield, formerly geologist and mine superintendent
at the Chemfos Ltd mine at Langebaanweg, whose contributions to the
Langebaanweg Research Project were of inestimable value.

Diagnosis

A species of Mellivora a little smaller than the extant M. capensis; P, and
M, absent; P, to M, relatively narrow; principal cusps of P, to P, sharp-pointed,
with sharp anterior and posterior keels; anterior and posterior accessory cusps
of P, relatively small; M, talonid relatively short, narrow and sectorial; internal
lobe of M?! only slightly expanded, lacking prominent cingulum round the
protocone; mandibular condyle not elevated above cheekteeth.

Age
Langebaanian (latest Miocene/early Pliocene), between 3,5 and 7 Ma.

Description

L6385 has already been described and discussed in detail (Hendey 19745),
and the paragraphs which follow summarize and supplement the earlier account
of the Langebaanweg Mellivora.

The lower canine and cheekteeth of Mellivora benfieldi are superficially
similar to those of the living honey badger, M. capensis. They differ in being
smaller and narrower (Table 7), with cusps more sharp-pointed and keels more
sharp-edged (Fig. 7). In these respects M. benfieldi is clearly the more primitive
(less specialized) of the two species. They resemble one another in the number
and arrangement of the teeth in the jaw and in the cusps on individual teeth.
The P, accessory cusps are, however, smaller in M. benfieldi, while the M,
talonid is smaller, narrower and without the basin-shaped depression situated
lingually in M. capensis.

The internal (lingual) lobe of the isolated M! (L50541), like the M, talonid,
is relatively small (Fig. 7). Its length (5,2 mm) is only slightly greater than that

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 345

of the buccal lobe (4,2 mm), whereas in M. capensis the corresponding figures
are 8,3 and 4,4 mm (n = 8). The expansion of the internal lobe in the M! of
M. capensis is due to the development of a prominent cingulum round the
protocone, a feature which is lacking in L50541. The fossil tooth is also narrower

qn TELLIER LLL IL

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Fig. 7. A. Occlusal and buccal views of Mellivora benfieldi holotype
(L42838). B. Occlusal view of M? (L50541). Both from Langebaanweg.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 347

than the M! of M. capensis, measuring 9,5 mm as against the mean of 10,9 mm
in the comparative series.

The inferior margins of the fossil mandibles are slightly convex, whereas
in the M. capensis comparative series they are straight or slightly concave. In
both species the number and position of mental foramina are variable. In
M. benfieldi there is only a slight elevation of the inferior margin towards the
angle and the condyle is not raised above the level of the cheekteeth as in
M. capensis. The larger of the fossil mandibles presumably belonged to males
and are comparable in size to the smaller mandibles in the M. capensis com-
parative series, which belong to females.

The ulna, L40080 (Fig. 8), and radius, L45384, are the only postcranial
bones in the Langebaanweg fossil assemblage which have so far been tentatively
assigned to Mellivora. They are essentially similar to their counterparts in
M. capensis, but are relatively short and stout (Table 8). Their size is in keeping
with that of the lower jaws described above.

Discussion

It was earlier concluded that the ‘E’ Quarry Mellivora was closely related to,
and possibly conspecific with, the Indian Mio/Pliocene M. punjabiensis (Hendey
1974b). It is now clear that the two species are not conspecific. M. benfieldi
differs in having a smaller canine and shallower mandibular corpus, it lacks P,
and has a narrower P,.

The loss of P, is an advanced character in mellivorines and in this respect
at least, M. benfieldi is closer to M. capensis. Since the only known canine of
M. benfieldi is probably that of a female (L42838), while the M. punjabiensis
holotype could be a male, the canine size difference is not necessarily as great as
would appear at first sight. Nevertheless, the difference is probably more than
would be encountered in a single species. The large canine and relatively deep

Fig. 8. Ulnae of Mellivora benfieldi (L40080) from Langebaanweg
and M. capensis (ZM36867).

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LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 349

mandibular corpus may also be primitive characters in M. punjabiensis, since
they are features of Eomellivora, the genus from which M. punjabiensis may have
been derived. The relatively short and broad P, of the latter is, however, more
specialized than that of M. benfieldi.

To sum up, M. punjabiensis is an inappropriate structural ancestor for
M. benfieldi and there was probably no direct phylogenetic connection between
them. On the other hand, M. benfieldi is suitable in all observable respects to be
ancestral to M. capensis. It follows that the suggestion, first made by Pilgrim
(1932), that M. punjabiensis may have been an early ancestor of M. capensis,
can now be dismissed. It also follows that the suggestion that the original
separate generic status of ‘M.’ punjabiensis was unwarranted (Hendey 19745),
is no longer acceptable. This species should once again be identified as
Promellivora punjabiensis. The origins of Mellivora will be discussed again below.

The M. benfieldi sample is divisible into two units on stratigraphic grounds.
The holotype (L42838) and L6385 are from bed 3aS of the Pelletal Phosphorite
Member, while L31273 and L50443 are from the stratigraphically higher (and
younger) bed 3aN. It was mentioned earlier that the interval between deposition
of these beds may have been appreciable and that evidence exists of evolutionary
changes in taxa common to both.

This applies in the case of M. benfieldi, since the P,’s and M,’s of the
bed 3aN specimens are intermediate in breadth between those of the bed 3aS
specimens and modern M. capensis (Table 7). In the case of M,’s there are
overlaps in the ranges of the three samples, but the mean figures show a breadth
increase from the 3aS sample (1 : 0,45), through the 3aN sample (1 : 0,47), to
M. capensis (1 : 0,49). The differences are thus in the expected order given the
relative ages of the three samples. Although the mean values for both P, and M,
are discrete, the differences are small and appreciable overlaps in ranges would
be expected of larger samples.

It is worth noting here that the P, length : breadth ratio in Promellivora
punjabiensis (1 : 0,74) sets this species apart from the M. benfieldi-M. capensis
combination.

The new Mellivora specimens from ‘E’ Quarry confirm the earlier conclusion
that the ‘E’ Quarry species is definitely not conspecific with the Mel/livora from
the nearby Baard’s Quarry (Hendey 19746, 1978a). The latter is clearly more
advanced and closer to, if not conspecific with, M. capensis.

Subfamily Lutrinae
Enhydriodon africanus Stromer, 1931

Material

L9138. Right mandible fragment with part of P, (Hendey 19746: 72-74,
fig. 7).

L50000. Left-mandible fragment with P, to M, and isolated left P*.

Various postcranial bones, including a femur (L41523), distal radii (L50001)
and an astragalus (L50117), are tentatively assigned to this species.

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality and horizon

Varswater Formation, “E’ Quarry, Langebaanweg. L9138 and L41523 are
from bed 3aS, and the remaining specimens are from bed 3aN, both units of the
Pelletal Phosphorite Member.

Age
Langebaanian (latest Miocene/early Pliocene), between 3,5 and 7 Ma.

Description

The mandible L9138 was eae earlier and identified with Enhydriodon
africanus, a species otherwise known only from Kleinzee, which is also on the
west coast of the Cape Province, but about 400 km north of Langebaanweg
(Hendey 19746).

The new mandible, L50000, is a better specimen than L9138 and has
served to confirm the identification with E. africanus, since it is very similar to
the type specimen of this species. It lacks P,, has a single-rooted P, and a
double-rooted P;. The P, has a principal cusp, a posterior accessory cusp and a
prominent cingulum round its circumference. The P, of L9138 is higher crowned
and has a more prominent posterior accessory cusp than those of L50000
(Fig. 9) and the E. africanus holotype. The significance of this difference will be
discussed below.

The M, of L50000 is large, with the paraconid, acoiieeeen and metaconid
more or less equally developed (Table 9, Fig. 9). These cusps are low-crowned
and bulbous. The talonid is large and basin-shaped, with the hypoconid
covering about half its area. There is a prominent cingulum encircling the
paraconid and extending posteriorly to the talonid on the buccal margin of the
tooth. The M, is single-rooted and slightly elongated transversely, with little
relief on the occlusal surface.

The isolated P* (Table 9, Fig. 10), which evidently belongs to the same
individual as the new mandible, is an important specimen since the P* of
E. africanus was not previously known. It has a small parastyle, the paracone is
the most prominent cusp, the metastyle is very short, the protocone is large and
nearly as prominent as the paracone and it is flanked posteriorly by a large, but
low hypocone. There is a prominent cingulum encircling much of the tooth.

The various postcranial bones tentatively identified with FE. africanus are
similar to corresponding bones of the living clawless otter, Aonyx capensis. They
are distinguished principally by their larger size. For example, the femur,
L41523 (Fig. 11), has an overall length of 165 mm, compared with a mean
length of 114 mm in a series of four A. capensis specimens.

Discussion

Repenning (1976) has dealt in detail with recorded representatives of
Enhydriodon and concluded that there were two late Tertiary lineages of this
genus. One led to E. sivalensis and ‘can be characterized by the presence of a

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 551

nA Mm nn

mn

TN

nm

HH

Fig. 9A. Occlusal and buccal views of Enhydriodon africanus mandible
(L50000). B. Buccal view of E. africanus mandible (L9138) (reversed).
Both from Langebaanweg.

parastyle on P* and by the location of the protocone of this tooth which is
located as far lingually as the hypocone’ (Repenning 1976: 305). The second
lineage led to the living sea otter, Enhydra lutris.

On the basis of the P* characters, FE. africanus evidently belongs with the
group which includes E. sivalensis, a conclusion already reached by Repenning
(1976) on other evidence. E. sivalensis is the more advanced of the two species,
since it has a broader P* (Table 9), with a more quadrate outline due to a greater
development of the hypocone. In addition, the cheekteeth of EF. africanus
apparently have more strongly developed cingula, which is a primitive condition
in the Enhydriodon/Enhydra group.

E. africanus is here regarded as a structural and temporal intermediate

ANNALS OF THE SOUTH AFRICAN MUSEUM

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bulbous, or mastoid, tooth cusps’, which are characteristic of Enhydra
(Repenning 1976: 306). The fact that the P, of the bed 3aN L50000 has lower
crowned and more bulbous cusps than the bed 3aS L9138 may therefore be
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of Aonyx capensis (ZM36254).

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

interval between deposition of beds 3aS and 3aN. The bed 3aN E. africanus is
apparently morphologically closer to the Kleinzee representative of this species
than is the one from bed 3aS.

GENERAL DISCUSSION

The Mustelidae comprise a relatively small element in the carnivore fauna
of the Varswater Formation, being made up of only 3 of the approximately 29
recorded species (Table 10). In terms of the numbers of specimens known they
form an almost insignificant part of the assemblage of carnivore material.
Further collecting, and analysis of collected specimens, is unlikely to change this
pattern of representation. Mustelids are a relatively uncommon element in the
Pleistocene and Recent faunas of Africa as well.

The presence in the Langebaanweg fauna of an otter and a honey badger
is not surprising, since both have counterparts in the modern fauna of the
region, and Pleistocene representatives of these animals are also known (Hendey
1974b). The post-Pliocene species concerned are Aonyx capensis and Mellivora
capensis, both of which were first recorded from the ‘Cape of Good Hope’
(Ellerman ef al. 1953), the type specimens probably having come from the
region in which Langebaanweg is located.

The only other mustelid, living or fossil, from this region is the musteline,
Ictonyx striatus. It is still one of the more commonly occurring small carnivores
in the vicinity of Langebaanweg. Small carnivores are well represented in the
Varswater Formation fauna, but all are viverrids and the apparent absence of
even one musteline is notable.

Wolverines are today known only from Arctic and sub-Arctic regions, so
the presence of a fossil wolverine at Langebaanweg at about 33°S would seem
at first sight to be extraordinary. It is, however, but one of several species in the
Varswater Formation which have living counterparts on continents other than
Africa. Other examples are a bear (Hendey 1972, 1977) and a peccary (Hendey
1976a). Such species belong to groups which had a much wider distribution in
the Old and New Worlds during the late Tertiary than was the case subsequently.
There was evidently appreciable faunal interchange between Africa and Eurasia
at certain times during the late Tertiary. Greater climatic uniformity and
differences in the pattern of zoogeographic barriers contributed to the existence
of a more widespread and cosmopolitan fauna at that time.

The wolverines of that period were obviously not inhabitants of cold
regions as they are today, and no satisfactory explanation can be offered for
their failure to maintain their position other than in northern high latitudes.
The living wolverine is a remarkably strong and resourceful animal, and one of
the more extreme examples of an opportunistic feeder. It is difficult to conceive
of its large late Tertiary forebears becoming extinct through unsuccessful
competition with other carnivores or because of a decline and extinction of a
preferred prey species. Similarly, since the wolverine today flourishes in the
most rigorous of climates, the climatic deterioration late in the Tertiary, and

a

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 355

subsequently, is unlikely to have adversely affected its viability. The failure of
the bear-like wolverine, and of true bears, to survive in Africa is one of the more
curious aspects of the later history of mammals on this continent.

No such problems exist in the case of the honey badgers. Mellivora capensis
is still found over much of Africa and parts of southern Asia (Dorst & Dandelot

TABLE 10
Carnivora of the Varswater Formation, Langebaanweg.

Quartzose Pelletal Phosphorite
Sand Member
Member bed3aS __ bed 3aN

Canidae

Vulpes sp. : 5 : : aioe ae! rere: x x
Ursidae

Agriotherium africanum : : : : : x x
Mustelidae

Plesiogulo monspessulanus . ; ; : : ; x ?

MRCITORGDCHIICIAE 4k a we Le x x

agpvyariodon Gfricanus . - = s «= «. « x x
Phocidae

Prionodelphis capensis . : ; P : ; x x x
Viverridae

Viverra leakeyi x » S<

Genetta sp. x

Herpestes sp. A x x

Herpestessp.B x Xx

Herpestinae spp. C, D, E : ‘ : : ; x

Herpestinae (not studied) . : : : : : Ss x
Hyaenidae

Adcrocuta australis : : : : : x ? q

Ictitherium preforfex . ; ‘ , ~ x

Hyaenaabronia . : : : z : < x x

Hyaenictitherium AIRE : x

Euryboas sp. . ; : : : ; : ‘ : x x x

Hyaenidae sp.E . : : 3 : : : x

Hyaenidae (not studied) : : ‘ : : ; x K
Felidae

‘“Machairodus’ sp. ‘<

Homotherium sp. x x

Felis sp. (small) x

Felis aff. issiodorensis x x

Felis obscura : : : : E x

Dinofelis aff. Perse ; P : : : : x ~ x

Felidae (not studied) sx x

Unclassified Carnivora

Gen. et sp. indet. (Canidae or Viverridae) 3 : x
Gen. et sp. indet. (?Procyonidae) : ‘
Gen. et sp. indet. (?Lutrinae) : . , 3 x

Gen. et sp. indet. (?Otariidae) . : ‘ S<

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

1970), although it is probably nowhere common. Mellivora has a poor fossil
record, but the genus has probably had an uninterrupted tenure in Africa since
late in the Miocene. Apart from the Langebaanweg species and Pleistocene
representatives of the genus in South Africa (Hendey 1974a, 19745), there is an
approximately 10 m.y. old Mellivora recorded from the Ngorora Formation in
Kenya (Bishop & Pickford 1975). The Ngorora species is undescribed, but it is
apparently the earliest record of the genus anywhere. This, together with the
suggestion made above that Asian Eomellivora and Promellivora were not
directly related to Mellivora, suggests that the latter genus may have had its
origins in Africa. The ultimate origins of Mellivora are obscure, but there is
now a middle Miocene mellivorine recorded from South West Africa (Hendey
19786), which probably is a suitable structural ancestor. This animal, which is
tentatively identified with Ischyrictis, might also be ancestral to Plesiogulo.

Unlike the situation with Plesiogulo, the extinction of Enhydriodon in the
late Pliocene or early Pleistocene could be explained by unsuccessful competition.
The appearance in Africa of both Lutra and Aonyx was apparently more or less
coincident with the disappearance of Enhydriodon. Otters are, however, not
common as fossils in Africa and much has yet to be learnt of the history of the
subfamily on this continent.

The ‘E’ Quarry mustelids provide little information on the problem of the
age of the Varswater Formation. The three species are consistent with a
post-‘Pikermian’ and pre-‘Villafranchian’ age, but only when their broadly
contemporaneous counterparts from securely dated contexts elsewhere are
better known will they be useful for relative dating purposes.

On the other hand, Mellivora benfieldi and Enhydriodon africanus are
significant in providing evidence that there was an appreciable time interval
between deposition of beds 3aS and 3aN of the Pelletal Phosphorite Member.
Other supporting evidence has yet to be published and it is intended to delay a
detailed consideration of this matter until more of the taxa common to these
units have been studied. There is little prospect yet of being able to establish the
duration of the time interval, but this may change as more is learnt of late
Miocene and Pliocene faunas elsewhere in Africa.

ACKNOWLEDGEMENTS

I am greatly indebted to Dr Elaine Anderson (Denver, Colorado) for
assistance in preparing the manuscript of this paper and for providing the data
on Gulo gulo used in Table |. I also thank Mr C. A. Repenning (U.S. Geological
Survey, Menlo Park) for providing Gulo gulo comparative material, Miss
T. Salinger for the photographs and Mrs P. Eedes for typing the tables.

The Langebaanweg Research Project is supported by Chemfos Ltd., the
South African Council for Scientific and Industrial Research and the Wenner-
Gren Foundation for Anthropological Research (Grant no. 2752-1834), and
the assistance of these organizations is gratefully acknowledged.

LATE TERTIARY MUSTELIDAE FROM LANGEBAANWEG, SOUTH AFRICA 357

REFERENCES

ANDERSON, E. 1977. Pleistocene Mustelidae (Mammalia, Carnivora) from Fairbanks, Alaska.
Bull. Mus. comp. Zool., Harv. 148: 1-21.

BisHop, W. W. & PICKFORD, M. 1975, Geology, fauna and palaeoenvironments of the Ngorora
Formation, Kenya Rift Valley. Nature, Lond. 254: 185-192.

Byork, P. R. 1970. The Carnivora of the Hagerman Local Fauna (Late Pliocene) of south-
western Idaho. Trans. Am. phil. Soc. 60: 1-54.

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

ELLERMAN, J., MORRISON-ScoTT, T. C. S. & HAYMAN, R. W. 1953. Southern African Mammals
1758 to 1951: a reclassification. London: British Museum (Natural History).

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

HENDEY, Q. B. 1974a. New fossil carnivores from the Swartkrans australopithecine site
(Mammalia: Carnivora). Ann. Transy. Mus. 29: 27-48.

Henpbey, Q. B. 19746. The late Cenozoic Carnivora of the south-western Cape Province.
Ann. S. Afr. Mus. 63: 1-369.

HENDEY, Q. B. 1976a. Fossil peccary from the Pliocene of South Africa. Science 192: 787-789.

HENDEY, Q. B. 19766. The Pliocene fossil occurrences in ‘E’ Quarry, Langebaanweg, South
Africa. Ann. S. Afr. Mus. 69: 215-247.

HENDEY, Q. B. 1977. Fossil bear from South Africa. S. Afr. J. Sci. 73: 112-116.

HENDEY, Q. B. 1978a. The age of the fossils from Baard’s Quarry, Langebaanweg, South
Africa. Ann. S. Afr. Mus. 75: 1-24.

HENDEY, Q. B. 19785. Preliminary report on the Miocene vertebrates from Arrisdrift, South

West Africa. Ann. S. Afr. Mus. 76: 1-41.

HENDEY, Q. B. 1978c. Late Tertiary Hyaenidae from Langebaanweg, South Africa, and their
relevance to the phylogeny of the family. Ann. S. Afr. Mus. 76: 265-297.

HoweELi, F. C. & PETTER, G. 1976. Carnivora from Omo Group formations, southern
Ethiopia. In: Coppens, Y. et al., eds. Earliest man and environments in the Lake Rudolf
Basin: 314-331. Chicago: University Press.

KurTEN, B. 1970. The Neogene wolverine Plesiogulo and the origin of Gulo (Carnivora,
Mammalia). Acta zool. fenn. 131: 1-22.

KurtTEN, B. 1971. The Age of Mammals. London: Weidenfeld & Nicolson.

KourtTEN, B. & RAuScH, R. 1959. Biometric comparisons between North American and
European mammals. Acta arctica 11: 1-44.

MATTHEW, W. D. 1907. A Lower Miocene fauna from South Dakota. Bull. Am. Mus. nat. Hist.
23: 169-219.

Novikov, G. A. 1962. Fauna of the U.S.S.R. 62. Carnivorous mammals. Jerusalem: IPST Press
(Israel Program for Scientific Translations Ltd).

PILGRIM, G. E. 1932. The fossil Carnivora of India. Mem. geol. Sury. India Palaeont. indica (n.s.)
18: 1-232.

REPENNING, C. A. 1976. Enhydra and Enhydriodon from the Pacific Coast of North America.
J. Res. U.S. geol. Surv. 4: 305-315.

STROMER, E. 1931. Reste stisswasser- und land bewohnender Wirbeltiere aus den Diamanten-
feldern Klein- Namaqualandes (Siidwest-afrika). Sber. bayer Akad. Wiss. 1931: 17-47.
VirET, J. 1939. Monographie paléontologique de la faune de vertébrés des sables de Montpellier.

III. Carnivora Fissipedia. Trav. Lab. géol. Fac. Sc. Lyon 37: \-26.

Wess, S. D. 1969. The Burge and Minnechaduza Clarendonian mammalian faunas of north-

central Nebraska. Univ. Cal. Publ. geol. Sci. 78: 1-484. |~| |.

q
»
*

6. SYSTEMATIC papers must conform to 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. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
, Figs 14-1SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *.. . the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

(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 but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

Q. B. HENBEY

LATE TERTIARY MUSTELIDAE
(MAMMALIA, CARNIVORA)
FROM LANGEBAANWEG, SOUTH AFRICA

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