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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 104 Band

February 1995 Februarie

Rane 27 ae Deel

DEVONIAN CHONETACEAN BRACHIOPODS

FROM SOUTH AFRICA

NORTON HILLER

Cape Town Kaapstad

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D3586

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA

NORTON HILLER

Department of Geology, Rhodes University, Grahamstown, South Africa, 6140

(With 10 figures)

[MS accepted 3 October 1994]

ABSTRACT

Six species of chonetacean brachiopods are recorded from the Lower-Middle Devonian

Bokkeveld Group of South Africa, with the bulk of the material coming from the Emsian-

Eifelian Ceres Subgroup. Kentronetes africanus and Aseptonetes capensis are described as

new species, and Anoplia is recorded from South Africa for the first time, extending its range

within the Malvinokaffric Realm. Most specimens, except some of the larger Notiochonetes

and Pleurochonetes, come from fine-grained deposits, suggesting a preference for the tranquil

subtidal conditions of benthic assemblages 4 and 5. Although much less diverse, the fauna

shows strong palaeobiogeographic affinities with those from South America and the Falkland

Islands.

CONTENTS

PAGE

MRIERQOUCHOM cre ete cance tice sie cionsticin oem ace sew on cisions sis sacise ei eraass cise ear aacle naan neu ene 159

EFAR DEAD NIC CISETIDUEIONY se cceee sees see stent cis core cto ctersieeoeie sister tettietenre sta STIG sees eat 160

Eaunallistor Devonianchonetids from/SouthvAtricay 2.2. .sseeseecssccee- teen oes ecaneeseciee 162

AIACOCCOLO LY oat cre ioe. sotinets ses sssicwiaie secic edie cleus clase WOE nie na nloecslasentcoae Mets eaeuc acca samementanG 163

EAlACODIOL COP TAD IC AlMItleS ea ceee eceine esac cee crete ce sa en Clee Ne oe sere ene 164

MPUSICIALICAPAlACON(OlOPY) a asescscmscconcoecoaceec Sune eesemae neciets vi cas nelesaterets anes mune taraunere ae 165

MOHENISIOH Ste. tees ces ee romans aoce see eaees cee eee ae ORR Ne A ee ea ese Ree Seen 178

PAC MMOWICOD EIMEHES ree pea rc sees nctoncla nein cte ois Sie oes atta enistnen aide Siameald aut eloactuiaiatanmare ane 178

| FSS (Caen arse CBC eC eS ICTS OT ISIC NPG ACCS Tt sa RDI at es aro nie LN ta 179

INTRODUCTION

Brachiopods usually form an important part of the marine fauna found in

fossiliferous Palaeozoic rocks and, in many cases, they were the dominant

elements in shallow-water benthic communities of the time. In Devonian rocks,

chonetaceans often form a significant proportion of the brachiopods recovered

and this is certainly true for the Bokkeveld Group of South Africa.

The fossiliferous nature of the Bokkeveld Group sediments was recognized

in the early part of the nineteenth century, but few attempts at systematic des-

criptions of the fossil remains were made until Andrew Geddes Bain submitted a

collection of specimens to the Geological Society of London in 1852. The

brachiopods, described by Sharpe (1856), included nine species, two of which

were assigned to Chonetes.

* Present address: Department of Geology, University of Canterbury, Private Bag 4800,

Christchurch, New Zealand

159

Ann. S. Afr. Mus. 104 (7), 1995: 159-180, 10 figs.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

During subsequent years, extensive mapping by the Geological Commission

of the Cape of Good Hope led to the discovery of many more fossils and a rep-

resentative collection was sent to England for identification. Reed (1903)

described the brachiopods and increased the number of species present to 23.

The work of Ulrich (1893) in Bolivia drew attention to the close similarity

between that country’s Devonian fauna and others known from South America,

the Falkland Islands and South Africa. As more work was carried out on the

South American faunas by Thomas (1905), Knod (1908), Clarke (1913) and

Kozlowski (1913, 1923), and with further collecting in South Africa, so Reed

continued to revise and add to the Bokkeveld brachiopod fauna (1906, 1907).

Reed published his final revision of the Bokkeveld fauna in 1925, in which

the number of brachiopod species recorded was increased to 37, including six

chonetaceans. Since then little formal taxonomic work has been done on the

brachiopods apart from Boucot ef al. (1963), Boucot et al. (1965) and Hiller

(1987). Mention has been made of particular South African forms in Cloud

(1942) and Boucot & Harper (1968) but without detailed descriptions. Other

references to the Bokkeveld brachiopods have come in papers dealing with

palaeobiogeography and/or palaeoecology, e.g. Boucot ef al. (1969), Boucot

(1970, 1971, 1975a), Copper (1977) and Hiller & Theron (1988), but sys-

tematic descriptions were not included in any of these.

Reed (1925: 28) noted that many of the South African species were

established on poorly preserved material and that it would have been better had

specific names not been applied. He made the same comment about the South

American faunas and Branisa (1965) proposed nomima nuda for some taxa with-

out proper descriptions. A revision of the Bokkeveld brachiopods is therefore

warranted. In light of more recent work in South America, e.g. Isaacson (1977),

Racheboeuf & Branisa (1985) and Racheboeuf (1992), this paper makes a start

with the chonetacean representatives in the fauna. This follows from an earlier

paper (Hiller 1987), in which an attempt was made to clear up the confusion

regarding the identity of those specimens that had been referred to Chonetes

falklandicus Morris & Sharpe and Notiochonetes.

Specimens described are housed either in the South African Museum, Cape

Town (SAM numbers) or in the Albany Museum, Grahamstown (AM numbers).

STRATIGRAPHIC DISTRIBUTION

The currently recognized lithostratigraphical subdivision of the Bokkeveld

Group (Fig. 1) was established by Theron (1970, 1972) and revised by the

South African Committee for Stratigraphy (1980). The Bokkeveld Group is the

middle division of the tripartite Cape Supergroup and appears to have been

deposited during a period of relative crustal instability. Lithologically the group

consists of a number of predominantly argillaceous units alternating with pre-

dominantly arenaceous units to which Theron (1970) assigned formational

status. The sediments are arranged into a number of coarsening-upward

sequences that imply phases of tectonically controlled regressions and trans-

gressions coupled with delta out-building and reworking (Tankard et al. 1982:

352). Southwards the arenaceous units die out and the most southerly outcrops

appear to represent a deeper-water shelf facies.

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 161

FORMATION

FAMENNIAN

: S$WARTRUGGENS - PRASNIeN

WAGEN = DRIFT 7 «

KAROOPOORT

[=2 OSBERG =..).: GIVETIAN

KLIPBOKKOP

WUPPERTAL ~

———F

WABOOMBERG 4

WITTEBERG

SUBGROUP

BIDOUW

__BOPLAAS . -

= TRA-TRA 4

EIFELIAN

SS ag

SUBGROUP

BOKKEVELD

VOORSTEHOEK <4

56a)

(LIT) me

G) Roel oer oreo Sa KAn As PNAu

S |-’-’- RIETVLEI -::.

fae | eieikes eteeicaiielehac neueine

CCC DO UeCHnCIMECh ON Ln iC)

Statiethalion iene eh eee

Fig. 1. Stratigraphic column of the Devonian rocks in South Africa showing

the ranges of species described in this paper. Black triangles indicate the

formations from which chonetid brachiopods have been recorded. K—Kentro-

netes; An—Anoplia; As—Aseptonetes; P—Pleurochonetes; N—Notiochonetes;

Au—Australostrophia. Stipple shading indicates sandstone-sdominated

horizons; horizontal lines indicate finer-grained units.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

That the Bokkeveld Group is of Devonian age was established by the early

workers, e.g. Reed (1903), but a more precise dating proved to be something of

a problem. The ages of the various stratigraphic units indicated in Figure | are

based on arguments put forward by Boucot et al. (1969), Theron (1970, 1972)

and Boucot (1975a) on the basis of the marine invertebrate fossils of the Ceres

Subgroup. More recently, Cooper (1986) used eustatic changes in sea-level to

effect a correlation with European and North American sequences and suggested

that the marine portion of the Bokkeveld Group was late Emsian-Eifelian in

age.

Marine invertebrate fossils are characteristic of the lower part of the Bokke-

veld Group, up to and including the fourth shale horizon, i.e. the whole of the

Ceres Subgroup, plus the Waboomberg Formation (Fig. 1). However, by far the

greatest number of specimens has been recovered from the Gydo and Gamka

formations at the base of the group (Hiller & Theron 1988). Chonetacean

brachiopods are here recorded from the lowest three formations of the

Bokkeveld Group plus the Tra-Tra and Waboomberg formations. Possible

chonetaceans have also been noted from the Rietvlei Formation at the top of the

Table Mountain Group (Hiller & Theron 1988) and from the Wagen Drift

Formation at the base of the Witteberg Group (Hiller & Dunlevey 1978).

FAUNAL LIST OF DEVONIAN CHONETIDS FROM SOUTH AFRICA

The South African Devonian brachiopod fauna is, overall, a very conserva-

tive one, showing much lower diversity in terms of genera and species than

contemporary faunas from Europe or North America. It is also apparently less

diverse than equivalent South American faunas. Among the chonetaceans only

six genera, each represented by a single species, are currently recognized. The

equivalent fauna in Bolivia contains 20 species belonging to 11 genera of

chonetaceans (Isaacson 1977; Racheboeuf & Branisa 1985; Racheboeuf 1992).

The South African Devonian chonetaceans are represented by the following

taxa:

Family Strophochonetidae

Kentronetes africanus sp. nov.

Australostrophia clarkei Racheboeuf & Herrera, 1994

Family Anopliidae

Subfamily Anopliinae

Anoplia cf. A. belenensis Racheboeuf & Branisa, 1985

Family Chonetidae

Subfamily Chonetinae

Aseptonetes capensis sp. nov.

Subfamily Notiochonetinae

Notiochonetes skottsbergi (Clarke, 1913)

Pleurochonetes falklandicus (Morris & Sharpe, 1846)

Apart from these, other Malvinokaffric taxa may be present in South Africa.

Reed (1925) described and figured the internal mould of a pedicle valve that he

identified as Chonetes (Eodevonaria) aff. arcuata Hall. However, he stated that

the specimen did not show any spines or crenulations (denticles) along the hinge

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA - 163

line, a feature diagnostic of Eodevonaria. Muir-Wood (1962: 93) included

Reed’s record in the list of species to be included in Eodevonaria, but Boucot &

Harper (1968: 151) categorically rejected the specimen from the genus and

stated that Eodevonaria had not been recognized in the Malvinokaffric Realm.

Subsequently, Isaacson (1977) discovered a species of Eodevonaria in Peru and

again raised the possibility (p. 174) that Reed’s specimen may belong in the

genus. Herrera (1991) has recorded Eodevonaria from Argentina and it has now

been found in Bolivia (P. R. Racheboeuf pers. comm. 1994).

Reed (1925: 46) mentioned that a specimen from the southern Cape in the

Sedgwick Museum, Cambridge, did show denticles along the hinge line. The

author has examined two pedicle valve internal moulds labelled as

C. (Eodevonaria) aff. arcuata in the Sedgwick Museum, although they were

originally identified as C. falklandicus. One of them possessed some denticles

along one-half of the hinge line but assignment to Eodevonaria would only be

tentative at best. Therefore, unless better-preserved material is found, the

presence of the genus in South Africa must be regarded as still unsubstantiated.

Other Bolivian forms, such as Austronoplia and Babinia, have not been

positively identified from South African rocks but the possibility exists that,

with further collecting, they may well turn up in the Bokkeveld fauna. Several

small specimens that the author originally identified as Babinia lacked an

enlarged median costa and hence are regarded as juveniles of Kentronetes, but

Reed (1925) described Chonetes ruecki medialis as a new subspecies of

C. ruecki Ulrich differing from it principally in possessing an enlarged median

rib. Reed’s specimen is shown here as Figure 3H. Racheboeuf & Branisa (1985)

placed C. ruecki in Quadrikentron and erected Babinia for small specimens

possessing an enlarged median costa and lacking a dorsal median septum.

Reed’s specimen (1925, pl. 4 (figs 7-8)) may well belong in Babinia, but it is

not sufficiently well preserved for such an assignment to be certain.

Reed (1925: 44) transferred several small shells that he had originally (Reed

1903) identified as Chonetes aff. C. setiger Hall to Chonetes stuebeli Ulrich,

which is now the type species for the genus Austronoplia Isaacson, 1977.

However, Reed did not describe the interior of the brachial valve and, from his

illustrations, the pedicle valve could just as well belong to Aseptonetes.

PALAEOECOLOGY

The Bokkeveld Group chonetacean brachiopods show a preference for

environments in which finer-grained clastic sediments were deposited. As can

be seen in Figure 1, they tend to occur in those formations that are predomi-

nantly shale and mudstone, although they are occasionally found in sandier

horizons within these formations. Only Pleurochonetes and Notiochonetes have

been recovered from the sandstone-dominated Gamka Formation.

Hiller & Theron (1988) outlined a number of benthic communities that can

be recognized within the Bokkeveld Group, which has been interpreted as a

series of stacked deltaic cycles (Tankard & Barwis 1982). Chonetacean

brachiopods are found in each of the depositional settings associated with the

deltas, but particularly in the distributary mouth bar, delta slope, and prodelta

shelf communities. Notiochonetes appears to have preferred the shallow-water

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

distributary mouth bar and delta-slope environments, probably the equivalent of

benthic assemblages 3 and 4 of Boucot (1975a). Evidently the large flattish

Australostrophia and Kentronetes, and the small Aseptonetes and Anoplia

preferred the deeper, less-turbulent settings of the lower delta slope and shelf as

they are only known from muddy horizons, i.e. benthic assemblages 4 and 5.

Pleurochonetes seems to have tolerated a wider range of conditions as it has

been recovered from distributary mouth bar, delta slope and shelf sediments,

i.e. benthic assemblages 3-5. No chonetaceans are known for certain from the

tidal-flat community but, if their occurrence in the Wagen Drift Formation of

the Witteberg Group can be confirmed, then they may be found in such a

setting. Similarly, the possible occurrence of chonetaceans in the Rietvlei

Formation at the top of the Table Mountain Group suggests their presence in a

lower shoreface community (Hiller & Theron 1988: 238).

When any of the species associated with muddy environments is found in

sandy horizons within the shaly units, they usually occur as dissociated valves

resting in a convex-up position on a bedding surface. This may indicate some

transport before relatively slow burial. In contrast, when they are found in the

mudstones they often occur as complete shells, suggesting fairly rapid burial.

Their orientation within the sediment is usually random, however, probably as a

result of the intense bioturbation normally observed in these units (Toots 1965).

The larger, flatter forms probably lived free on the sea-floor, using their

spines for stabilization in the manner suggested by Brunton (1972) and Rache-

boeuf (1990), but the discovery of a complete specimen of Aseptonetes capensis

sp. nov. in close association with a plant fragment raises the question as to

whether the shell was attached to the plant during life. Plant fragments are not

uncommon in these shallow clastic environments and the juxtaposition of the

specimens may simply be a chance occurrence, but the possibility remains that

at least some small chonetaceans may have made use of suitable substrates for

attachment if such were available. Racheboeuf (1990: 165) has suggested that

small chonetaceans, especially those with long asymmetrical spines, probably

lived epiplanktonically, attached to algae or floating organisms.

PALAEOBIOGEOGRAPHIC AFFINITIES

The Devonian fossils of the Bokkeveld Group have long been regarded as

belonging to, and typical of, the Malvinokaffric palaeobiogeographic realm

(Boucot et al. 1969; Boucot 1988). It is not surprising, therefore, that the

chonetacean brachiopods within the Bokkeveld fauna show greatest similarity

with those in other parts of this realm, including the Falkland Islands, Bolivia,

and the Paranda Basin of Brazil. Of the forms described here, Australostrophia

and Notiochonetes are endemic to the Malvinokaffric region, having been

recorded from Bolivia, Brazil, the Falklands, South Africa, and possibly Argen- —

tina and Uruguay (Boucot et al. 1969: 27; Melo 1988: 695). Pleurochonetes and

Aseptonetes are similarly likely to be Malvinokaffric endemics. The former

shows the same distribution as Australostrophia and Notiochonetes, whereas

Aseptonetes is so far known only from Bolivia and South Africa.

The genus Kentronetes has recently been erected for Malvinokaffric realm

strophochonetids (Racheboeuf & Herrera 1994) and is thus also endemic to the

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 165

realm. Racheboeuf & Branisa (1985) recorded it from Bolivia as Quadrikentron.

Its occurrence in South Africa extends the geographic range of the genus within

the Malvinokaffric Realm. Anoplia is a genus with an almost cosmopolitan

distribution during Early Devonian times. It occurs in the Eastern American

Realm, the Rhenish-Bohemian and Tasman regions of the Old World Realm,

and the Malvinokaffric Realm (Boucot et al. 1969). The South African

occurrence extends its range further.

“woes mmm

0 200 km

Fig. 2. Map showing the outcrop of the Bokkeveld Group rocks in the Cape Province and

indicating the positions of towns mentioned in the text. A—Alexandria, C—Ceres,

CT—Cape Town, Cw—Clanwilliam, G—Grahamstown, M—Montagu, P—Prince Albert,

PA—Port Alfred, PE—Port Elizabeth, S—Steytlerville, TR—Touws River, U—Uitenhage,

Ud—Uniondale, W—Willowmore, Wo—Worcester, Wu—Wuppertal.

SYSTEMATIC PALAEONTOLOGY

Suborder CHONETOIDEA Muir-Wood, 1955

Superfamily CHONETACEA Bronn, 1862

Family Strophochonetidae Muir-Wood, 1962

Genus Kentronetes Racheboeuf & Herrera, 1994

Kentronetes africanus sp. nov.

Figs 3-4

Chonetes falklandicus non Morris & Sharpe, 1846. Reed, 1903: 171-172, pl. 20 (figs 9-10).

Material and horizon

Holotype. AM 5187 in the Albany Museum, Grahamstown. Internal mould

of a brachial valve from the Voorstehoek Formation, Alexandria district,

eastern Cape (see Fig. 2).

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes. AM 5175-1, AM 5188-AM 5191, SAM-13400: internal and

external moulds of 14 pedicle valves and 4 brachial valves from the Gydo

Formation in the Wuppertal and Ceres districts of the western Cape, and

21 pedicle and 3 brachial valves from the Voorstehoek Formation in the

Alexandria district of the eastern Cape.

Diagnosis

A species of Kentronetes with markedly transverse elliptical outline

(length/width ratio about 0.60), three spines on either side of umbo in

adult stages, and lacking any sort of median septum or ridge in the brachial

valve.

Description

Small to medium sized (maximum length 17.6 mm) plano- to gently

concavo-convex Shells with transversely sub-rectangular to sub-elliptical

outlines; maximum width at a little over two-fifths of valve length from umbo;

cardinal extremities rounded or obtuse. Pedicle valve about three-fifths as long

as wide; very gently convex in both lateral and anterior profiles. Umbo small;

interarea short, flat, apsacline; delthyrium with small apical pseudodeltidium.

Brachial valve interarea very short, catacline; no chilidium observed. Radial

ornament of fine, low, rounded capillae that increase by branching and

intercalation to number 100-120 at anterior margin, with a density of 16-19

(most commonly 17) ribs in a 5-mm sector at the 10-mm growth stage. Three

small specimens show development of enlarged median costa with rounded

cross-section and extending to valve margin from about one-third valve length.

Two, or maximum three, spines on either side of umbo, at high angles

(66°-75°) to hinge line. Spines about 2 mm apart, nearest 5 mm from umbo.

Pedicle valve interior with small, obliquely elongate hinge teeth directed

anterolaterally, without dental plates. Muscle field broadly triangular, two-thirds

as long as wide and almost two-fifths as long as valve; anterior margin weakly

impressed, posterior bounding ridges low, gently curved and diverging at about

100°. Median septum low, narrow, up to one-quarter as long as valve. Large

triangular diductor scars enclose small, elongate, oval adductor scars.

Brachial valve interior without median septum. Bilobed cardinal process

fused to proximal ends of long, widely divergent, curved inner socket ridges.

Sockets moderately deep, curved. Anderidia long, very narrow, diverging at

about 55°-60°. Muscle field about four-fifths as long as wide and about

one-third as long as valve, bisected by shallow median groove extending

anteriorly from in front of cardinal process.

Dimensions (in mm)

Pedicle valves length width

AM 5175 335 248

AM 5189 13:9 28.7

AM 5190 14.4 30:5

SAM-13400 12a 24.9

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 167

Fig. 3. A-G. Kentronetes africanus sp. nov. A. AM5187, holotype, internal mould of a

brachial valve, x 2.9. B. AM5175-1, internal mould of conjoined pedicle and brachial

valves, x 4.1. C-D. AM5188, internal mould and latex cast of a brachial valve, x 2.1

and 2.2, respectively. E-F. AM5190, internal mould and latex cast of a pedicle

valve, x 2.1. G. AM5191, latex cast of a brachial valve exterior, x 2.1. H. Babinia?

medialis. SAM-7779, external mould of a brachial valve, x 4.2 (Reed’s holotype of

Chonetes ruecki medialis). A and C-G are from the Voorstehoek Formation in the

Alexandria district, eastern Cape; B is from the Gydo Formation, near Wuppertal, western

Cape; H is probably from the Gydo Formation in the Hex River Valley.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

The specimens described here are most similar to Quadrikentron ruecki

(Ulrich, 1893), described from Bolivia by Racheboeuf & Branisa (1985). The

main difference with the South American species is that the South African shells

are slightly larger and possess a significantly more transverse outline. They

also lack a dorsal median septum, even the wide low ridge seen in the Bolivian

specimens. In most other respects they are very similar. Quadrikentron

ruecki has recently been transferred to Kentronetes by Racheboeuf & Herrera

(1994).

Fig. 4. Kentronetes africanus sp. nov. A. SAM-13400, internal mould of a pedicle

valve, x 2.6. B. AM5189, internal mould of a pedicle valve, x 1.9. A is probably

from the Gydo Formation in the Ceres district, western Cape; B is from the

Voorstehoek Formation in the Alexandria district, eastern Cape.

Genus Australostrophia Caster, 1939

Remark

The decision to place Australostrophia in the family Strophochonetidae

follows Racheboeuf & Herrera (1994).

Australostrophia clarkei Rachebeouf & Herrera, 1994

Fig. 5

Stropheodonta cf. S. concinna (Morris & Sharpe). Reed, 1903: 169, pl. 22 (fig. 6).

Stropheodonta (Leptostrophia) concinna (Morris & Sharpe). Reed, 1925: 41.

Australostrophia clarkei Rachebeouf & Herrera, 1994: 556, fig. Sa-f.

Material and horizon

Internal and external moulds of 9 pedicle valves and 2 brachial valves from

the Gydo Formation, north of Clanwilliam in the western Cape Province

(Big. 2):

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 169

Fig. 5. Australostrophia clarkei Racheboeuf & Herrera, 1994. A-B. AM 5186-2,

internal mould and latex cast of a brachial valve, x 3.1 and 2.3, respectively.

C. AM5186-5, latex cast of a pedicle valve exterior, x 1.7. D. AM5186-1, internal

mould of a pedicle valve, x 2.1. E-F. AM5185-1, latex cast and internal mould of a

pedicle valve, x 2.4. G. AM 5184, external mould of a pedicle valve, x 2.5.

H. AM5185-2, internal mould of a pedicle valve, x 2.4. I. AM5186-3, internal

mould of a pedicle valve, x 3. All from the Gydo Formation north of Clanwilliam,

western Cape.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Medium to large plano-convex shells with semi-circular outline, maximum

width along hinge line; cardinal extremities slightly acute to slightly obtuse.

Two pairs of spines situated on either side of umbo. Pedicle valve about

two-thirds as long as wide; gently convex in both lateral and anterior profiles

with greatest convexity in umbonal region, becoming flatter towards the

posterolateral margins. Ventral interarea short, apsacline. Delthyrium closed

by large convex pseudodeltidium. Brachial valve flat. Dorsal interarea very

short; anacline. Surface ornament of fine rounded costellae, numbering about

22 in a 5-mm sector at 10-mm growth stage and increasing by branching

and intercalation, crossed by fine concentric filae giving ribs a beaded

appearance.

Pedicle valve interior with stout hinge teeth diverging at 90°-100° and

supported by reduced dental lamellae. Large, roundly triangular, flabellate

muscle field about two-thirds as long as valve, bounded posteriorly by low

ridges and bisected by low, slender median myophragm.

Brachial valve interior with prominent bilobed cardinal process situated on

low notothyrial platform. Strong inner and outer socket ridges bound large oval

sockets. Short anderidia and low median ridge extend anteriorly from

notothyrial platform to bound anterior adductor muscle scars.

Dimensions (in mm)

Pedicle valves length width

AM 5184 18.3 25.4

AM 5185 10.6 16.1

9.2 13.4

AM 5186 3h<7/ 21.6

7.4 10.9

14.4 19.8

19.1 30.9

Ted 11.4

Discussion

The specimens described herein are referable to Australostrophia because of

their distinctive ornament and chonostrophiid-like cardinalia. Only three species

are currently assigned to this genus, the type species Australostrophia

mesembria (Clarke, 1913), the much smaller A. senegalensis Racheboeuf &

Villeneuve, 1989, and A. clarkei Racheboeuf & Herrera, 1994. Of these the last

most closely resembles the specimens described here. Indeed, they are virtually

identical to specimens described from Bolivia by Isaacson (1977) and -

Racheboeuf & Branisa (1985) as A. mesembria but subsequently treated as a

separate species by Racheboeuf & Herrera (1994).

The pedicle valve described and figured by Reed (1903) as Stropheodonta

cf. S. concinna cannot be assigned to that species because it lacks the

denticulate hinge line of Protoleptostrophia to which that species belongs. The

specimen is better placed in A. clarkei, as it bears close resemblance to the

specimens described here.

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 171

Family Anopliidae Muir-Wood, 1962

Genus Anoplia Hall & Clarke, 1892

Anoplia cf. A. belenensis Racheboeuf & Branisa, 1985

Fig. 6

Compare:

Anoplia belenensis Racheboeuf & Branisa, 1985: 1437, figs 7.1-7.7, table 5.

Material and horizon

Twelve pedicle and 9 brachial valves from the Voorstehoek Formation near

Uitenhage, eastern Cape Province (Fig. 2).

Description

Small concavo-convex shells with transverse semicircular outline, maximum

width at hinge line; cardinal extremities rounded. Pedicle valve about three-

quarters as long as wide; moderately convex in lateral profile, height about one-

third of valve length. Umbo small; interarea short, flat, apsacline with small

convex pseudodeltidium covering apical portion of delthyrium. Brachial valve

gently concave with very short, flat, hypercline interarea. Shell exterior smooth.

Pedicle valve interior with small hinge teeth. Myophragm short, about one-

third as long as valve. Muscle field well impressed, about as long as wide,

bounded posteriorly by strong ridges diverging at about 80°. Diductor scars

Fig. 6. Anoplia cf. A. belenensis Racheboeuf & Branisa, 1985. A. AM5181, internal

mould of a pedicle valve, x 3.8. B. AM5180, external mould of a brachial

valve, x 4. C. AM5183, internal mould of a pedicle valve, x 4. D. AM5182-2,

internal mould of a pedicle valve, x 3.8. E. AM5182-1, internal mould of a pedicle

valve, x 4. F. AM5182-3, external mould of a brachial valve, x 4.3. All from the

Voorstehoek Formation near Uitenhage, eastern Cape.

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

elongate, triangular in outline with rounded anterior margins; diverging

anteriorly. Adductor scars small, narrow. Roots of up to four spines observed

along posterior margin on either side of beak.

Brachial valve interior poorly preserved. Cardinal process very small,

bilobed. No median septum. One pair of accessory septa about one-half as long

as valve, diverging anteriorly at 22°.

Dimensions (in mm)

Pedicle valves length width

AM 5181 529 8.5

AM 5182 6.0 8.0

CEO @

AM 5183 4.5 6.6

Discussion

The lack of radial ornament and absence of a dorsal median septum, but

with a pair of diverging accessory septa, suggest placement in Anoplia. In size

and shape, the specimens described here are very similar to those described

from Bolivia by Racheboeuf & Branisa (1985), and they must be regarded as

being closely related. However, more and better-preserved material is required

before unequivocal assignment to the species is warranted. The specimens can

be separated from the other Bolivian species, A. mariae Racheboeuf, 1992,

which has two pairs of accessory septa in the dorsal valve and only two spines

along the posterior margin.

These specimens represent the first record of the genus from South Africa.

Family Chonetidae Bronn, 1862

Subfamily Chonetinae Bronn, 1862

Genus Asepronetes Isaacson, 1977

Aseptonetes capensis sp. nov.

Fig. 7

Chonetes aff. setiger Hall: Reed, 1903: 174, pl. 21 (figs 4-5).

Chonetes aff. setiger Hall: Reed, 1907: 170, 223.

Chonetes sttibeli non Ulrich. Reed, 1925: 44.

Material

Holotype. AM 5174 in the Albany Museum, Grahamstown. Internal and —

external moulds of joined pedicle and brachial valves from the Gydo Formation

at Wuppertal (Fig. 2).

Paratypes. AM 5175-AM 5179, 19 brachial and 20 pedicle valves from the

Gydo Formation at Wuppertal in the western Cape and three pedicle valves

from the same formation in the Cockscomb area, south-east of Steytlerville in

the eastern Cape. Further material is from the Voorstehoek Formation near

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 173

Uitenhage, eastern Cape (3 brachial and 14 pedicle valves) and from two

localities within the Waboomberg Formation in the western Cape, including a

pedicle valve from Tafelberg in the Ceres district, plus a pedicle and a brachial

valve from the Hex River Valley, between Worcester and Touws River, both in

the western Cape (Fig. 2).

Fig. 7. Aseptonetes capensis sp. nov. A. AM5174-1a, holotype, internal mould of a

brachial valve, x 3.8. B. AM5174-2, internal mould of a pedicle valve and a brachial

valve, x 3.7. C. AM5174-2, latex cast of brachial valve interior, x 3.8.

D. AM5175-2, external mould of a pedicle valve, x 5.3. E. AM5178, internal mould

of a brachial valve, x 5. F. AM5174-3, internal mould of conjoined pedicle and

brachial valves, x 4.8. G. AM5179, internal mould of a pedicle valve, x 3.

H. AM5175-3, external mould of a brachial valve, x 5.1. I. AM5177, latex cast of a

pedicle valve interior, x 4.3. J. AM5176, internal mould of conjoined pedicle and

brachial valves, x 4.5. A-D, F and H-J come from the Gydo Formation at

Wuppertal, western Cape; E and G come from the Voorstehoek Formation near

Uitenhage, eastern Cape.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Small chonetids with fairly coarse V-shaped ribs and lacking dorsal median

septum. Anderidia and ventral median myophragm better developed than in the

closely related A. boucoti Isaacson.

Description

Small to medium concavo-convex shells with transversely subrectangular

outline; maximum width along hinge line. Pedicle valve between two-thirds and

three-quarters as long as wide; strongly and evenly convex in lateral profile but

anterior profile with strongly convex median portion and more gently convex

flanks. Ventral interarea curved orthocline to apsacline; delthyrium partially

closed apically by convex triangular pseudodeltidium. Dorsal interarea flat,

hypercline; chilidium strongly convex. Radial ornament of fairly coarse

V-shaped costae and costellae increasing by bifurcation and intercalation to

number 32-48 in shells between 5 mm and 11 mm long, with a density of

2-4 per millimetre at anterior margin. Four or five spines, seen as oblique

openings on valve interiors, more or less symmetrically arranged on each side

of umbo with the inner three being more closely spaced than the outer two.

Pedicle valve interior with stout, widely divergent, elongate hinge teeth

supported by reduced dental lamellae. Muscle field not clearly defined but

broadly triangular, flabellate, less than half as long as valve. Short median

myophragm divides muscle field and extends for about one-third of valve length.

Brachial valve interior with well-developed, high, distally bilobed cardinal

process and well-defined alveolus. Deep oval sockets bounded by strong, widely

divergent inner socket ridges and less well-developed outer ones. Anderidia

short, slender, diverging at an angle of about 35° and extending for about one-

quarter of valve length. Median septum absent. Accessory septa originate

anterior to alveolus, diverge at about 20° and extend for four-fifths of valve

length.

Dimensions (in mm)

Pedicle valves length width

AM 5174 ay 7.8

4.2 62

AM 5175 5.0 6.6

AM 5176 4.6 a9

AM 5177 6.5 9.4

AM 5179 US? 11.0

Discussion

The small size, relatively coarse V-shaped radial ornament, and lack of -

median septum in the brachial valve suggest that these specimens belong in

Aseptonetes. However, Racheboeuf & Branisa (1985) suggested that the speci-

mens originally described by Isaacson (1977) were immature and did not show

the adult characteristics. They described and figured somewhat larger shells that

show a well-developed median septum but lack accessory septa. They stated that

these specimens occurred on the same slabs as specimens bearing a close resem-

blance to Isaacson’s type specimens and therefore represent later growth stages.

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 175

Fig. 8. Notiochonetes skottsbergi (Clarke, 1913). A-B. AM 4805, internal mould and

latex cast of a brachial valve, x 1.5. C. AM5192, external mould of a brachial

valve, x 2.0. D. SAM-6713, latex cast of a brachial valve exterior, x 1.3.

E. SAM-3391, internal mould of a pedicle valve, x 1.9. F. AM 4808, latex cast of a

brachial valve exterior, x 1.4. A, B, and F are from the Tra-Tra Formation, and C is

from the Waboomberg Formation, all from north-east of Montagu, western Cape;

D-E are from the Gydo Formation in the Ceres district, western Cape.

The Bokkeveld Group specimens described here overlap in size range with

both Isaacson’s (1977) and Racheboeuf & Branisa’s (1985) specimens, but none

are as large as their biggest shells. All bear a close resemblance to A. boucoti as

described by Isaacson but none are similar to those described by Racheboeuf &

Branisa. They also occur on the same pieces of rock as larger chonetids that do

show the development of a median septum. However, there is no overlap in

sizes, the larger shells have rounded rather than V-shaped ribs, and there are

certainly no intermediate morphologies. The larger specimens are referable to

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pleurochonetes falklandicus Morris & Sharpe but the smaller ones cannot be

regarded as juveniles of this species.

To this author, Aseptonetes is a valid genus as defined by Isaacson (1977),

and the larger specimens described and figured by Racheboeuf & Branisa

probably should not be assigned to it. This being the case, then the subfamilial

placing of the genus should revert to the Chonetinae as originally assigned by

Isaacson.

As previously mentioned, the South African specimens are very similar to

Isaacson’s A. boucoti but they are rather more coarsely ribbed and their

anderidia and ventral median myophragms are longer. They are thus placed in a

separate species.

Subfamily Notiochonetinae Racheboeuf, 1992

Genus Notiochonetes Muir-Wood, 1962

Notiochonetes skottsbergi (Clarke, 1913)

Figs 8-9

For synonymy see Hiller (1987: 1152).

Dimensions (in mm)

Pedicle valves length width

SAM-3391 722 2920

AM 4806-2 Pile 34.4

AM 4806-3 25.4 38.5

Fig. 9. Notiochonetes skottsbergi (Clarke, 1913). A. AM4802-2, internal mould of a

pedicle valve, x 1.7. B. AM 4806-3, internal mould of a pedicle valve, x 1.8. Both

are from the Tra-Tra Formation north of Montagu, western Cape.

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 177

Fig. 10. Pleurochonetes falklandicus (Morris & Sharpe, 1846). A-B. AM 4811, internal

mould and latex cast of a brachial valve, x 2. C. SAM-13399, internal mould of a

pedicle valve, x 2.9. D. AM4812, internal mould of a pedicle valve, x 2.2.

E-F. SAM-13399, internal mould and latex cast of a brachial valve, x 3.

G-H. AM 4802, internal mould and latex cast of a brachial valve, x 1.8. I. AM 4801,

latex cast of a brachial valve interior, x 2. A, Band D are from the Gamka Formation in

the Hex River Valley area of the western Cape; C-F are from the western Cape but

locality and horizon are uncertain; G, H and I are from the Uitenhage district, eastern

Cape (horizon is uncertain but probably the Gydo Formation).

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This species was fully described by Hiller (1987) and nothing more is added

here. It has been recorded from the Gydo and Voorstehoek formations in the

Ceres district, the Voorstehoek Formation near Uitenhage, and the Tra-Tra

Formation north-east of Montagu in the western Cape (Fig. 2).

Genus Pleurochonetes Isaacson, 1977

Pleurochonetes falklandicus (Morris & Sharpe, 1846)

Fig. 10

For synonymy see Hiller (1987: 1149).

Dimensions (in mm)

Pedicle valves length width

AM 4812 13.0 20.0

SAM-13399 13 18.0

Remarks

This species was fully described by Hiller (1987) and nothing further is

added here. It is well represented in widespread exposures of the Gydo, Gamka,

and Voorstehoek formations, particularly in the western and southern Cape.

CONCLUSIONS

This study expands the known diversity of chonetacean brachiopods in the

South African Devonian and draws attention to the close similarity between the

Bokkeveld Group fauna and the Malvinokaffric Realm faunas of Bolivia, Brazil,

Argentina, Paraguay, Uruguay and Peru. Unfortunately, knowledge of

chonetaceans in South American countries other than Bolivia and Argentina is

incomplete, because of a lack of recent publications. Therefore, the significance

of the apparent absence of genera such as Austronoplia, Babinia and

Eodevonaria cannot be properly assessed at this stage. Certainly more work is

required on the South African fauna, especially from those stratigraphic units

that are under-represented in present collections. Only then will we be in a

position to better understand Lower—Middle Devonian biogeography.

ACKNOWLEDGEMENTS

I wish to thank the directors and curatorial staff of the South African

Museum, Cape Town, the Sedgwick Museum, Cambridge, and the Natural -

History Museum, London, for allowing me access to the collections in their

care. In particular I am indebted to Drs J. N. Theron and J. E. Almond of the

South African Geological Survey for their willingness to share their knowledge

of fossiliferous localities within the Bokkeveld Group of the western Cape and

for permitting access to their collections. Sincere thanks are due to A. J.

Boucot, P. E. Isaacson and P. R. Racheboeuf who reviewed the manuscript and

whose comments helped improve the final product.

DEVONIAN CHONETACEAN BRACHIOPODS FROM SOUTH AFRICA 179

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THOMAS, I. 1905. Neues Beitrage zur Kenntnis der devonischen Fauna Argentiniens.

Zeitschrift der Deutschen geologischen Gesellschaft 57: 233-290.

Toots, H. 1965. Orientation and distribution of fossils as environmental indicators.

Guidebook. 19th Annual Field Conference, Wyoming Geological Association: 219-229.

ULRiIcH, A. 1893. Palaeozoische Versteinerungen aus Bolivien. Jn: STEINMANN, G. Beitrdge _

zur Geologie und Paldontologie von Stidamerika. Neues Jahrbuch fiir Mineralogie,

Geologie und Paldontologie (Beilagebande) 8: 5-116.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particu-

larly 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 trans-

ferred 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 specimens

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, descrip-

tion 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 Eliza-

beth (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. DuToit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably 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. The generic name should

not be abbreviated at the beginning of a sentence or paragraph.

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.