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

Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

bered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 X 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

into consideration.

All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 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.:

‘Smith (1969) describes... .’

‘Smith (1969: 36, fig. 16) describes . . .’

‘As described (Smith 1969a, 1969b; Jones 1971)’

‘As described (Haughton & Broom 1927)...”

‘As described (Haughton et al. 1927)...”

Note: no comma separating name and year

pagination indicated by colon, not p.

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, 1969b) 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 (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number 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 la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHER, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthro-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

February 1989 Februarie

Part 7 Deel

THE AMMONITE SUBFAMILY

LABECERATINAE SPATH, 1925:

SYSTEMATICS, PHYLOGENY, DIMORPHISM

AND DISTRIBUTION

(WITH A DESCRIPTION OF A NEW SPECIES)

HERBERT CHRISTIAN KLINGER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

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

6(1, t.-p.i.), 7(1—4), 8) 9(1-2, 7), 10(1-3),

11(1-2, 5, 7, t.—-p.i.), 14(1-2), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33, 36(2), 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 101 6

Printed in South Africa by In Suid-Afrika gedruk deur

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

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE AMMONITE SUBFAMILY LABECERATINAE SPATH, 1925:

SYSTEMATICS, PHYLOGENY, DIMORPHISM AND DISTRIBUTION

(WITH A DESCRIPTION OF A NEW SPECIES)

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum,

Cape Town

(With 18 figures)

[MS accepted 20 October 1988]

ABSTRACT

Systematics of the subfamily Labeceratinae Spath, 1925, are discussed. Data from Zulu-

land suggest that the subfamily probably consists of a single, dimorphic genus, to which the

names Labeceras and Myloceras have been applied. Distribution of the subfamily shows con-

centration in two main areas—southern Africa—Madagascar and Australia. Comparisons of the

faunas of these suggest a high degree of endemism, and also show that post-mortal drift is neg-

ligible. Absence of labeceratine faunas from the south Atlantic—with the exception of a single

locality in the Austral Basin of southern Patagonia—may possibly be linked to the opening of

the Atlantic and the associated different environments, but also the differential ecological

requirements of the different ammonite taxa, of which we know nothing.

CONTENTS

PAGE

pL Lg oh lat bP ar 189

Systematics, phylogeny and dimorphism ................... 190

NE RRR ae ee lee LY sj cis ite 8S oles bd 2 else's ~ 200

Biogeographic and stratigraphic implications ............... 204

Perennial GO! ANEW SICCICS 5... os - ee ee cc eee eeeees ait

ee MM IIEENAIS, Gig glo es ee oe ee eine ee teens 214

Ce SUS ee Ee ZAS

INTRODUCTION

The ammonite subfamily Labeceratinae Spath, 1925, is a small group of

heteromorphs, restricted to the Upper Albian, and generally assumed to be

south Gondwanid or Austral in distribution (Collignon 1932: 25; Venzo 1936:

124; Collignon 1950: 84; Wiedmann 1965: 443; Kennedy 1972: 402; Forster

1975: 173; 1981: 168; Klinger 1976: 37; McNamara 1978: 240). The size of the

group, conspicuous morphology, and short temporal and limited geographical

occurrence is ideal for studying some identifiable factors that may have in-

fluenced the palaeobiogeographical distribution.

189

Ann. S. Afr. Mus. 98 (7), 1989: 189-219, 18 figs.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATICS, PHYLOGENY AND DIMORPHISM

Taking Wright’s (1957: L231—L232) classification as a starting point, the

systematics of the group may be discussed.

Superfamily SCAPHITACEAE Meek, 1876

Family Labeceratidae Spath, 1925

(= Aleteceratidae Whitehouse, 1926; Myloceratidae Spath, 1939)

Genus Labeceras Spath, 1925

Subgenus Labeceras (Labeceras) Spath, 1925

Subgenus Labeceras (Appurdiceras) Whitehouse, 1926

Genus Myloceras Spath, 1925

Genus Ellipsoceras Collignon, 1950

(= Abadieceras Collignon, 1950)

? Genus Hamitoides Spath, 1925

This classification was based mainly on ideas of Spath (1925: 191, et seq.;

1930: 57; 1939: 600-601), many of which no longer hold true or need updating.

The alleged scaphitoid affinity of the subfamily as advocated by Spath (1925:

191; 1939: 601), and followed by Wright (1957: L231) and Luppov & Drusch-

chits (1958: 125), was based on superficial homoeomorphy and incorrect

interpretation of sutural data. It was questioned by Schindewolf (1961: 109) and

convincingly disproved by Wiedmann (1962: 84; 1965: 443) and Wiedmann &

Dieni (1968: 74), who would rather place the group in the family Anisocerati-

dae, a view followed by Reyment (19646: 37), Klinger (1976: 36) and

McNamara (1978: 232). But, as Forster (1975: 172) has pointed out, the trifid

lobes are more like those of Ancyloceratinae rather than Anisoceratidae with

predominantly bifid lobes. However, the stratigraphic gap between the last

occurrence of Ancyloceratinae at the top of the Lower Albian, and the first

occurrence of Labeceratinae at the base of the Upper Albian has to be bridged.

I previously (Klinger 1976: 41, pl. 12 (figs 3-4, 6), text-figs 7i-j, 8a),

recorded Labeceras sp. nov. aff. L. crassicostatum from the third, possibly fourth,

division of the Albian of Zululand. Additional material (Fig. 1) has since been

found that suggests the presence of an as yet unnamed heteromorph with labe-

ceratid coiling and simple sharp ribbing, and another with apparent crioceratitid

coiling and ventrally tuberculate fine ribbing. The sutures have asymmetrically

trifid lobes L, U and I. It would be tempting to regard these as the connecting

link between the last Ancyloceratinae and the first Labeceratinae, as envisaged

by Forster (1975: 172). However, apart from the suture lines, the regular tuber-

culation, style of ribbing and planispiral coiling are very difficult to reconcile

with the first true Labeceratinae in the Upper Albian of Zululand, and I would

rather regard them as homoeomorphic, but ancestrally distinct, hamitids or aniso-

ceratids. Similar faunas were described from the Middle Albian of the Samana

Range of India by Spath (1930) and are worth reinvestigating.

In Zululand, the first forms definitely identifiable with Labeceras and Mylo-

ceras, Labeceras crassetuberculatum Klinger, 1976 (Fig. 2) and Myloceras

THE AMMONITE SUBFAMILY LABECERATINAE 191

rotundum Klinger, 1976 (Figs 3C—E, 4), occur in the lower part of the fifth divi-

sion of the Albian. Next, Labeceras plasticum Spath, 1925 (Fig. 3A) Myloceras

serotinum Spath, 1925 (Figs 5—6), and M. cornucopia Spath, 1925, are common.

This association is followed by common Labeceras inflatum Forster 1975 (= Labe-

ceras ovale Klinger, 1976) (Fig. 7B), L. rectum Klinger, 1976 (Fig. 3B), and

Myloceras besairiei Collignon, 1932. Rare elements in the latter association also

include Ellipsoceras expansum Collignon, 1950. This is slightly oversimplified

(cf. Klinger 1976) but several distinct morphological trends can be observed

(Fig. 8). These are from oldest to youngest.

Labeceras

1. Decrease in overall size.

. Lateral lobe (L) changes from

asymmetric to symmetrically trifid.

. Whorl section becomes more com-

pressed.

. Coiling initially typically labecera-

tid, with curved shaft and inward-

facing aperture. Later forms with

straight shaft, aperture parallel to

shaft, very similar or identical to

Myloceras, and can only be distin-

guished by virtue of the lateral

tubercles.

. Ornament becomes subdued in

later forms and umbilical tubercles

reduced.

. Coiling on

Myloceras

1. Decrease in overall size.

- Lateral lobe. (ij) changes from

asymmetric to symmetrically trifid.

. Whorl section becomes more com-

pressed.

criocone whorls

becomes more regular and com-

pact; distinct change in whorl

section near point of uncoiling.

Shaft possibly tends to recurve in

Ellipsoceras.

. Ornament initially very irregular

with variable number of minor to

major and/or tuberculate to non-

tuberculate ribs. Ribbing later

more regular and tending to

become more uniform.

Given the range of diversity and also overlap of coiling in Labeceratinae, I

doubt if Ellipsoceras deserves separate generic rank. Apart from the curious

coiling, which has so far been observed only in the holotype of Ellipsoceras

expansum (Collignon, 1950, pl. 14(5) (fig. 2)), Ellipsoceras has ornament of the

type of M. besairiei Collignon, and is here included in the synonomy of Myloceras.

Against this background it is now possible to look at the systematic position

of the poorly known genus Hamitoides Spath, 1925, which had in the past either

been referred to the Labeceratinae with doubt (Spath 1925: 191; 1930: 57; 1939:

600; Wright 1957: L232; Klinger 1976: 36-37) or was considered to be a link

between Hamites and Labeceras (Spath 1925: 191; 1939: 601; Haas 1942: 187;

Wiedmann 1962: 98; Wiedmann & Dieni 1968: 74).

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Gen. et. sp. indet (Hamites?), SAM—KK154. Material previously described by

Klinger (1976: 41, pl. 12 (figs 3-4, 6), text-figs 7i-j], 8a) as Labeceras sp. nov. aff.

L. crassicostatum from Albian HI-IV? Note association with Lyelliceras lyelli. X 1.

The type species of Hamitoides, H. studerianus as interpreted in terms of

the lectotype of the species (in Pictet 1847, pl. 15 (fig. 1)), is certainly not a

labeceratid. The straight shaft and rounded whorl section are combinations as

yet unknown in Labeceratinae, especially in view of the fact that the type

material ranges from the Middle Albian to the lower Upper Albian.

Several alleged Hamitoides have been recorded from the Middle Albian of

Madagascar: Hamitoides? madagascariensis Breistroffer (1936: 174, pl. 20

(figs 6-9, fig. 101); Collignon 1963: 45, pl. 258 (figs 114-115)), Hamites studeri

(Collignon, 1932: 22, pl. 4 (figs 10-13)), Hamitoides aff. studeri (Collignon

1963: 43-45, pl. 257 (figs 1110-1111), pl. 258 (fig. 1116)).

The specimens referred to the type species by Collignon (1963) may,

according to Wiedmann & Dieni (1968: 74), possibly belong to Eoscaphites

tenuicostatus (Pervinquiére) and not involve Labeceratinae at all. Hamitoides

madagascariensis 1s too poorly known for definite comment.

Venzo (1936: 111 (53)) recorded two specimens from Zululand as Hamites

studeri. These are misidentified shafts of Labeceras plasticum. Hamitoides sp.

THE AMMONITE SUBFAMILY LABECERATINAE 193

Fig. 2. Labeceras crassetuberculatum Klinger, 1976. A-C. NMB-D578.

D-F. SAM-PCZ205/5/7/, the holotype. G—J. NMB—D2372.

All x 1.

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. A. Labeceras plasticum Spath, 1925. SAS-EM127. x 1. B. Labeceras rectum

Klinger, 1976. NMB-D736. x 2. C-E. Myloceras rotundum Klinger, 1976. NMB-—D496.

x 1.

THE AMMONITE SUBFAMILY LABECERATINAE 195

Fig. 4. Myloceras rotundum Klinger, 1976. A-C. NMB-D429. D-F. NMB-D294a.

G-I. NMB-D294b. All x 1.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

ind. from the Samana Range of India (Spath 1930: 62) was neither figured nor

described and I cannot comment on it.

Hamitoides angolanus Haas (1942: 187, pl. 45 (fig. 3a—e), text-figs 28a—b)

is from the Upper Albian of Hanha, Angola. The holotype consists of a

recurved hook and part of the shaft. It is wholly septate, thus definitely excluding

labeceratine affinities—rather suggesting ptychoceratid affinities. Thus inter-

preted, Hamitoides definitely does not belong in the Labeceratinae.

McNamara (1980: 147) has recently shown that the type species of Labe-

ceras (Appurdiceras) Whitehouse, 1926—Ancyloceras corcycepoides Etheridge

(1905: 14, pl. 1 (figs 3-5), pl. 2 (fig. 4)), is in fact an anisoceratid and should be

removed from Labeceratinae. The subfamily Labeceratinae is thus effectively

reduced to the two genera, Labeceras and Myloceras.

Below generic rank, systematics of the Labeceratinae becomes difficult, due

to the low specific diversity but extreme intraspecific variation. However, one of

the most consistent features of labeceratid systematics is the consistent associa-

tion of Labeceras and Myloceras (e.g. McNamara 1978: 241) throughout the

known stratigraphic range of the group.

Distinct successive pairs of Labeceras and Myloceras can be recognized. In

both there seems to be an increasing trend towards hydrodynamic stability

and/or ‘streamlining’ of ornament. This leads to surprising homoeomorphy

between Labeceras and Myloceras (Fig. 7).

This raises the question of dimorphism in Labeceratinae—i.e. whether

Myloceras is the macroconch and Labeceras the microconch.

Apart from the scaphitids (see e.g. Makowski 1963; Cobban 1969), dimor-

phism has only recently been recognized in heteromorph ammonites and

remains largely to be explored in Cretaceous groups (see Callomon 1981: 267).

Distinct dimorphism has been recognized in the baculitid genera Scipono-

ceras (Marcinowski 1980: 253; see also Kennedy & Juignet 1983: 17), Lechites

(Cooper & Kennedy 1977), Baculites (Kennedy 1984: 143—in Baculites incurva-

tus; Klinger & Kennedy in prep.—in Baculites capensis), and in the hetero-

ceratine species Colchidites vulanensis (Aguirre Urreta & Klinger 1986: 350). It

has also been suggested for several other groups, e.g. Bostrychoceras (Kennedy

1986: 95), Eubostrychoceras (Kennedy 1986: 101), Hamites (Cooper, unpub-

lished data; Kennedy & Juignet 1983: 12), Tridenticeras (Kennedy 1984: 138),

and ‘Neancyloceras’ (Klinger 1982: 229).

In most of these groups, dimorphism manifests itself mainly by differences

in size and, where preserved, in apertural modifications. This would correspond

more or less to Type II dimorphism of HouSa (1965), where the sexes differ not

only in size, but where the male has apertural outgrowths, such as lateral lappets

or a ventral rostrum.

Dimorphism has previously been suggested for Labeceratinae but, in my

opinion, for the wrong reasons. Spath (1925: 192) suggested that forms with and

those without croziers (in Myloceras) might be dimorphs. This would be a situa-

tion analogous to that found in Macroscaphites and Costidiscus, as suggested by

THE AMMONITE SUBFAMILY LABECERATINAE 197

Fig. 5. Myloceras serotinum Spath, 1925. SAS-—Z174. x 1.

Callomon (1981: 267). However, the Zululand material does not support this

view.

Forster (1975: 176) suggested that L. plasticum crassum, which consists of

predominantly large, inflated, strongly tuberculate forms, may differ from the

generally smaller and more weakly tuberculate L. plasticum plasticum on

account of sexual dimorphism. By analogy with the scaphitids (e.g. Makowski

1963: 31, et seq.; Cobban 1969), this is the situation one would expect in Labe-

ceras. Also, by analogy with the scaphitids, we would expect the range in size to

LOS ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Myloceras serotinum Spath, 1925. A-B. SAS—Z174. x 1.

be such that the largest microconchs would be larger than the smallest macro-

conch, but not as large as the largest macroconch. In the available material we

find that the coarsely ornamented forms are generally larger than the more

finely ornamented forms, but that the size overlap of these forms is virtually

1:1. Size distribution patterns of L. crassetuberculatum, L. plasticum, L. rectum

and Myloceras serotinum are shown in Figure 9. Apertural lappets are generally

taken to be characteristic of micromorphs. Spath (1925: 192, pl. 31 (fig. 4b-c))

noticed that in Labeceras plasticum ‘The aperture is provided with a short dorsal

THE AMMONITE SUBFAMILY LABECERATINAE 199

Fig. 7. A. Myloceras serotinum Spath, 1925. SAM-PCZ7665. B. Labeceras

inflatum Forster, 1975. NMB-D2702. Both x 1. Note homoeomorphy between

Labeceras and Myloceras suggesting dimorphism.

and two longer lateral lappets’ and in Myloceras serotinum (1925: 193, pl. 33

y (fig. 1)) ‘The aperture, as restored .. . shows a slight ventral and two more

| prominent lateral lappets’. Subsequently Wright (1953: 473) noted that *...

Labeceras has lateral lappets, whereas the probably closely related Myloceras

has an aperture merely with a sinuous border’.

Material from Zululand shows that lateral lappets occur in both Labeceras

and Myloceras but, that in relation to overall shell size, those of Labeceras are

much larger than those of Myloceras (Figs 13-15).

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Thus, unless we are dealing with isochronous parallelism or homoeomorphy,

it seems plausible to regard Labeceras and Myloceras as a dimorphic pair—the

former being the microconch, the latter the macroconch.

I admit that this may appear as an extreme form of dimorphism not pre-

viously recorded in other heteromorph groups. It would be ideal to show that

the nuclei of Labeceras and Myloceras are identical, thus lending further cred-

ibility to the assumption that they are dimorphs. Unfortunately nuclei are not

available and, at the smallest diameters preserved, Myloceras is distinctly tuber-

culate and Labeceras non-tuberculate. However, if they are indeed dimorphs,

the comparisons proposed by Forster (1975: 172-173) between Myloceras and

Australiceras, and Labeceras and Toxoceratoides, may not be as preposterous as

they seem—although the taxonomic implications are frightening.

This systematic arrangement of the subfamily Labeceratinae seems far

removed from that adopted by Wright (1957: L231—L232) but, in the light of

current data and from a phylogenetic point of view, makes much more sense.

This effectively reduces the subfamily Labeceratinae to one dimorphic genus.

Until details of the complex synonymies have been worked out, I prefer to

retain both generic names Labeceras and Myloceras in this discussion, albeit

mainly for the sake of taxonomic ‘neatness’.

Within this systematic framework it is now possible to look at the strati-

graphic and geographic distribution of the subfamily Labeceratinae.

DISTRIBUTION

Records of Labeceras from the Albian of England (Owen 1971: 195) are

based on misidentifications of a labeceratid homoeomorph, /diohamites ellip-

ticoides Spath, 1939 (see Kennedy 1972: 400-404). All other records of Labe-

ceratinae are of Gondwanid origin (Fig. 10). These include:

1. Australia. Occurrences here are in the Great Artesian Basin of Queensland

and in South Australia (see Ludbrook 1966 for summary). Individual references

include McCoy (1867), Etheridge (1909—South Central Queensland), White-

house (1926—South Central Queenland), Reyment (1964a, 1964b—South Aus-

tralia), and McNamara (1978—Central Queensland).

Precise data are not available, but the Myloceras—Labeceras assemblages

have been dated as middle Late Albian (McNamara 1978: 231), or as being

equivalent to the varicosum—aequatoriale subzones of the Gault of England

(Spath 1925: 194) or possibly the orbignyi subzone (Whitehouse 1926: 197; also

Reyment 1964a: 34).

2. New Guinea. Glaessner (1958: 217-218) recorded Myloceras davidi White-

house, Myloceras cf. flindersi (McCoy), and Labeceras trifidum Whitehouse

from the Albian of New Guinea.

3. New Zealand? Wellman (1959: 121) and Henderson (1973: 99) recorded a

single questionable fragment of Myloceras from the Clarence Series at Waira-

rapa on North Island.

THE AMMONITE SUBFAMILY LABECERATINAE 201

Labeceras rectum

Labeceras plasticum

Labeceras crassetuberculatum

Myloceras rotundum

Fig. 8. Suggested phylogenetic sequence of Labeceras and Myloceras in Zululand. All reduced

by various fractions to fit diagram.

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

4. Antarctica? Thomson (1984: 89) mentioned small heteromorphs ‘possibly

Labeceras’ from the Albian of Lost Valley on James Ross Island in Lesser Ant-

arctica.

5. Somalia? Tavani (1942: 30 (116), pl. 3 (pl. 12) (fig. 8)) recorded a single

specimen from the Albian of Scec Gure (Bugda Acable), Somalia, as Labeceras

plasticum. The specimen is very poorly preserved but the presence of straight,

near-parallel shafts seems to indicate ‘Hamitoides’ rather than Labeceras plasti-

cum affinities.

6. Madagascar. A very well-documented labeceratine fauna is known from

Madagascar. Individual references include Boule et al. (1907—Andrakaka,

Diego-Suarez), Besairié (1932), Collignon (1932—Mont Raynaud), Collignon

(1936— Maniamba-amba), Collignon (1950—Mokaraha), Collignon (1951—

Andranofotsy, Manja), Collignon (1963—Diego-Suarez, Betioky (Vohimara-

nitra), Andronofotsy (Manja), Mokaraha (Soalala)), and Besairié & Collignon

(1972)

This fauna is well dated and, according to the biozonation compiled for

Madagascar by Collignon (1963), ranges from the lowermost zone of the Upper

Albian, Zone a Dipoloceras cristatum, through the Zone a Hysteroceras binum to

the penultimate Zone a Pervinquieria inflata.

The systematic position of Euhemihoplites paradoxus Collignon (1964: 38,

pl. 326 (fig. 1457)), reported from the Lower Cenomanian, Zone a Mantelli-

ceras mantelli et Calycoceras newboldi, should be mentioned here. According to

oOo

Oo = NS tor ee On) > 3) (COL sO

er pe ee ar iss

NSS 20 21225723 eke 25 70 80 90 100 110 120 130140

Labeceras rectum Myloceras serotinum

n= n=8

WIDTH CROZIER IN MM/NUMBER SPECIMENS

oOo

‘oO

So Ro) Ww) ee nl oS =a

sy SS IS (6 Way eS IE oe)

20) 29) 2223) 24 25) 26) 2728) 29) 30) 31) 032838) 34535) 3637 26 27 2 29 30 31 32 33 34 35 36 37 38 39 40 41 42

Labeceras plasticum Labeceras crassetuberculatum

n=36 n= 21

Fig. 9. Size distribution in Labeceras and Myloceras.

G 24

THE AMMONITE SUBFAMILY LABECERATINAE 203

Kennedy (1985), this species, based on a single fragment, may in fact be a Mylo-

ceras, probably derived from Upper Albian strata. It is similar to a new species

described below in possessing umbilical tubercles.

7. Mozambique. The original labeceratid material on which Spath’s (1925: 191,

et seq.) descriptions were based, are probably from near Catuane in southern

Mozambique—Forster’s (1975: 32) locality Lagoa Catuine. Other localities

yielding labeceratids include Rio Zepundune, Road Metal Quarry 9,5 km north

of Catuane, Rio Comane, and Rio Changalane.

The stratigraphic range of labeceratids in southern Mozambique was com-

pared to the standard European succession of varicosum to aequatoriale

subzones of the Mortoniceras inflatum Zone.

8. Zululand. Labeceratids are locally very common in Kennedy & Klinger’s

(1975: 276) fifth division of the Albian. References include Besairié (1930:

622—Mzzinene), Venzo (1936: 113 (55), et seqg.—Ndabana-Mzinene), Kennedy

& Klinger (1975: 276), Klinger (1976: 36, et seqg.—Mzinene River, Ndambana

Creek, Ndumu, Mkuze Game Reserve, Munywana River), and Haughton (1936:

292— Pongola River).

Definite Labeceras and Myloceras first occur in the fifth division of the

Albian in association with Dipoloceras cristatum, ‘Deiradoceras’ , ‘Drepanoceras’

and ‘Rhytidoceras’. Later forms of the group Myloceras besairiei, with extremely

compressed whorl section and regular ornament, and Labeceras rectum occur in

association with Pervinquieria (P.) and P. (Styphloceras). In comparison with

oe Labeceras

eae

Fig. 10. Palaeobiogeographic distribution of Labeceras and Myloceras. Map based after Smith

& Briden (1973).

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Madagascar, this would suggest a definite range of Upper Albian, Zone a Dipo-

loceras cristatum to Zone a Pervinquieria inflata.

9. Argentina (Patagonia). Leanza (1970: 202, 209) recorded two new genera,

Calliscaphites, type species Calliscaphites andinus Leanza, 1970, and Paralepto-

ceras, type species Paraleptoceras singulare Leanza, 1970, from the Albian of

Estancia La Vega, in the Santa Cruz Province of Patagonia. These are clearly

synonyms of Myloceras Spath and Labeceras Spath and, from the limited figured

material available, seem close to Myloceras rotundum Klinger and Labeceras

crassetuberculatum Klinger.*

Subsequent reports from the Santa Cruz Province include Nullo et al. (1981,

pl. 2 (fig. 10—Labeceras? sp., fig. 11— Calliscaphites andinus)) and Riccardi &

Rollieri (1980: 1198).

Medina & Rinaldi (1986) recorded both Myloceras and Labeceras from

Estancia La Vega in Santa Cruz Province, as do Riccardi et al. 1988. Unfortu-

nately, this is the only known locality in the Austral Basin with labeceratids and,

in addition, the outcrop is rather small.

BIOGEOGRAPHIC AND STRATIGRAPHIC IMPLICATIONS

Distribution of the Labeceratinae is strictly Gondwanid—more specifically

south Gondwanid. Two features are conspicuous:

1. Concentration of Labeceratinae into two main regions—south-eastern

Africa—Madagascar and Australia.

2. Conspicuous absence of Labeceratinae in India and, with the exception of

the Austral Basin of Patagonia, absence in the South Atlantic.

South-eastern Africa—Madagascar and Australia

Correlation of Labeceratinae faunas between south-eastern Africa—Mada-

gascar and Australia is tenuous, mainly due to a combination of taphonomy,

differential preservation and systematics. Complete specimens of Myloceras and

Labeceras are rare, and most descriptions are based on incomplete material.

To this must be added the effects of disparate sizes and intraspecific varia-

tion. Some of the Australian species of Myloceras are gigantic, e.g. Myloceras

axononoides (Etheridge, 1909, pl. 44 (fig. 2)) and Myloceras nautiloides (Ether-

idge, 1909, pl. 45), and no Afro—Malagassy specimens are known that even

approximate the Australian giants in size (Figs 11-12).

It is difficult to evaluate the reports of species common to both areas (e.g.

Venzo 1936: 55 (113); Reyment 1964a: 23; Forster 1975: 176; Klinger 1976).

From these records it appears that the only species probably common to south-

eastern Africa-Madagascar and Australia, are Labeceras bryani Whitehouse

and L. plasticum Spath.

Out of a total of about 40 described species, this is indeed a very low corre-

lation factor. However, even given the effects of intraspecific variation, analysis

of the faunas still shows some differences.

* See p. 219 for note added at proof stage.

THE AMMONITE SUBFAMILY LABECERATINAE 205

oO cm 10

Fig. 11. Variation in size in Myloceras. A. Myloceras serotinum Spath, 1925. SAS—Z174.

B. ‘Ellipsoceras’ expansum Collignon, 1950. Holotype. C. M. axonoides Etheridge, 1909.

Holotype. Half size compared to other specimens. D. M. serotinum Spath, 1925. SAM- 7665.

E. M. cornucopia Spath, 1925. Holotype. F.M. amaltheia Spath, 1925. Holotype.

G. M. mokaharense Collignon, 1964. Holotype. H.M. auritulum McNamara, 1978.

Holotype. I. M. serotinum rugosa Spath, 1925. Holotype. J. M. serotinum Spath, 1925.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

eee. ae»

ge shee ea

Fig. 12. Variation in size in Labeceras (and one specimen of Myloceras). A. L. plasticum

Spath, 1925, the holotype. B. L. gracilis Collignon (1964, fig. 1140). C. Myloceras serotinum

elliptica Collignon (1965, fig. 1137). D.L. plasticum crassa Spath, 1925, the holotype.

E. L. etheridgei Whitehouse, 1926 (pl. 38 (fig. 2)). F. L. plasticum (Collignon, 1964,

fig. 1125). G. L. papulatum Whitehouse (1926, pl. 39 (fig. 3)). H. L. hourcqi Collignon (1950,

pl. 14 (fig. 1)). I. L. compressum Whitehouse, 1926 (1950, pl. 39 (fig. 5)). J. L. crassicostatum

Collignon, 1950 (pl. 13 (fig. 5)).

THE AMMONITE SUBFAMILY LABECERATINAE 207

Fig. 13. A. Sketch of aperture of Myloceras serotinum. SAM-7665. X 3,9.

B-C. Labeceras plasticum. B. SAM-Z450. C. A2173.

The only reliable feature in the systematics of Myloceras appears to be the

whorl section (see also Reyment 1964a: 31). Taking the phylogenetic sequence

(Fig. 8) as a basis for comparison, three groups can be recognized: Myloceras gr.

ex. rotundum with round to reniform whorl section, Myloceras gr. ex. serotinum

with rectangular depressed to compressed whorl section, and Myloceras gr. ex.

besairiei with rectangular to very compressed whorl section. A similar crude

analysis can be done on Labeceras, based on the nature of the shaft— Labeceras

gr. ex. crassetuberculatum with incurved, asymmetrically coiled shaft, Labeceras

gr. ex. plasticum with distinctly curved shaft, and Labeceras gr. ex. rectum with

straight shaft and very often a distinct nick-point and coiling, virtually as in

Myloceras.

Within this crude framework Australian and Afro—Malagassy forms can be

compared. No forms similar to Myloceras gr. ex. besairiei or Labeceras gr. ex.

rectum are known from Australia, i.e. comparisons are possible amongst strati-

graphic early forms but not amongst the later, established ones.

Though fully aware of the possible errors in the systematics of the sub-

family, the above data do point to significant differences between the Australian

and Afro—Malagassy faunas, and seem to suggest considerable endemism in the

two labeceratine populations, especially as far as the apparent phylogenetically

more advanced forms are concerned. This also implies restricted necroplanktonic

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. Sketch of aperture of Labeceras

crassetuberculatum. EM 151. X 3,9.

dispersal in at least part of the population, which is at variance with the views

held by Reyment (1964a: 33).

India

Data on the Cretaceous of southern India mainly date back to Forbes

(1846), Stoliczka (1863-1865), and Kossmat (1895-1898). The most recent strati-

graphic syntheses of the Cretaceous of southern India are by Sastry et al. (1968)

and Gupta (1975). The lowermost Utatur Group is a mixture of Upper Albian

and Cenomanian. The lower Schloenbachia inflata Zone contains a mixture of

Upper Albian and Lower Cenomanian faunas. According to Spath (1925: 196)

the sequence only starts with the aequatoriale subzone, i.e. the uppermost part

of the European inflatum zone. Thus, on stratigraphic data alone, chances of

finding Labeceratinae in southern India are very slender. New collections being

studied by Dr M. A. Ghare (letter 17.7.84) contain numerous new records of

heteromorph genera, but no labeceratines.

South Atlantic

Apart from one locality, Estancia La Vega, in the Austral Basin of southern

Patagonia, no Labeceratinae are known from the South Atlantic.

The stratigraphic range of the Labeceratinae coincides with part of.the

opening-up of the South Atlantic Ocean. For discussions see e.g. Beurlen

THE AMMONITE SUBFAMILY LABECERATINAE 209

Fig. 15. Sketch of aperture of Labeceras crassetuberculatum.

HOTS. x 3,9:

(1961), Reyment & Tait (1972), Van Andel et al. (1977), Sclater et al. (1977),

Melguen et al. (1979), Reyment (1980), Rand & Mabesoone (1983), etc. It is

possible that the distribution of the group may have been influenced in part by

these circumstances (Fig. 16).

Whatever the exact dates of North and South Atlantic marine connections

may be, it seems reasonable to accept that surface currents existed (Berggren &

Hollister 1974; Lloyd 1982) and that free faunal migration was possible between

south-eastern Africa—Madagascar, via the Cape Sea Route, through the Atlan-

tic, as far north as the Venezuelan Andes, Mexico and Texas during the Upper

Albian (e.g. Kennedy & Cooper 1975: 284-285; Kennedy & Cobban 1976: 77;

Forster 1978: 158; 1981: 167-168; Renz 1981: 201; 1982).

Absence of records of Labeceratinae from the Cape Basin is due to the

absence of onshore exposures. No onshore Albian exposures are known

between southern Angola and Zululand, a distance of some 3 500 km. Offshore

drilling on the Orange River delta may hold some prospects. However, the

absence of Labeceratinae from the Upper Albian of the Brazil-Angola Basin

seems real, and not due to collecting failure, non-exposure of relevant strata or

misidentifications, as various extensive faunal lists from this region seem to sub-

stantiate, e.g. Spath (1922), Haas (1942), Howarth (1965), Reyment (1955,

1956, 1978, 1981), Cooper (1978), Collignon (1978), and Cooper & Kennedy

(1979).

Prevailing conditions in the Upper Albian in the Brazil-Angola Basin seem

to have been favourable for development of endemic faunas, e.g. the mortoni-

ceratine genus Elobiceras (e.g. Reyment 1981: 149), but unfavourable for the

entry of others, e.g. the Labeceratinae, while apparently having no effect on

others.

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

Deep water

Shallow shelf

No sedimentary record

R.G.R/W.R.-Rio Grande Rise/Walvis Ridge

F.P. Falkland Plateau

A.P. Agulhas Plateau

E - Elobiceras

NIGERIA

ANGOLA

MOZAMB IQUE

ZULULAND

Neuquén Basin

San Jorge Basin

Austral or Magallanes

Basin

Fig. 16. Palaeo-reconstruction of opening of the Atlantic Ocean. Map based mainly on

Dingle et al. 1983.

There seems to be little consensus on the physical size and effect of the

Walvis Ridge separating the Angola Basin from the Cape Basin, ranging from a

submerged chain to a series of islands (e.g. various references in Sclater et al.

1977; Bolli et al. 1978; Reyment 1980; Rage 1981), nor on its effects on the dis-

tribution of invertebrate faunas (e.g. Scheibnerova 1978, 1981; Tambareau

1982; Dingle 1984). However, as the South Atlantic progressively opened, circu-

lation in the Cape Basin seems to have kept one step ahead of that of the

Angola Basin, especially after the Falkland Plateau had cleared the tip of Africa

at about 100 Ma.

THE AMMONITE SUBFAMILY LABECERATINAE pA

In addition, climatic effects have to be taken into consideration. Plant

material from DSDP site 361 suggests a temperate climate and well-vegetated

onshore environment off the Cape Basin (although this is questioned by

McLachlan & Pieterse (1978), who suggest a harsh, probably dry climate),

whereas in Angola the climate was probably warm and arid to semi-arid (e.g.

Siesser 1978: 970; Natland 1978; Melguen 1978). Apart from differences in

water temperature and also probably salinity and density, these climatic dif-

ferences are clearly reflected by the different dominant sedimentary facies in the

two basins—that of the Cape Basin being predominantly terrigenous (reflecting

active erosion and transport by river systems), whereas that of the Angola Basin

is more calcareous and pelagic, in accordance with arid climate and low sedi-

ment supply. This in turn may be related to differential nutritional resources in

the basins.

To summarize, it would seem simplistic to seek a single cause for the lack of

Labeceratinae in the eastern South Atlantic. Prevailing surface currents may

have been adequate to assist migration of the Labeceratinae into the Atlantic,

but a combination of factors, including physical and chemical differences in the

Argentine—Cape and the Brazil—Angola Basins, separated by the aseismic Rio

Grande Rise and Walvis Ridge, may have acted selectively in producing this

palaeobiogeographical distribution pattern. To these must be added the prime,

albeit virtually unknown, factors of differential ecological requirements of the

different ammonite taxa, which probably completely overshadow all the other

restricting factors.

DESCRIPTION OF A NEW SPECIES

Myloceras bituberculatum sp. nov.

Figs 17-18

Type

Holotype is SAM—PCZ7299a (ex Van Hoepen Collection, uncatalogued

specimen); probably from the lower reaches of the Mzinene River, Zululand,

Mzinene Formation, Upper Albian.

Material

Paratypes are SAM-—PCZ7299b, PCZ7300-—7304, PCZ8331—8338 (ex Van

Hoepen Collection, uncatalogued specimens); probably all from the lower

reaches of the Mzinene River, Zululand, Mzinene Formation, Upper Albian.

Description

The material consists of crioceratitid phragmocones only; the uncoiled

section is unknown. Coiling is relatively tight for Myloceras, with successive

whorls just touching, and the ventral ribs and tubercles of the preceding whorls

slightly impressed in the dorsum of succeeding whorls. Umbilical width increases

212 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. Myloceras bituberculatum sp. nov. A-—C. SAM-—PCZ7301. D-E. SAM—PCZ8331.

D. xX 2, E. x 1. F-G. SAM-PCZ8332. F. x 2, G. x 1. H. SAM—PCZ7299a, holotype. x 1.

I-J: SAM=PCZ8333. T..* 2/3. x 1.

THE AMMONITE SUBFAMILY LABECERATINAE 2

ae ay,

Fig. 18. Myloceras bituberculatum sp. nov. A. Suture line of

SAM-PCZ7301. B. Whorl section of SAM-—PCZ7301.

Scale bar to size for A and B.

slightly with growth, in the available material ranging from 36 to 41 per cent of

the diameter.

The whorl section is trapezoidal, higher than wide to very compressed, with

a narrower venter than dorsum. The flanks are virtually flat (Fig. 18B).

Ornament is very robust, consisting of prominent umbilical tubercles, bifur-

cating and intercalated ribbing and ventral tubercles. The umbilical tubercles are

situated on the outer edge of the umbilical wall, and are conical to radially

elongated. From these a prorsiradiate, slightly sinusoidal pair of ribs generally

arises. In addition, intercalatory ribs may arise at the umbilical edge or up to a

midflank position. On the early whorls of some specimens single ribs, arising at

the umbilical tubercles, alternate with intercalatory ribs arising at midflank.

Ventral tuberculation is variable, both within the same or between different

specimens. All ribs may bear a pair of distinctly clavate tubercles or major ribs,

with prominent clavate tubercles, may alternate with weaker ribs, either without

tuberculation at all, or only with weak, conical ventral tubercles. All ribs cross

the venter with a slight forward curvature and broadening.

The suture line is only partially exposed in PCZ7301 (Fig. 18A).

ci.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions

Specimen D Wb Wh Wb/Wh U

number

PCZ7299a 62 19 (30,6) 21 (33,8) 0,90 24 (33,8)

PCZ7299b 61 16 (26,2) 21 (34,4) 0,76 25 (41)

PCZ7300 18 (1) 23 (—) 0,78 —

PCZ7301 51 15 (29,4) 18 (33,3) 0,83 21 (41,2)

PCZ13502 36 10 (27,8) 15.(4157) 0,67 13:(36, 1)

PCZ7303 — 12 (—) 16 (—) 0,75 —

PCZ7304 24 7 (29,2) 9:(37,3) 0,78 O(375)

PCF S552 26 6 (23,0) 10:G8,5) 0,60 10 (38,5)

PCZS8333 21 5,2 (20,2) 8 (38,1) 0,69 8,5 (40,5)

Discussion

This species and M. paradoxus (Collignon, 1964) (see Kennedy 1985:

307-309, figs 11-12) are unique amongst Myloceras in possessing distinct

umbilical tubercles. Myloceras bituberculatum differs from M. paradoxus in

having much stronger and wider-spaced ribbing. As far as whorl section and fre-

quency of ribbing on the venter are concerned, affinities may be sought with

M. besairiei Collignon and M. robustum Klinger, but the presence of umbilical «|

tubercles clearly distinguishes the present material.

Forster (1975: 185) described part of a crozier with incipient umbilical

tubercles from Mozambique as M. cornucopium. This fragment has a whorl-

section that is wider than high, and I doubt if it can be referred to the present

species.

Due to the imprecise locality data I do not know what—if at all, the

accompanying Labeceras dimorph is.

Occurrence

Upper Albian of Zululand.

ACKNOWLEDGEMENTS

A condensed version of this paper was presented in Tiibingen in 1985 at the

Second Cephalopod Symposium. Assistance by the Alexander von Humboldt

Foundation, the Council for Scientific and Industrial Research, and the South

African Museum is gratefully acknowledged. Thanks are also due to Drs A. Ric-

cardi (La Plata) and M. B. Aguirre Urreta (Buenos Aires), and Dra G. Blasco

de Nullo (Buenos Aires) for showing me the material from Estancia La Vega. I

am grateful to Drs W. J. Kennedy, C. W. Wright (Oxford), R. A. Henderson

(Townsville), and R. McNamara (Perth), who commented on the manuscript;

THE AMMONITE SUBFAMILY LABECERATINAE 21D

however, I take responsibility for the opinions expressed. Technical assistance

by Mss S. Dove, J. Blaeske, and M. Joubert (S.A. Museum) is also gratefully

acknowledged. Thanks are due to Mrs S. Saven for typing several drafts of the

manuscript.

REFERENCES

AGUIRRE UrreTA, M. B. & KLINGER, H. C. 1986. Upper Barremian Heteroceratinae (Cepha-

lopoda, Ammonoidea) from Patagonia and Zululand, with comments on the systematics of

the subfamily. Annals of the South African Museum 96 (8): 315-358.

BERGGREN, W. A. & Ho.uister, C. D. 1974. Paleogeography, paleobiography and the history

of circulation in the Atlantic Ocean. In: Hay, W. W. ed. Studies in paleo-oceanography.

Special Publications. Society of Economic Paleontologists and Mineralogists 20: 126-186.

Besairi£, H. 1930. Les rapports du Crétacé Malgache avec le Crétacé de l’Afrique australe.

Bulletin de la Société géologique de France (4) 30 (7): 613-643.

BEsairi£E, H. 1932. Fossiles caractéristiques du nord et du nord-ouest de Madagascar. Annales

géologiques du Service des mines, Madagascar 2: 37-53, pls 5-6.

BESAIRIE, H. & COLLIGNON, M. 1972. Géologie de Madagascar. I. Les terrains sédimen-

taires. Annales géologiques de Madagascar 35: 1-463, pls 1-89.

BEURLEN, K. 1961. Die palaogeographische Entwicklung des Sudatlantischen Ozeans. Nova

acta Leopoldina (Neue Folge) 24 (154): 1-36.

Boul, H. M., Ryan, W. B. F., FORESMAN, J. B., HoTTMAN, W. E., KAGAmi, H., LONGorIA,

J. F., McKnicut, B. K., MELGUEN, M., NATLAND, J., PRoto-DEcIMA, F. & SIESSER,

W.G. 1978. Initial Reports of the Deep Sea Drilling Project 40: 1-1079.

Boute, M., Lemoine, P. & THEVENIN, Q. 1906-1907. Paléontologie de Madagascar. III.

Céphalopodes crétacés des environs de Diego-Suarez. Annales de paléontologie 1 (1906):

1-20 (173-192); 2 (1907): 21-76 (1-56).

BREISTROFFER, M. 1936. Fossiles de l’albien moyen (niveau supérieur). /n: BESAIRIE, H. La

géologie du nord-ouest. Mémoires de l’Académie malgache 21: 167-176.

CALLomoNn, J. H. 1981. Dimorphism in ammonoids. /n: House, M. R. & SENIor, J. R. The

Ammonoidea. Systematics Association. Special Volume 18: 257-273.

CosBAN, W. A. 1969. The Late Cretaceous ammonites Scaphites leei Reeside and Scaphites

hippocrepis (DeKay) in the Western Interior of the United States. Professional Papers.

United States Geological Survey 619: 1-29.

CoLiicNon, M. 1932. Les ammonites pyriteuses de l’Albien supérieur du Mont Raynaud a

Madagascar. Annales géologiques du Service des mines, Madagascar 2: 5-35.

CoLLIGNoNn, M. 1936. Fossiles de l’Albien supérieur de Maniamba-amba. /n: BEsairi£, H.

La géologie du nord-ouest de Madagascar. Mémoires de l’Académie malgache 21:

190-198.

CoLLicGNon, M. 1950. Rechérches sur les faunes albiennes de Madagascar. Annales géo-

logiques du Service des mines, Madagascar 17: 6-85.

COLLIGNON, M. 1951. Rechérches sur les faunes albiennes de Madagascar. V. L’albien

supérieur d’Andranafotsy (Cercle de Manja). Annales géologiques du Service des mines,

Madagascar 19: 7-40.

CoLLicNon, M. 1963. Atlas des fossiles caractéristiques de Madagascar (Ammonites). X.

Albien. Tananarive: Service Géologique.

CoLLicNon, M. 1964. Atlas des fossiles caractéristiques de Madagascar (Ammonites). X1.

Cénomanien. Tananarive: Service Géologique.

CoLLiGNon, M. 1978. Ammonites du Crétacé Moyen-Supérieur de l’Angola. Estudos de

Geologia e Paleontologia e de Micologia, Lisboa 1977: 1-75.

Cooper, M. R. 1978. The mid-Cretaceous (Albian—Turonian) biostratigraphy of Angola.

Annales du Museum d’histoire naturelle de Nice 4 (1976): XVI.1—XVI.22.

Cooper, M. R. & KeNnnepy, W. J. 1977. A revision of the Baculitidae of the Cambridge

Greensand. Neues Jahrbuch fiir Geologie und Paldontologie. Monatshefte 11: 641-658.

Cooper, M. R. & KENNEDY, W. J. 1979. Uppermost Albian (Stoliczkaia dispar Zone) ammo-

nites from the Angolan littoral. Annals of the South African Museum 77 (10): 175-308.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

DINGLE, R. V. 1984. Mid-Cretaceous Ostracoda from southern Africa and the Falkland

Plateau. Annals of the South African Museum 93 (3): 97-211.

DINGLE, R. V., StessER, W. G. & Newron, A. R. 1983. Mesozoic and Tertiary geology of

southern Africa. Rotterdam: Balkema.

ETHERIDGE, R. 1905. Contributions to the palaeontology of South Australia. No. 14. Creta-

ceous fossils from Dalhousie Springs. Parliamentary Paper South Australia 71: 13-17.

ETHERIDGE, R. 1909. Lower Cretaceous fossils from the sources of the Barcoo, Ward and Nive

rivers south central Queensland. Part II. Cephalopoda. Records of the Australian Museum

7 (3): 135-165, pls 30-49.

Forses, E. 1846. Report on the fossil invertebrata from Southern India, collected by Mr Kaye

and Mr Cunliffe. Transactions of the Geological Society (2) 7 (3): 97-174, pls 7-19.

FORSTER, R. 1975. Die geologische Entwicklung von Stid-Mozambique seit der Unterkreide

und die Ammoniten-Fauna von Unterkreide und Cenoman. Geologisches Jahrbuch 12:

3-324.

FORSTER, R. 1978. Evidence for an open seaway between northern and southern proto-

Atlantic in Albian times. Nature 272 (5649): 158-159.

FOrSTER, R. 1981. Mozambique. /n: REYMENT, R. A. & BENGtTson, P. eds. Aspects of mid-

Cretaceous regional geology: 161-174. London, New York, Toronto, Sydney, San Fran-

cisco: Academic Press.

GLAESSNER, M. F. 1958. New Cretaceous fossils from New Guinea. With a contribution on a

new ammonite genus by R. Casey. Records of the South Australian Museum 13 \2):

199-226, pls 24-26.

Gupta, V. J. 1975. Indian Mesozoic stratigraphy. Delhi: Hindustan Publishing Corporation.

Haas, O. 1942. The Vernay Collection of Cretaceous (Albian) ammonites from Angola.

Bulletin American Museum of Natural History 81: 1-224, pls 1-47.

HauGuTon, S. H. 1936. Account of the geology of the Cretaceous beds and a preliminary

analysis of the associated ammonite fauna. Jn: RENNIE, J. V. L. Lower Cretaceous lamelli-

branchia from northern Zululand. Annals of the South African Museum 31 (3): 283-297.

HENDERSON, R. A. 1973. Clarence and Raukumara Series (Albian—?Santonian). Ammonoi-

dea from New Zealand. Journal of the Royal Society of New Zealand 3 (1): 71-123.

HouSsa, V. 1965. Sexual dimorphism and the system of Jurassic and Cretaceous Ammonoidea

(preliminary note). Casopis Ndrodniho musea 134: 33-351.

Howartn, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bulletin of the

British Museum (Natural History) (Geology) 10 (10): 337-412.

KENNEDY, W. J. 1972. The affinities of Jdiohamites ellipticoides Spath (Cretaceous Ammonoi-

dea). Palaeontology 15 (3): 400-404.

KENNEDY, W. J. 1984. Systematic palaeontology and stratigraphic distribution of the ammo-

nite fauna of the French Coniacian. Palaeontological Association. Special Papers in

Palaeontology 31: 1-160.

KENNEDY, W. J. 1985. A note on Euhemihoplites paradoxus Collignon, 1964 (Cretaceous

Ammonoidea). Cretaceous Research 6 (3): 307-309.

KENNEDY, W. J. 1986. Campanian and Maastrichtian ammonites from northern Aquitane,

France. Palaeontological Association. Special Papers in Palaeontology 36: 5-145.

KENNEDY, W. J. & CoBBAN, W. A. 1976. Aspects of ammonite biology, biogeography, and

biostratigraphy. Palaeontological Association. Special Papers in Palaeontology 17: 1-94,

pls 1-11.

KENNEDY, W. J. & Cooper, M. R. 1975. Cretaceous ammonite distribution and the opening of

the South Atlantic. Journal of the Geological Society, London 131 (3): 283-288.

KENNEDY, W. J. & JUIGNET, P. 1983. A revision of the ammonite faunas of the type Cenoma-

nian: 1. Introduction, Ancyloceratina. Cretaceous Research 4 (1): 3-83.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, Stratigraphy. Bulletin of the British Museum (Natural History)

(Geology) 25 (4): 263-315, pl. 1.

KLINGER, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Memoirs. Geo-

logical Survey, South Africa 69: 1-142.

KLINGER, H. C. 1982. Revision of Ancyloceras bipunctatum Schliter, 1872 (Cephalopoda,

Ammonoidea) and discussion of the validity, phylogeny and limits of the genus Neancylo-

ceras Spath, 1926. Annals of the South African Museum 90 (5): 219-239.

THE AMMONITE SUBFAMILY LABECERATINAE Pal

Kossmat, F. 1895-1898. Untersuchungen tber die Siidindische Kreideformation. Beitrage zur

Paldontologie und Geologie Ostereich-Ungarns und des Orients 9 (3-4) (1895): 97-203,

pls 15-25; 11 (1) (1897): 1-46, pls 1-8; 12 (3) (1898): 89-152, pls 14-19.

LeanzA, A. F. 1970. Ammonitos nuevos 0 poco conocidos del Aptiano, Albiano y Cenoma-

niano de los Andes Australes con notas acerca de su posicion estratigrafica. Revista de la

Asociacion geologica argentina 25 (2): 197-261.

Lioyp, C. R. 1982. The mid-Cretaceous; paleogeography; ocean circulation and temperature;

atmospheric circulation. Journal of Geology 90 (4): 393-413.

Lupsrook, N. H. 1966. Cretaceous biostratigraphy of the Great Artesian Basin in South

Australia. Bulletin. Geological Survey of South Australia 40: 1-223.

Luprov, N. P. & Drusucuits, V. V. 1958. [Mollusca—Cephalopoda II. Ammonoidea (Cerati-

tida and Ammonitida). Endocochlia, Coniconchia.] Jn: OrLov, Yu. Q. Fundamentals of

Paleontology 6: 9-190. (In Russian.)

MakowskI, H. 1963. Problem of sexual dimorphism in ammonites. Palaeontologica polonica

12: 1-92, pls 1-20.

MarcinowskI, R. 1980. Cenomanian ammonites from German Democratic Republic, Poland,

and the Soviet Union. Acta geologica polonica 30 (3): 215-325.

McCoy, F. 1867. On the discovery of /chtyosaurus and Plesiosaurus in Australia. Annals and

Magazine of Natural History (3) 19: 355-356.

McLacuian, I. R. & Piererse, E. 1978. Preliminary palynological results: site 361, leg 40,

Deep Sea Drilling Project. Jn: Botui, H. M., er AL. eds. Initial Reports of the Deep Sea

Drilling Project 40: 857-882.

McNamara, K. J. 1978. Myloceras (Ammonoidea) from the Albian of central Queensland.

Alcheringa 2: 231-242.

McNamara, K. J. 1980. Heteromorph ammonites from the Albian of South Australia. Trans-

actions of the Royal Society of South Australia 104 (5/6): 145-159.

MEDINA, F. & RINALDI, C. A. 1986. Ammonoideos del Albiano Superior en la Formacion Rio

Mayer de la Estancia La Vega, Provincia de Santa Cruz. Instituto Antartica Argentina.

Contribucion cientifica 324: 1-34, pls 1-5.

MELGUEN, M. 1978. 29. Facies evolution, carbonate dissolution cycles in sediments from the

eastern South Atlantic (DSDP Leg 40) since the Early Cretaceous. Jn: Boxu, H. M., ET

AL. Initial Reports of the Deep Sea Drilling Project 40: 981-1024.

MELGUEN, M. M., PicHon, X. LE. & SipueT, J. C. 1979. Paleonenvironnement de I’atlantique

sud. Bulletin de la Société Géologique de France 20: 471-489.

NATLAND, J. H. 1978. Deposition, provenance, and diagenesis of Cretaceous clastic sediments

drilled on the Atlantic continental rise off Southern Africa, DSDP site 361—Implications

for the early circulation of the South Atlantic. Jn: BoLut, H. M., ET AL. Initial Reports on

the Deep Sea Drilling Project 40: 1025-1062.

mero. F. E., Proserpio, C. A. & NuLLo, G. B. DE. 1981. El Cretacico de la Cuenca

Austral. Comité Sudamericano del Juradsico y Cretacico: Cuencas sedimentarias del Jura-

sico y Cretacico de América del Sur 1: 181-220.

Owen, H. G. 1971. The stratigraphy of the Gault in the Thames Estuary and its bearing on

the Mesozoic history of the area. Proceedings of the Geologists’ Association 82: 187-207.

PicteT, F.-J. 1847. Description des mollusques fossiles qui se trouvent dans les gres verts des

environs de Genéve. Mémoires de la Société de physique et @histoire naturelle de Genéve

11 (2): 257-412, pls 1-15.

Race, J. C. 1981. Les continents péri-atlantiques au crétacé supérieur: migrations des faunes

continentales et problémes paléogéographiques. Cretaceous Research 2 (1): 65-84.

Ranp, H. M. & Masesoong, I. M. 1983. Northeast Brazil and the final separation of South

America and Africa. Palaeogeography Palaeoclimatology Palaeoecology 38 (3-4):

163-183.

RENZ, O. 1981. Venezuela. In: REYMENT, R. A. & BENGTSON, P. Aspects of Mid-Cretaceous

Regional Geology: 197-220. London, New York, Toronto, Sydney, San Francisco:

Academic Press.

RENZ, O. 1982. The Cretaceous ammonites of Venezuela. Basel, Boston, Stuttgart: Birk-

hauser Verlag.

REYMENT, R. A. 1955. The Cretaceous Ammonoidea of southern Nigeria and the southern

Cameroons. Bulletin. Geological Survey of Nigeria 25: 1-112, pls 1-25.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

ReYMENT, R. A. 1956. On the stratigraphy and palaeontology of the Cretaceous of Nigeria

and the Cameroons, British West Africa. Geologiska foreningens i Stockholm férhand-

lingar 78 (1): 17-96.

ReEYMENT, R. A. 1964a. Albian ammonites from Fossil Creek, Oodnadatta, South Australia.

Transactions of the Royal Society of South Australia 88: 21-36, pls 1-S.

REYMENT, R. A. 19646. Coiling and form in South Australian Labeceratidae (Albian; Creta-

ceous). Transactions of the Royal Society of South Australia 88: 37—40, pls 1-2.

REYMENT, R. A. 1978. The mid-Cretaceous of the Nigerian coastal basin. Annales du

Muséum d’histoire naturelle de Nice 4 (1976): XX.1-—XX.15.

REYMENT, R. A. 1980. Paleo-oceanology and paleobiogeography of the Cretaceous South

Atlantic Ocean. Oceanologica Acta 3 (1): 127-133.

REYMENT, R. A. 1981. West Africa. Jn: REYMENT, R. A. & BENGTSON, P. eds. Aspects of

Mid-Cretaceous Regional Geology: 133-160. London, New York, Toronto, Sydney, San

Francisco: Academic Press.

REYMENT, R. A. & Tait, E. A. 1972. Biostratigraphical dating of the early history of the

South Atlantic Ocean. Philosophical Transactions of the Royal Society of London (B. Bio-

logical Sciences) 264 (858): 55-95.

Riccarpi, A. C., AGUIRRE URRETA, M. B. & MeEpINnA, F. A. 1987. Aconeceratidae (Ammoni-

tina) from the Hauterivian—Albian of southern Patagonia. Palaeontographica. ag zur

naturgeschichte der Vorzeit (A) 196 (4-6): 105-185.

Riccarpi, A. C. & Ro.ieri, E. O. 1980. Cordillera Patagonica Austral. /n: Geologia

Regional Argentino: 1173-1306.

Sastry, M. V. A., Rao, B. R. J. & Mamaain, V. D. 1968. Biostratigraphic zonation of the

Upper Cretaceous Formations of Trichinopoly district, South India. Memoirs of the

Geological Society of India 2: 10-17.

SCHEIBNEROVA, V. 1978. 17. Aptian and Albian benthic foraminifers of leg 40, sites 363 and

364, Southern Atlantic. Jn: Botu, H. M., et AL. Initial Reports of the Deep Sea Drilling

Project 40: 741.

SCHEIBNEROVA, V. 1981. Palaeogeographical implications of Cretaceous benthic Foraminifera

recovered by the Deep Sea Drilling Project in the Western South Atlantic Ocean. Creta-

ceous Research 2 (1): 1-18.

SCHINDEWOLF, O. H. 1961. Studien zur Stammesgeschichte der Ammoniten. I. Abhand-

lungen. Mathematisch-Naturwissenschaftlichen Klasse. Akademie der Wissenschaften und

der Literatur, Mainz 1960 (10): 3-109, pls 1-2.

ScLATER, J. G., HELLIMGER, S. & Tapscott, C. 1977. The palaeobathymetry of the Atlantic

Ocean from the Jurassic to the Present. Journal of Geology 85 (5): 509-552.

SIESSER, W. W. 1978. 40. Results in relation to continental shelf and onshore geology. Jn:

Bou, H. M. Et AL. Initial Report Deep Sea Drilling Project 40: 965-979.

SmitH, A. G. & BriDEN, J. C. 1973. Mesozoic and Cenozoic paleocontinental-maps. Cam-

bridge, London, New York, Melbourne: Cambridge University Press.

SpATH, L. F. 1922. On Cretaceous Ammonoidea from Angola, collected by Professor J. W.

Gregory, D.Sc., F.R.S. Transactions of the Royal Society of Edinburgh 53 (1): 91-160.

SpATH, L. F. 1925. On Upper Albian Ammonoidea from Portuguese East Africa, with an

appendix on Upper Cretaceous ammonites from Maputoland. Annals of the Transvaal

Museum 11 (3): 179-200, pls 28-37.

SpATH, L. F. 1930. The fossil fauna of the Samana Range and some neighbouring areas.

Part V. The Lower Cretaceous Ammonoidea; with notes on Albian Cephalopoda from

Hazara. Memoirs of the Geological Survey of India. Palaeontologia indica (n.s.) 15: 51-66.

SpaTH, L. F. 1939. Ammonoidea of the Gault. 13. Palaeontographical Society (Monographs)

92: 541-608, pls 59-64.

STOLICZKA, F. 1863-1866. The fossil Cephalopoda of the Cretaceous rocks of Southern India.

Memoirs of the Geological Survey of India. Palaeontologica indica 1 (1863): 41-56, pls

26-31; 3-5 (1864): 57-106, pls 32-54; 6-9 (1865): 107-154, pls 55-80; 10-13 (1866):

155-216, pls 81-94.

TAMBAREAU, Y. 1982. Les ostracodes et l’histoire géologique de |’Atlantique sud au Crétace.

Bulletin des Centres Rechérches Exploration-Production. Elf-Aquitaine 6 (1): 1-37.

TAVvANI, G. 1942. Paleontologia della Somalia. III. Fossili del Cretaceo. 6. Molluschi del

Cretaceo della Somalia. Palaeontographia italica 32 (suppl. 4): 7-47 (93-133).

THE AMMONITE SUBFAMILY LABECERATINAE 219

Tuomson, M. R. A. 1984. Preliminary ammonite zonation of the mid-Cretaceous rocks of

James Ross Island. Bulletin. British Antarctic Survey 64: 85-91.

Van ANDEL, J. H., THIEDE, J., SCLATER, J. G. & Hay, W. W. 1977. Depositional history of

the South Atlantic Ocean during the last 125 million years. Journal of Geology 85 (6):

651-698.

VENZO, S. 1936. Cefalopodi del Cretaceo medio-superiore dello Zululand. Palaeontographia

italica 36: 1-75 (59-133).

WELLMAN, M. W. 1959. Divisions of the New Zealand Cretaceous. Transactions of the Royal

Society of New Zealand 87 (1-2): 99-163, pls 10-12.

WHITEHOUSE, F. W. 1926. The Cretaceous Ammonoidea of eastern Australia. Memoirs of the

Queensland Museum 8: 195-242, pls 34-41.

WIEDMANN, J. 1962. Unterkreide-Ammoniten von Mallorca. 1. Lieferung: Lytoceratina,

Aptychi. Abhandlungen. Mathematisch-Naturwissenschaftlichen Klasse. Akademie der

Wissenschaften und der Literatur, Mainz, 1962 1: 3-148, pls 1-10.

WIEDMANN, J. 1965. Origin, limits and systematic position of Scaphites. Palaeontology 8:

397-453, pls 53-60.

WIEDMANN, J. & DieENnI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeonto-

graphia italica 64: 1-171, pls 1-18.

WricuT, C. W. 1953. Notes on Cretaceous ammonites. I. Schaphitidae. Annals and Maga-

zine of Natural History (12) 6 (66): 473-476.

WricuT, C. W. 1957. [Cretaceous Ammonoidea.] Jn: Moore, R. C. ed. Treatise on Inverte-

brate Paleontology, Part L, Mollusca 4. Cephalopoda, Ammonoidea: L1-L490. Boulder,

Lawrence: Geological Society of America & University Kansas Press.

* Note added at proof stage (see p. 204).

New material has been described from Patagonia (Aguirre Urreta, M. B. &

Riccardi, A. C. 1988. Albian heteromorph ammonoids from southern Pata-

gonia, Argentina. Journal of Paleontology 62 (4): 598-164). This includes

Labeceras crassetuberculatum Klinger magnum ssp. nov., L. singulare (Leanza,

1970) and Myloceras (Calliscaphites) andinus Leanza. Calliscaphites is retained

as a subgenus of Myloceras on account of its alleged crioceratitid coiling.

Judging by the compressed whorl section and regular ornament of the type and

only species, M. (C.) andinus, it appears to be close to the group of M. besairiei

and ‘Ellipsoceras’ expansum. On the whole, description of this fauna does not

change any of the opinions expressed in this article.

6. SYSTEMATIC papers must conform to the International 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

C.8. . 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 ecdacs 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.

HERBERT CHRISTIAN KLINGER

THE AMMONITE SUBFAMILY

LABECERATINAE SPATH, 1925:

SYSTEMATICS, PHYLOGENY, DIMORPHISM

AND DISTRIBUTION

(WITH A DESCRIPTION OF A NEW SPECIES) —

UME 98 PART 8 FEBRUARY 1989 ISSN 0303-2515

LIBRARY

MAy 04

1989

BYARD

ANNALS

E SOUTH AFRICAN

F ice MUSEUM

UNIV.

CAPE 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

Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Jntroduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

bered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and ~

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

into consideration.

All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 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.:

‘Smith (1969) describes .. .’

‘Smith (1969: 36, fig. 16) describes .. .’

‘As described (Smith 1969a, 1969b; Jones 1971)’

‘As described (Haughton & Broom 1927)...’

‘As described (Haughton et al. 1927)...”

Note: no comma separating name and year

Pagination indicated by colon, not p.

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, 1969b) 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 (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number in parentheses, pagination (first and

last pages of article).

Examples (note capitalization and punctuation)

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

FiscHER, P. H: 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHer, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

TuIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische und anthro-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgefiihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

February 1989 Februarie

Part 8 Deel

LANTERNFISHES OF THE SOUTHERN

BENGUELA REGION. PART 2.

GYMNOSCOPELUS (GYMNOSCOPELUS)

BOLINI ANDRIASHEV IN SOUTH AFRICAN

WATERS, WITH COMMENTS ON THE

DISTRIBUTION OF SUBANTARCTIC

MYCTOPHIDS IN THE EASTERN SOUTH

ATLANTIC

P. ALEXANDER HULLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

1, 2(13; 528) SGD, 4508, Cpa Se 5

6(1, t--p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3),

11(1-2, 5, 7, t.-p.i.), 14(1-2), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33, 36(2), 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 100 8

Printed in South Africa by In Suid-Afrika gedruk deur

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

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

—

LANTERNFISHES OF THE SOUTHERN BENGUELA REGION.

PART 2. GYMNOSCOPELUS (GYMNOSCOPELUS) BOLINI

ANDRIASHEV IN SOUTH AFRICAN WATERS, WITH COMMENTS ON

THE DISTRIBUTION OF SUBANTARCTIC MYCTOPHIDS IN THE

EASTERN SOUTH ATLANTIC

P. ALEXANDER HULLEY

Department of Marine Biology, South African Museum, Cape Town

(With 10 figures and 3 tables)

[MS accepted 17 August 1988]

ABSTRACT

Three large fish specimens, taken with a 180-foot bottom trawl in 780-825 m, west of Cape

Point, South Africa, are identified as Gymnoscopelus (Gymnoscopelus) bolini Andriashev,

1962, despite the fact that they are severely damaged and totally lack body photophores. They

represent not only the most northern record of this Subantarctic species, but also the first

record of the species in southern African waters. The distribution of Subantarctic lanternfishes

in the eastern South Atlantic is discussed. The northern boundary zone for mesopelagic semi-

Subantarctic and bathypelagic Subantarctic species at about 18°S matches the limits for tropical

(holotropical), broadly tropical (thermophilic eurytropical and extended Agulhas), and subtrop-

ical (bisubtropical and south subtropical) species. The boundary limit for mesopelagic

holosubantarctic species lies farther to the south, at about 30°S, and appears to be associated

with the northern limit of relatively fresh Antarctic Intermediate Water in this region. Broadly

tropical (thermophobic eurytropical) species also demonstrate a southern distributional limit at

this latitude in the eastern South Atlantic.

CONTENTS

PAGE

aU URI PRMTIT Fg cna aloha saveeasvece 221

Peet aAPCMSL OOS. oo dat. eee cc ch ccc wa civecasscne 222

eee ram Ol Mateiial 20, ee ee eee dee ees 223

EL DLL a hg a 226

POEMMOICOCCINCINES Cys ec ee bee eee ewes eee eee ees 237

ee dda dele ewes we ned 237

INTRODUCTION

Biannual, demersal fish-sampling cruises over the South African west coast

shelf region (Orange River to Cape Agulhas) have been undertaken on a

routine basis by the Sea Fisheries Research Institute (SFRI), Cape Town, since

January 1983 (Payne et al. 1987). The lanternfishes (Myctophidae) caught during

these cruises have not only provided specimens for taxonomic investigation, but

have also yielded data for distributional analyses of species in the family (Hulley

1986a, 1986b). Biological hypotheses for the distribution of the pseudoceanic

species, Lampanyctodes hectoris, which has accounted for up to 42,4 per cent

pea

Ann. S. Afr. Mus. 98 (8), 1989: 221-240, 10 figs, 3 tables.

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

(451 000 metric tons) of the total annual South African pelagic catch, have been

advanced recently by Hulley & Prosch (1987).

During the January 1987 and July 1987 sampling cruises of R.V. Africana,

several stations were occupied with a 180-foot bottom trawl at depths greater

than the usual 500-m isobath limit. Amongst the material from one of these

deeper stations were three large, but very badly skinned, myctophid specimens.

Since they were totally devoid of body photophores and since myctophid taxo-

nomy is based mainly on photophore grouping and orientation, the identification

of the specimens offered a challenge to an otherwise ‘Gestalt’ process. The identi-

fication, coupled mainly with additional data from the 1986 and 1987 SFRI

demersal cruises, calls for comment on the oceanic zonal patterning of meso-

pelagic fishes in the eastern South Atlantic, and the penetration of Subantarctic

and Antarctic faunal elements into the region.

MATERIAL AND METHODS

In addition to the three specimens from the eastern South Atlantic, meas-

urements were taken on 27 specimens of Gymnoscopelus bolini (SAM-30694,

30695, 30696, 30697, 30698, 30699, 30700, 30701) and 20 specimens of Gymno-

scopelus (Gymnoscopelus) braueri (SAM-30702, 30704, 30705, 30706, 30707)

from FRV Walther Herwig stations in the western South Atlantic. Meristics for

specimens of the two species from the same region were extracted from labora-

tory work-sheets. Distributional records for G. bolini have been taken from

both published and unpublished sources.

When possible, all counts and measurements were made on the left side.

Gill-raker counts (GR) for the first arch do not include stubs. The ultimate ray

of both the dorsal and anal fin is double, but has been counted as a single

element. Measurements, which were taken to the nearest 0,1 mm with needle-

point sliding calipers, included: standard length (SL); head length (HL)—tip of

snout to extreme posterior margin of operculum; length of upper jaw (JL)—

anterior tip of premaxilla to its posterior end; diameter of eye (ED)— horizontal

distance between opposite margins of the socket; depth of head (HD)—-vertical

through the posterior end of the premaxilla; depth of body (BD)—vertical

through the upper base of the pectoral fin; depth of caudal peduncle

(CPD)—least vertical depth; predorsal length (PreD)—distance between tip of

snout and vertical through base of first dorsal ray; prepectoral length (PreP)—

distance between tip of snout and vertical through upper base of pectoral fin;

preventral length (PreV)—distance between tip of snout and vertical through

base of outermost ray of ventral fin; preanal length (PreA)—distance between

tip of snout and vertical through base of first anal ray; pre-adipose length

(PreAd)—distance between tip of snout and vertical through origin of adipose

fin; and length of caudal peduncle (CPL)—distance between vertical through

base of last anal fin ray and end of lateral line.

Photophore groupings and terminology are in accordance with Paxton

(1972).

SOUTHERN BENGUELA LANTERNFISHES 223

DESCRIPTION OF MATERIAL

Gymnoscopelus (Gymnoscopelus) bolini Andriashev, 1962

Figs 1-10

Lampanyctus nicholsi: Norman, 1930: 326 (partim).

Gymnoscopelus (Gymnoscopelus) bolini Andriashev, 1962: 272, fig. 34 (holotype ZIANL

36383; 53°01’S 109°30’W). Parin et al., 1973: 118, fig. 20; 1974: 109, fig. 11. Krefft, 1974:

232. Wisner, 1976: 210, fig. 197. Post, 1979: 19. Hulley, 1981: 250, fig. 120; 1986c: 299,

fig. 86.51. McGinnis, 1982: 61, fig. 50. Bekker, 1983: 220, fig. 90; 1985: 161. Bekker &

Evseenko, 1987: 16.

Material

Three immature female specimens, SAM-—31035 (SL 197,7 mm; 220,7 mm;

249,8 mm) trawled west of Cape Point, South Africa; R.V. Africana (SFRI

station A5280 050 E16; 34°21,8’S 17°38,4’E; 18 January 1987; 19h30-20h30

(local); BT-180’; 780-825 m; bottom temperature not measured).

Description

Measurements expressed as a percentage of SL are given in Table 1.

D 21-22; A 20-22; P 12-13; V8; GR 6+ 1+ 15, total 22; AO and Prec

photophores missing.

“TABLE 1

Measurements expressed as a percentage of the standard length

(SL) for Gymnoscopelus (Gymnoscopelus) bolini (SAM-—31035)

from the eastern South Atlantic.

SL 197,7 mm 220,7 mm 249.8 mm

HL 25,9 24,3 255

IL 19,8 19,1 19,7

ED 5,2 5,6 5,1

HD 18,4 17,2 i

BD 17,5 18,7 1.1

CPD 10,1 9,6 —

CPL 20,1 18,6 20,6

PreD 36,2 39,3 38,8

PreP 24,4 255 25,8

PreV 40,2 43,8 40,9

PreA 573 59,4 58,6

PreAd 78,4 79,1 70,4

Body fusiform and somewhat laterally compressed, length of head 3,9-4,1

in SL. Mouth terminal, slightly oblique and extending well behind vertical

through posterior margin of orbit; length of upper jaw 5,1—-5,2 in SL and 1,3 in

head length; anterior dentigerous region of premaxilla not expanded; maxilla

only slightly expanded posteriorly. Teeth on premaxilla and dentary small and

conical, none enlarged; palatines with series of minute teeth along anterior two-

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

(ey YL A KE eta eeee eaten 6%

Fig. 1. Gymnoscopelus (Gymnoscopelus) bolini (SAM-31035). Lateral view of specimens.

SL 220.7 mm; SL 197,7 mm; 249,8 mm (top to bottom). Scale = 40 mm.

SOUTHERN BENGUELA LANTERNFISHES Dey

Fig. 2. Gymnoscopelus (Gymnoscopelus) bolini (SAM-31035). Lateral view of head.

SL 220,7 mm. Scale = 10 mm.

thirds, some teeth at antero-dorsal margin enlarged and hooked; mesoptery-

goids with small teeth. Eye large, its diameter 4,3—-5,1 in length of head, situated

normally so that vertical distance between upper lip and lower margin of eye at

middle of orbit, about equal in length to pectoral base. Postero-dorsal margin of

operculum evenly concave. Origin of dorsal fin in front of vertical through outer

base of ventral fin. Pectorals damaged, but probably short and reaching only

about one-half to two-thirds the distance to ventral origin. Ventrals reaching to

anus. Anal origin under posterior third of base of dorsal fin; length of base of

anal fin about equal to dorsal base. Origin of adipose fin on or only slightly in

advance of vertical through base of last anal ray. Length of caudal peduncle

about equal to or only slightly longer than length of upper jaw. Seven dorsal and

9-10 ventral procurrent caudal rays, well developed, spine-like and posteriorly

curved.

Antorbital luminous tissue well developed at antero-ventral margin of orbit,

extending dorsally between nasal capsule and eye to about level of dorsal margin

of capsule, and surrounded by black-pigmented skin. Three Br photophores on

each side, under lower jaw. Op: small, opposite posterior end of upper jaw; Op2

large, above and behind Op: at overlap of dorsal and ventral preopercular

flanges. Body photophores missing, specimens very badly skinned.

Colour. Flesh pale, but head with torn patches of dark-brown skin; iris

chocolate-brown; fin rays pigmented; mouth and gill cavities dark.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

DISCUSSION

The body form and fin ray distributions, the presence of an adipose dorsal

fin, three Br photophores under the lower jaw, and Op; and Op: photophores

confirm the family identity of the specimens. Further, the length of the upper

jaw and the structure of the dorsal and ventral procurrent rays suggest inclusion

within the subfamily Lampanyctinae. Within the subfamily, only the genera Noto-

scopelus, Scopelopsis and Gymnoscopelus (Gymnoscopelus) have 21 or more

dorsal fin rays, and only Notoscopelus and Gymnoscopelus (Gymnoscopelus)

have 20 or more anal fin rays (Paxton 1972, table 8; Hulley 1981). In both

Notoscopelus and Scopelopsis the origin of the anal fin is on a vertical through

the middle of the dorsal fin base, whereas in Gymnoscopelus the origin is on a

vertical through the posterior third of the dorsal fin base. This feature is evident

even in larval forms of these genera (Moser & Ahlstrom 1972, figs. 12A, D; 1974,

fig. 12; Moser et al. 1984, figs 120F, 121B, D). Further, whereas the Op: is nor-

mally positioned in Scopelopsis, the Op2 is absent from its position under the

overlap region between the dorsal and ventral preopercular flanges. It may be

represented by a photophore that is reduced in size to that of a ‘secondary’

photophore and is positioned postero-ventrally to the overlap region (SAM-—

28664—4 specimens; SAM -28657—1 specimen). The diagnostic character

given by Paxton (1972), namely the anterior part of premaxillary with (Noto-

scopelus) or without (Gymnoscopelus) expanded dentigerous area, is open

to question. Examination of specimens of Notoscopelus has revealed that the

dentigerous area is expanded in N. (Notoscopelus) caudispinosus (SAM-29245

—8 specimens), somewhat expanded in N. (Notoscopelus) kroeyerii (SAM-

29529—5 specimens) and N. (Pareiophus) bolini (SAM-—29497—2 specimens),

and without an anterior expansion in N. (Notoscopelus) resplendens (SAM-—

30824—4 specimens). Finally, the size of the three large specimens (198-

250 mm) precludes their identification in either Scopelopsis or Notoscopelus;

S. multipunctatus attains a maximum length of 81 mm, N. caudispinosus a

length of 140 mm, N. kroeyerii 143 mm, N. resplendens 95 mm, and N. bolini

102 mm, in the Atlantic Ocean (Hulley 1981).

The genus Gymnoscopelus comprises two subgenera, differentiated by the

position of PVO2 below (subgenus Gymnoscopelus—4 species) or above (sub-

genus Nasolychnus—4 species) the level of the upper base of the pectoral fin.

Unfortunately, this photophore is missing in the three specimens (SAM-31035).

However, only two species in the genus match the specimens in gill-raker count,

namely G. (G.) bolini with GR 6 (rarely 7)+1+15 (14-16, rarely 13); and

G. (G.) braueri with GR 6 (7) +1+4+ 15-16 (17, rarely 14 or 18). Distributions

for these counts are given in Table 2. GR: and anal fin ray counts partially

overlap in these two species and are therefore not entirely suitable as diagnostics

for the separation of these two species. However, the position of the origin of

the dorsal fin in relation to the origin of the ventral fin is diagnostic. In G. bolini

the dorsal origin is anterior to the vertical through the outer base of the ventral

SOUTHERN BENGUELA LANTERNFISHES 227

TABLE 2

Dorsal, anal and gill-raker counts for Gymnoscopelus (Gymnoscopelus) bolini and Gymno-

scopelus (Gymnoscopelus) braueri from the western South Atlantic, and Gymnoscopelus

(Gymnoscopelus) bolini (SAM-31035) from the eastern South Atlantic.

Number of specimens

Count See cedar ta eh to Ay ES 19) 2021 22

G. bolini & it 1-24 8

Dorsal SAM-—31035

G. braueri B28. AOD 25

—

G. bolini 1 9

Anal SAM-31035 1

G. braueri i AT 107 8 1

G. bolini 74 4

GRu SAM-31035 3

G. braueri 211. 31

G. bolini a i453 6

GR, SAM-31035 3

G. braueri Sige io 8 1

fin, whereas in G. braueri the origin is posterior to the vertical through the outer

base of the ventral fin. This is evident from the differences in the slopes of the

regressions of PreD against SL (Fig. 3) and of PreD against PreV (Fig. 4), since

the ventral fin is similarly positioned in the two species (Fig. 5). In addition, the

length of the caudal peduncle (CPL) in G. bolini is shorter than its length in

G. braueri (Fig. 6). Values of the meristics and morphometrics for the three

R.V. Africana specimens therefore confirm their identity as G. bolini. This con-

clusion is supported by the large size of the specimens—G. bolini attains a

maximum length of 280 mm, whereas G. braueri attains only a maximum length

of 132 mm—and by comparison of scatter-plots of their morphometrics with

those of other specimens of G. bolini (Figs 7-9).

The distribution of Gymnoscopelus (Gymnoscopelus) bolini is given in

Figure 10.

The species has been trawled pelagically between the Antarctic Polar Front

(APF) and the Subtropical Convergence (STC), and a Subantarctic distribution

pattern (Parin et al. 1974; Hulley 1981, in press) or west wind drift distribution

pattern (Krefft 1974) has been proposed. Size diminution with decreasing lati-

tude observed by McGinnis (1982) was not evident in the western South Atlantic

(Hulley 1981), where large specimens (greater than 130 mm) have been taken as

far north as the STC, but at depths greater than 600 m. Hulley (1981) therefore

proposed that the distribution of G. bolini could be correlated with the spread

of Antarctic Intermediate Water. He suggested temperature limits between

1,0°C-2,0°C and 5,0°C-5,5°C for adults, and pointed out that juveniles were

taken in water temperatures of 12°C-—13°C.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

+ G bolini

o G braueri

a2 104 156 208 260

Fig. 3. Regression of predorsal length (PreD) against standard length (SL) for

Gymnoscopelus (G.) bolini (+) and Gymnoscopelus (G.) braueri (OQ).

101

+ G bolini

o G. braueri

22 43 64 85 106

Fig. 4. Regression of predorsal length (PreD) against preventral length (PreV) for

Gymnoscopelus (G.) bolini (+) and Gymnoscopelus (G.) braueri (Q).

SOUTHERN BENGUELA LANTERNFISHES 229

+ G bolini

o G. braueri

52 104 156 208 260

Fig. 5. Scatter-plot of preventral length (PreV) against standard length (SL) for

Gymnoscopelus (G.) bolini (+) and Gymnoscopelus (G.) braueri (Q).

+ G bolini

o G. braueri

2V 104 156 208 260

3) i

Fig. 6. Regression of caudal peduncle length (CPL) against standard length (SL) for

Gymnoscopelus (G.) bolini (+) and Gymnoscopelus (G.) braueri (Q).

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

109 146 183 220 257

109 146 183 220 Fre T

B ok

Fig. 7. Scatter-plots of Gymnoscopelus (G.) bolinit. A. Head length (HL) and length of upper

jaw (JL) against standard length (SL). B. Eye diameter (ED) and head depth (HD) against

standard length (SL). Open and solid square symbols refer to SAM-—31035 specimens.

SOUTHERN BENGUELA LANTERNFISHES oat

109 146 183 220 257

A a

101]

o))

PreD, PreP

109 146 183 220 25]

Fig. 8. Scatter-plots of Gymnoscopelus (G.) bolini. A. Body depth (BD) and caudal peduncle

depth (CPD) against standard length (SL). B. Predorsal length (PreD) and prepectoral length

(PreP) against standard length (SL). Open and solid square symbols refer to SAM-31035

specimens.

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

125

PreV, PreA

109 146 183 220 259

a PreAd

4"CRL

159

PreAd, CPL

109 146 183 220 257}

B SL

Fig. 9. Scatter-plots of Gymnoscopelus (G.) bolini. A. Preventral length (PreV) and preanal —

length (PreA) against standard length (SL). B. Pre-adipose length (PreAd) and caudal |

peduncle length (CPL) against standard length (SL). Open and solid square symbols refer to |

SAM-31035 specimens. é

SOUTHERN BENGUELA LANTERNFISHES 233

An adult epibenthic distribution, with probable spawning at depth, has

been suggested by Hulley (1981, in press). In the Atlantic, large specimens of

G. bolini (usually greater than 200 mm) have been reported from benthic sam-

pling along the Argentinian slope north to about 38°S (Hulley 1981). More

recent unpublished data (ISH) extend this benthic range northwards

(WH 703/78: 35°49,2'S 52°49,7'W; 750m; Ty 4,2°C) to approximate more

closely the southern limit of the STC and the position of the pelagic biogeogra-

phic boundary in this region (Boltovskoy 1970, 1986). The species has also been

taken with a beam trawl in the southern Indian Ocean (48°43,5’S 71°06,5'E;

937 m) (Hulley et al. in press).

Therefore, the three R.V. Africana specimens not only represent the most

northerly record for this Subantarctic species and the first record in southern

African waters, but they also mirror the benthic distribution of G. bolini on the

South American slope.

The effect of meridional current flow skews the zonal patterns of the distri-

butions of oceanic fishes, particularly those of mesopelagic species (fide Hulley

1981, fig. 30). Drift-card observations have confirmed the northward flow of

surface water from the STC towards the South African coast in winter (Shannon

et al. 1973), while subsurface investigations have revealed a complex cold-core

eddy system to depths of 500m and more, north of the STC to about 31°S

(Visser 1969; Shannon & Van Rijswijck 1969; Welsh & Visser 1970; Henry

1975; Lenz 1975; Allanson et al. 1981; Lutjeharms 1981; Lutjeharms & Emery

1983; Lutjeharms et al. 1985). Recently, Gordon et al. (1987: 586) described

how South Atlantic Water, a blend of thermocline water and Subantarctic

Surface Water (identical in temperature and salinity to South Atlantic varieties

of Antarctic Intermediate Water—McCartney 1977), may be ‘swept into the

Retroflection region to the east of each of the Agulhas rings’. Pulsed intrusions

of cold Subantarctic Surface Water have been observed crossing the STC into

the southern Benguela Region as an inherent part of the spawning and separation

of an Agulhas ring (Lutjeharms & Meeuwis 1987; Lutjeharms & Van Balle-

gooyen in press, fig. 9). Isotherm and isohaline distributions given by Gorshkov

(1978) suggest that this ‘cold corridor’, linking the offshore Benguela System

with the Subantarctic, is a regular feature of the region. Biologically, this feature

is well exemplified by the distribution of the Subantarctic copepod Metridia

lucens (De Decker 1984, figure on p. 355), where the longitudinal axis of the

penetration of this species into the southern Benguela Region matches the

average longitude (18°25’E) of the cold wedge feature (Lutjeharms & Van Bal-

legooyen in press).

The occurrence of Antarctic and Subantarctic midwater fishes in the eastern

South Atlantic is therefore to be expected and is confirmed by myctophids and

Other species, e.g. Benthalbella macropinna (Scopelarchidae), Bathylagus ant-

arcticus (Bathylagidae), Melanostigma gelatinosum (Zoarcidae), Scopelosaurus

hamiltoni (Notosudidae), Borostomias antarcticus (Astronesthidae), Melanonus

gracilis (Melanonidae), Woodsia meyerwaardeni (Photichthyidae), Ceratias

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

tentaculatus (Ceratiidae), Paradiplospinus gracilis (Gempylidae) and Sio norden-

skjoeldi (Melamphaidae) (Hulley 1972; Johnson 1974; Bertelsen et al. 1976;

Krefft 1978; Anderson 1986; Cohen 1986; Gibbs 1986).

For the Myctophidae, Hulley (19865) has pointed out that holosubantarctic

and semi-Subantarctic species make up about 16 per cent by number of the

oceanic, mesopelagic myctophid fauna in the southern Benguela Region.

Although certain Subantarctic Pattern species (Protomyctophum choriodon,

P. gemmatum, Electrona subaspera, Gymnoscopelus fraseri, G. hintonoides,

G. microlampas), Broadly Antarctic Pattern species (Krefftichthys anderssoni,

Protomyctophum bolini, P. tenisoni, Gymnoscopelus braueri, G. nicholsi) and

Antarctic Pattern species (Electrona antarctica) have not yet been recorded, an

analysis of nine species representing these pattern types that do occur in the

eastern South Atlantic is presented in Table 3. However, the following should

be noted: firstly, a single specimen of Lampanyctus achirus was recorded from

21°05’S 02°00'W (outside the defined region) during Transect-II of FRV Walther

Herwig (Station WH 443/71); and secondly, Rubiés (1985) recorded Diaphus

hudsoni from 18°01'S and Metelectrona ventralis from 20°50'S off the South

West African—Namibian coast.

From Table 3, and with regard to the above, the northern limit of the

bathypelagic Subantarctic species, Lampanyctus achirus, extends to about 21°S_ |

and at depths below 300 m in the eastern Atlantic. With the exception of Gym-

noscopelus piabilis, this northern limit is matched by mesopelagic semi-

Subantarctic species. These species extend to about 18°S and at depths of

10-1 550 m during pelagic sampling and 229-823 m during benthic sampling. On

the other hand, mesopelagic holosubantarctic species appear to be limited to the

region south of 30°S and at deeper levels, i.e. 300-1 400 m (pelagic sampling)

and 780-825 m (benthic sampling). The concept of these two latitudinal bound-

ary zones is reinforced by the observations of Olivar (1987), who pointed out

that Diaphus hudsoni larvae were most abundant in two particular areas, one

around 20°S and the other between 29°S and 30°S, over bottom depths of

400-600 m. Hulley (1981: 284) pointed out that G. piabilis represents an ‘anom-

alous case of a semi-subantarctic species’, whose distribution is similar to that of

a holosubantarctic species. The results expressed in Table 3 confirm this obser-

vation. It should be noted that a Holosubantarctic Subpattern for the species

has been suggested by both McGinnis (1982)(= Pattern III) and Bekker &

Evseenko (1987)(= Notal).

Hulley (1981) has already commented on the distribution of Lampanyctus

achirus and on the ‘18°S’ boundary zone, the region where the Benguela

Current turns to flow westwards (Moroshkin et al. 1970). This zone, which may

vary seasonally in position and may be farther to the south closer inshore,

apparently limits the distribution of broadly tropical species (Thermophilic

eurytropical and Extended Agulhas subpatterns), tropical species (Holotropical

Subpattern) and subtropical species (Bisubtropical and South subtropical sub-

patterns). On the other hand, broadly tropical (Thermophobic eurytropical)

ZOD

SOUTHERN BENGUELA LANTERNFISHES

"WOOT 38 Soinjeroduia} SuNTuUN]— | ‘UW QOT Je sommjesroduis} Suryunty— » ‘UOISSNISIP ISYJINJ IOJ 1X9} 9os—,

€78-6C7 ASOcEL S,Z0c€Z +L79 Z ep o1yyueq

000 I-01 AZ1opl S,6161 I8€ 97 orsejod *CI sdooq snaoydojoquxg

STP—-O0E ASZ9L S,SToZE € Z oryyuaq

00L-O1 WLC: SiS leSe 96 4 orsejad SI ¢ SIDAJUAA DUOAIIAIAIAW

S6b—-09r AOSobT S,OT0€ € I o1yyueq

Oss I-CIT A,SE9T S,ZobE 9 € o1sejad a € siiquid snjadossoumdy

69L-STPr ASEEL S,9E9Z +71 6 oryyuaq

009-SZ «A LvoOL S,0781 8L9 18 o1sejod 4ST ¢ 1uospny snydviq

(onjorejURQnsS-IWas) sTjOIeJURGNS

00L-009 A,000ST S,00.8€ (6 ‘6 o1sejod 8 Z wunjayosvd wnydojrXKuojodg

OOF I A,€0.01 S,0€.67 I I o1sejad $e Z 1aayspipun wnydojaKuojodg

S78-O8L ABEoLT Si7ZZebE € I o1yuaq c‘¢ I 1u1j0qg snjadoossouudy

SOE A€pobl S,9ZopE€ Z I oisejad S 1849GS]AD9 DUOAIIALT

(onorejuRQnsojo}) 90I1eJURQnS

OIDV TadOSAWN

000 I-OZI xa,cCeS0 S,PIoLZ ae SI o1sejod 2) snaryoo snjakuoduvT

ororeyueqns

OIOV TAdAHLVA

(ur) fe 5) (93)

yidap uorjisod suowutoads suone}s odA} toddn IOMO] soroods

SuIYsIy A]JOYIIOU SOP] JO ‘ON JO ‘ON IeayH ‘dw ] /sunmry

‘(T861) AoyN Je somnjesrsdurs}

SUI}IW] pue dINJepOUSWOU Ud}}ed UOTNQIISIG “W 6ZZ ULY) 19]e919 suOT}E}s [Je 1e syJdap Wo}0g “(9g6[ UOSIoydoeP]) (BUOTEDIeg) Ie] [EP SPIOUDID op

ONIsUT pur ‘(T—6T AdT[NH) (Sinquieyy) rorsyosijaa¢g Inj inNsuy ‘(UMOT, oded) an sUT YoIeasoy satsoysiy eas ‘(UMOT oded) WNesnp] URDTTY YINOS

Aq sasinio Surjdwies sutmnp (4.07—.00 SoSh—o81) SULT} W YINOS UIO}sed 9Y} UI UDye} seprydoyoAP Jo sotoads onNoejURQns s13ejodAyjeq pue o1sejadosay

¢ aTaVL

ANNALS OF THE SOUTH AFRICAN MUSEUM

236

‘eyep poyst{qndun pur paystyqnd uo paseq iuijog (snjadossouwdy) snjadoosouwdy Jo uonnquisiq ‘OL ‘314

ool Ovi oz oor 09 | M oz

OOoL O@L

OvL

SOUTHERN BENGUELA LANTERNFISHES 257

species extend to about 30°S. Despite differences in the distributional limiting

temperatures (Table 3), the apparent distributional limit between about 30°S

and 38°S is difficult to reconcile with physical hydrographic conditions in the

region. However, Shannon & Hunter (in press) recently pointed out that rela-

tively fresh Antarctic Intermediate Water is present off the south-western Cape

between Cape Point (34°21’S) and Cape Columbine (32°50’S), and Chapman &

Shannon (1985) discussed ‘virgin’ Central Water upwelling in the region. It is

interesting to note that the proposed temperature limits, i.e. 1°C-—8°C for holo-

subantarctic species, and 5°C-15°C for semi-Subantarctic species (excluding

G. piabilis) (Table 3), approximate temperature values in the T-S curve for

Antarctic Intermediate Water (4°-6°C, <34,7 x 10-3) and South Atlantic

Central Water (6°C, 34,5 x 10° to 16°C, 35,5 x 10-3), respectively (Shannon

1985; Shannon & Hunter in press). According to the latter authors, statistically

significant seasonal differences in the occurrence of Antarctic Intermediate

Water were not apparent in the region west of 20°E, although water of lower

salinity may be present at 30°-32°S during the summer. Existing data on mycto-

phids preclude elucidation of seasonal trends, since regular sampling was

restricted to a few months only. Bathypelagic Subantarctic species were taken in

March, April and August; mesopelagic holosubantarctic species in January,

March and August; and mesopelagic semi-Subantarctic species in January, Feb-

ruary, March, April, May, June, July, August, September and November (SAM

data).

ACKNOWLEDGEMENTS

I thank Dr Andy Payne (SFRI) and the Captain and crew of R.V. Africana

for the collection of specimens during the Hake Survey cruises. My thanks are

also due to Drs Gerhard Krefft (Hamburg) and Tomasz Linkowski (Gdynia) for

data on lanternfishes caught during mesopelagic trawling in the western South

Atlantic and during bottom trawling off the Argentinian continental shelf; and

to Drs Johan Lutjeharms, Alfred Post and Vere Shannon for permission to view

and cite the manuscripts of papers in press. Mrs Michelle van der Merwe and Mr

Virgilio Branco assisted with the preparation of the figures. This study was made

possible through a grant from the CSIR Foundation for Research Development

and support from the South African Museum.

REFERENCES

ALLANSON, B. R., Hart, R. C. & LutTseHarmMs, J. R. E. 1981. Observations on the nutrients,

chlorophyll and primary production of the Southern Ocean south of South Africa. South

African Journal of Antarctic Research 10/11: 3-14.

ANDERSON, M. E. 1986. Family 94: Zoarcidae. Jn: SmirH, M. M. & HeEeEmstra, P. C. eds.

Smiths’ Sea Fishes: 342-343. Johannesburg: Macmillan.

ANDRIASHEV, A. P. 1962. Biological results of the Soviet Antarctic Expedition (1955-1958).

I. Bathypelagic fishes of the Antarctic. 1. Family Myctophidae. IJssledovaniya Fauny

Morei 1 (10): 216-294. (In Russian.)

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

BEKKER, V. E. 1983. Myctophids of the world ocean. Moskva: Akademiya nauk SSSR. (In |

Russian. )

BeKKeER, V. E. 1985. Distribution of myctophid fishes and the position of the biogeographical

border between the islands of Saint Paul and Kerguelen. Journal of Ichthyology 25 (2):

159-162.

Bekker, V. E. & EvsEENKO, S. A. 1987. Distribution of mesopelagic fishes and biogeographic

borders in the southern Pacific Ocean in January-February 1985. Journal of Ichthy-

ology 27 (1): 9-20.

BERTELSEN, E., KRerrt, G. & MARSHALL, N. B. 1976. Fishes of the family Notosudidae.

Dana Report 86: 1-114.

Bo.tovskoy, E. 1970. Masas de agua (caracteristica, distribucion, movimientos) en la super-

ficie del Atlantico sudoeste, segun indicadores biologicos—Foraminiferos. Publico

Servicio de Hidrografia Naval, Buenos Aires H643: 1-99.

Bo.tovskoy, D. 1986. Biogeography of the Southwestern Atlantic; overview, current prob-

lems and prospects. fn: PrERROT-BULTS, A. C., VAN DER SPOEL, S., ZAHURANEC, B. J. &

JOHNSON, R. K. eds. Pelagic biogeography. UNESCO Technical Papers in Marine

Science 49: 14-24.

CHAPMAN, P. & SHANNON, L. V. 1985. The Benguela ecosystem. Part II. Chemistry and

related processes. In: BARNES, M. ed. Oceanography and Marine Biology. An a

Review 23: 103-251. Aberdeen: University Press.

CoHEN, D. M. 1986. Family No. 62: Bathylagidae. Jn: SmirH, M. M. & HEEmstrRA, P. C. eds.

Smiths’ Sea Fishes: 216. Johannesburg: Macmillan.

De Decker, A. H. B. 1984. Near-surface copepod distribution in the south-western Indian

and south-eastern Atlantic Ocean. Annals of the South African Museum 93 (5): 303-370.

Gipss, R. H. yr. 1986. Family No. 69: Astronesthidae. Jn: SmirH, M. M. & HEEmstra, P. C.

eds. Smiths’ Sea Fishes: 231-234. Johannesburg: Macmillan.

Gorpbon, A. L., LUTJEHARMS, J. R. E. & GRUNDLINGH, M. L. 1987. Stratification and circula-

tion at the Agulhas Retroflexion. Deep-sea Research 34 (4): 565-599.

GorsHkov, S. G. 1978. World ocean atlas. 2. Atlantic and Indian oceans. Oxford: Pergamon.

Henry, A. E. 1975. Hydrology and nutrient salts of the south-east Atlantic and south-west

Indian oceans in 1968. Jnvestigational Report. Division of Sea Fisheries, Republic of South

Africa 95: 1-66.

Hu .tey, P. A. 1972. A report on the mesopelagic fishes collected during the deep-sea cruises

of R.S. ‘Africana’, 1961-1966. Annals of the South African Museum 6@ (6): 197-236.

Huey, P. A. 1981. Results of the research cruises of FRV ‘Walther Herwig’ to South

America. LVIII. Family Myctophidae (Osteichthyes, Myctophiformes). Archiv ftir Fischerei-

wissenschaft 31 (1): 1-300.

Huttey, P. A. 1986a. A taxonomic review of the lanternfish genus Triphoturus Fraser-

Brunner, 1949 (Myctophidae, Osteichthyes). Annals of the South African Museum 97 (4):

71-95.

Hu .tey, P. A. 19865. Lanternfishes of the southern Benguela Region. Part 1. Faunal com-

plexity and distribution. Annals of the South African Museum 97 (7): 227-249.

Huey, P. A. 1986c. Family No. 86: Myctophidae. Jn: SmirH, M. M. & HEEmstraA, P. C.

eds. Smiths’ Sea Fishes: 282-321. Johannesburg: Macmillan.

Hu ey, P. A. In press. Family Myctophidae. Jn: Gon, O. & HEEmstRA, P. C. eds. Fishes of

the Southern Ocean.

Huey, P. A., Camus, P. & DUHAMEL, G. In press. Ichthyological results of cruise MD-

42/SIBEX-II. Part 1. Fishes from RMT-8 stations, with additional records of lanternfishes

(Myctophidae: Osteichthyes) from the Indian Sector of the Southern Ocean. Cybium.

Hu ey, P. A. & Proscu, R. M. 1987. Mesopelagic fish derivatives in the southern Benguela

Upwelling Region. Jn: Payne, A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Ben-

guela and comparable ecosystems. South African Journal of Marine Science 5: 597-611.

JOHNSON, R. K. 1974. A revision of the alepisauroid family Scopelarchidae (Pisces: Myctophi-

formes). Fieldiana (Zoology) 66: 1-249.

KreFFt, G. 1974. Investigations on midwater fish in the Atlantic Ocean. Berichte der Deut-

schen Wissenschaftlichen Kommission ftir Meeresforschung 23 (3): 226-254.

Krerrt, G. 1978. Distribution patterns of oceanic fishes in the Atlantic Ocean. Revue de

travaux. Institut des péches maritimes 40 (3-4): 439-460.

SOUTHERN BENGUELA LANTERNFISHES 239

Lenz, W. 1975. Untersuchungen zur inneren hydrographischen Struktur des sudlichen und

- mittleren Atlantik (0-2 000 m Tiefe) mit zoogeographischen Anmerkungen. Berichte der

Deutschen Wissenschaftlichen Kommission fiir Meeresforschung 24 (1): 1-22.

LUTJEHARMS, J. R. E. 1981. Features of the southern Agulhas Current circulation from satel-

lite remote sensing. South African Journal of Science 177: 231-236.

LUTJEHARMS, J. R. E. & Emery, W. J. 1983. The detailed thermal structure of the upper

layers between Cape Town and Antarctica during the period Jan.—Feb. 1978. South

African Journal of Antarctic Research 13: 3-14.

LUTJEHARMS, J. R. E. & MEEuwIS, J. M. 1987. The extent and variability of South-east Atlan-

tic upwelling. In: Payne, A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and

comparable ecosystems. South African Journal of Marine Research 5: 51-02.

*LUTJEHARMS, J. R. E. & VAN BALLEGOOYEN, R. C. In press. The retroflection of the Agulhas

Current. Journal of Physical Oceanography.

LUTJEHARMS, J. R. E., WALTERS, N. M. & ALLANSON, B. R. 1985. Oceanic frontal systems

and biological enhancement. Jn: SIEGFRIED, W. R., Conny, P. R. & Laws, R. M. eds.

Antarctic nutrient cycles and food webs: 11-21. Berlin: Springer.

MACPHERSON, E. 1986. Resultados de las expediciones oceanografico-pesqueras ‘Benguela IIT’

(1981) a ‘Benguela VII’ (1984) y ‘Valdivia I’ realizadas en el Atlantico Sudoriental

(Namibia). Datos Informativos. Instituto de Ciencias del Mar, Barcelona 17: 1-344.

McCartney, M. 1977. Subantarctic Mode Water. Jn: ANGEL, M. ed. A voyage of discovery:

George Deacon 70th anniversary volume. Deep-sea Research (Supplement): 103-119.

Oxford: Pergamon.

McGinnis, R. F. 1982. Biogeography of lanternfishes (Myctophidae) south of 30°S. Antarctic

Research Series. National Academy of Sciences, Washington 35: 1-110.

MorosHkin, K. V., Busnov, V. A. & BuLatov, R. P. 1970. Water circulation in the eastern

South Atlantic Ocean. Oceanology 10 (1): 27-37.

Moser, H. G. & AntstroM, E. H. 1972. Development of the lanternfish, Scopelopsis multi-

punctatus Brauer, 1906, with a discussion of its phylogenetic position in the family

Myctophidae and its role in a proposed mechanism for the evolution of photophore pat-

terns in lanternfishes. Fishery Bulletin of the National Oceanic & Atmospheric Admini-

stration 70 (3): 541-564.

Moser, H. G. & Autstrom, E. H. 1974. Role of larval stages in systematic investigations of

marine teleosts: the Myctophidae, a case study. Fishery Bulletin of the National Oceanic &

Atmospheric Administration 72 (2): 391-413.

Moser, H. G., AHLSTRoM, E. H. & Paxton, J. R. 1984. Myctophidae: Development. In:

Moser, H. G., RicHarps, W. J., CoHen, D. M., FaHay, M. P., KENDALL, A. W. yr., &

RICHARDSON, S. L. eds. Ontogeny and systematics of fishes, based on an International Sym-

posium dedicated to the memory of Elbert Halvor Ahlstrom. Special Publication. American

Society of Ichthyologists and Herpetologists 1: 218-239.

Ouivar, P. 1987. Larval development and spawning of Diaphus hudsoni in the Benguela

Current region. Marine Biology 94: 605-611.

ParIN, N. V., ANDRIASHEV, A. P., BoropuLina, O. D. & TcHuvasov, V. M. 1974. Midwater

fishes of the South-western Atlantic Ocean. Trudy Instituta okeanologii, Akademiya nauk

SSSR 98: 76-140. (In Russian.)

ParIN, N. V., BEKKER, V. E., BoRODULINA, O. D. & TcHuvasov, V. M. 1973. Deep-sea

pelagic fishes of the South-eastern Pacific and adjacent waters. Trudy Instituta okeanolo-

gli, Akademiya nauk SSSR 94: 71-172. (In Russian.)

PaxTon, J. R. 1972. Osteology and relationships of the lanternfishes (family Myctophidae).

Bulletin of the Natural History Museum of Los Angeles County, Science 13: 1-81.

PayNnE, A. I. L., Rose, B. & Leste, R. W. 1987. Feeding of hake and a first attempt at deter-

mining their trophic role in the South African west coast marine environment. Jn: PAYNE,

A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and comparable ecosystems.

South African Journal of Marine Science 5: 471-501.

*Post, A. 1979. Artificial key to fishes in the New Zealand region (Southern Island). Depart-

ment of Ichthyology, Institut fiir Seefischerei, Hamburg: 1-131 (mimeograph).

Rusiés, P. 1985. Zoogeography of the lanternfishes (Osteichthyes, Myctophidae) of South-

west Africa. Simposio Internacional sobre las areas de afloramiento mas importantes del

Oeste Africano (Cabo Blanco y Benguela). Instituto de Investigaciones Pesqueras, Barce-

lona 1: 573-586.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

SHANNON, L. V. 1985. The Benguela ecosystem. Part I. Evolution of the Benguela, physical

features and processes. Jn: BARNES, M. ed. Oceanography and Marine Biology. An

Annual Review 23: 105-182. Aberdeen: University Press.

“SHANNON, L. V. & Hunter, D. 1987. The Antarctic Intermediate Water in the South-east

Atlantic and South-west Indian oceans. Handbook and Abstracts of Papers and Posters.

6th National Oceanographic Symposium, South African National Committee for Oceano-

graphic Research. Paper 320.

*SHANNON, L. V. & Hunter, D. In press. Notes on Antarctic Intermediate Water around

southern Africa. South African Journal of Marine Science 6.

SHANNON, L. V., STANDER, G. H. & CAMPBELL, J. A. 1973. Oceanic circulation deduced from

plastic drift cards. Investigational Report. Sea Fisheries Branch, Republic of South

Africa 108: 1-31.

SHANNON, L. V. & VAN Ruswuck, M. 1969. Physical oceanography of the Walvis Ridge

region. Investigational Report. Division of Sea Fisheries, Republic of South Africa 70:

1-19.

VissER, G. A. 1969. Hydrographical observations on the South-east Atlantic Ocean. 1. The

Schmitt-Orr Seamount area. Investigational Report. Division of Sea Fisheries, Republic of

South Africa 177: 1-23.

WELSH, F. G. & Visser, G. A. 1970. Hydrological observations in the South-east Atlantic

Ocean. 2. The Cape Basin. Investigational Report. Division of Sea Fisheries, Republic of

South Africa 83: 1-24. ;

WIsnER, R. L. 1976. The taxonomy and distribution of lanternfishes (family Myctophidae) of

the eastern Pacific Ocean. Navy Ocean Research and Development Activity Report 3:

1-229.

*—_quoted with permission of the authors.

6. SYSTEMATIC papers must conform to the /nternational 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.,

ete.

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 Hee 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.

P. ALEXANDER HULLEY

LANTERNFISHES OF THE SOUTHERN |

BENGUELA REGION. PART 2.

GYMNOSCOPELUS (GYMNOSCOPELUS) —

BOLINI ANDRIASHEV IN SOUTH AFRICAN ~

WATERS, WITH COMMENTS ON —

THE DISTRIBUTION OF SUBANTARCTIC

MYCTOPHIDS IN THE EASTERN |

SOUTH ATLANTIC

| ME 98 PART 9 NOVEMBER 1989 |

: MC Z ISSN 0303-2515

LIBRARY

JAN 23 1990

HARVARD

UNIVERSITY

ANNALS

THE SOUTH AFRICAN

MUSE

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

Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s) .

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

bered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

should be left to editor and publisher.

4, ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

into consideration.

All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The.number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 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.:

‘Smith (1969) describes... .”

‘Smith (1969: 36, fig. 16) describes. . .’

‘As described (Smith 1969a, 1969b; Jones 1971)’

‘As described (Haughton & Broom 1927)...’

‘As described (Haughton et al. 1927)...”

Note: no comma separating name and year

pagination indicated by colon, not p.

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, 1969b) 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 (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number in parentheses, pagination (first and

last pages of article).

Examples (note capitalization and punctuation)

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

FiscHER, P. H: 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHer, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthro-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

November 1989 November

Part 9 Deel

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

THE AMMONITE FAMILY

PLACENTICERATIDAE HYATT, 1900;

WITH COMMENTS ON THE

SYSTEMATIC POSITION OF THE GENUS

HY PENGONOCERAS SPATH, 1924

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

1, 201-3, 5-8), 30D aesee: tp ia oe

6(1, tepid), 7 8s 9, Tay:

11(1-2, 5, 7, t.-p.i.), 14(1-2), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33, 36(2), 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 103 2

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica’ ‘Press, “Pty.,” Ltd... Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH

AFRICA. THE AMMONITE FAMILY PLACENTICERATIDAE HYATT,

1900; WITH COMMENTS ON THE SYSTEMATIC POSITION OF THE

GENUS HYPENGONOCERAS SPATH, 1924

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 127 figures)

[MS accepted 27 May 1987|

ABSTRACT

The systematics of the ammonite family Placenticeratidae are complex. Most of the

confusion surrounding the plethora of names applied to the family has been cleared by Kennedy

& Wright (1983). This confusion was due mainly to failure to recognize intraspecific variation,

dimorphism, differing rates of ontogenetic development, slow phylogenetic change, and possibly

effects of ecophenotypic variation. The Zululand representatives of this family, Placenticeras

kaffrarium Etheridge, 1904, from the Middle Coniacian, and Hoplitoplacenticeras howarthi

Collignon, 1970, in the Upper Campanian, illustrate all these features and permit meaningful

discussion. A list of species that have been referred to the family is given. The appendix contains

original (or translations of) diagnoses of genera placed in the synonymy of the genus

Placenticeras. The systematic position of the genus Hypengonoceras, which in the past had often

been referred to the family Placenticeratidae, is discussed. It is referred to the Engonoceratidae

on account of its sutural development.

CONTENTS

PAGE

ea EEN Meee FG Ue Fes Bice no wes wees Sele donee ae ks 242

Sn SSS Sohn [et SS ea 242

8 RSET G2 Se ee ieee 242

(LS DS CIT) Sites) 012 Cia) 210 An ne ra 243

IIR RHIAN ee ysl vic ais give Ss koa eve ea ee eee wee 243

tse Ie Pee in cisdc. ciabie coed ajais fein os wate elds Oe we Re eG 243

ee ACORN ANN ENOIN oe ea eset Sige olan Wards Pew eae Re eee SH 244

oe HES Pp TEESE Ucalie ois Snnc 246

COLO U IRE NS (Cee 246

Dice Galcs OL CEVCIOPMENL. . 2... nce cade a ese sees Hs 246

ania Omi Lelative PEODOLIODS 15... - 56 ssc 6 le ee ei ne wee 258

PO BeMOMCICNQMGE oe ce een cet g ee ede ne aaa eee sins 258

CP 2 SOUS(E oS Re Oe 259

sutural ontogeny, phylogeny and systematics .................... 259

241

Ann. S. Afr. Mus. 98 (9), 1989: 241-408, 127 figs.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Systematic palacontolOgy: 22 ins wis oss Suneag oaeee ee ee 266

Genus Placenucerds:«.. ick ais oh 2 cNedaas a ee ee 266

Placenticeras kaffraruum Etheridge, 1904... .......0. «0. 0.406 ne 268

Genus Hoplitoplacenticeras®. 3. .-.. 355. overs Sie oan eh ee 355

Hoplitoplacenticeras howarthi Collignon, 1970 ............... 358

The systematic position of Hypeneonoceras: .:. 3. 02.42 aso es eee 361

Speculations on intraspecific variation in Placenticeratidae ............ 383

List of species referred to: Placenticeratudde «5.4... o.. «, 05.40 st aoe 384

Acknowletigements ”.. 2.).2.5.625). a ymin sauna Geen eae ee 391

REREERCES e.5co5 5/5 dace Sido Ne ok Sees le BRR ey ee 392

Appendix—Original diagnoses of genera here placed in the synonymy

OF PIGCEMUECENGS 3 5i5.60.45. coo ioe nia Ee Oe Se 399

INTRODUCTION

In Zululand and Natal, South Africa, the family Placenticeratidae is repre-

sented by two species only, Placenticeras kaffrarium Etheridge, 1904, in the

Middle Coniacian (sensu Klinger & Kennedy 1984), and Hoplitoplacenticeras

howarthi Collignon, 1970, in the Upper Campanian. The large population of

P. kaffrarium permits meaningful discussion on the taxonomy of the family,

which should be seen as supplementary to Kennedy & Wright’s (1983) pioneering

study of Ammonites polyopsis Dujardin.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of the

material studied:

BMNH British Museum (Natural History), London

EMP _ Ecole des Mines Collections, Université Claude-Bernard, Lyons

FSR Faculté des Sciences, Rennes

MNHP Muséum National d’Histoire Naturelle, Paris

NMB_ National Museum (Bloemfontein) (presently in the collections of the

South African Museum, Cape Town)

NMP Natal Museum, Pietermaritzburg

OUM = Oxford University Museum

SAM — South African Museum, Cape Town

SAS Geological Survey of South Africa, Pretoria

SP Collections of the Sorbonne, now in Université Pierre et Marie Curie,

Paris

FIELD LOCALITIES

Details of field localities are given by Kennedy & Klinger (1975); fuller

descriptions of these localities are deposited in the Department of Palaeontology,

British Museum (Natural History), London; Geological Survey of South Africa,

Pretoria, and the South African Museum, Cape Town.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 243

DIMENSIONS OF SPECIMENS

All dimensions are given in millimetres: D = diameter; Wb = whorl breadth;

WH = whorl height; U = umbilical diameter. Ut, Lt and Vt respectively refer to

the number of umbilical, lateral and ventral tubercles. Figures in parentheses are

dimensions as a percentage of the total diameter.

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916) reviewed and discussed by

Kullmann & Wiedmann (1970) is followed here: I = interal lobe, U = umbilical

lobe, L = lateral lobe, E = external lobe. Where necessary, however, Ruzhent-

sev’s (1949, 1957) terminology is also given.

DISCUSSION

Failure to recognize the wide range of intraspecific variation in members of

the ammonite family Placenticeratidae, combined with the effects of dimorphism

and an extremely slow rate of evolutionary change in many lineages, has resulted

in a large number of specific and generic names and concomitant taxonomic

confusion. During the last decade, placenticeratids have been distributed over

two subfamilies and about two dozen genera. Most of this taxonomic confusion

has been cleared by Kennedy & Wright (1983) in their pioneering study of

Placenticeras polyopsis (Dujardin, 1837). The Zululand representatives of this

family, Placenticeras kaffrarium Etheridge, 1904, and Hoplitoplacenticeras

howarthi Collignon, 1970, especially the former, display the above-mentioned

features admirably, and permit detailed description as well as discussion on the

systematics of the family supplementary to the studies of Kennedy & Wright

(1983). Study of this material, as well as a review of the literature, point to several

problem areas in the taxonomy of the family. These include:

1. Separation of Placenticeratidae and Engonoceratidae is by no means as clear as

some authors would suggest. This problem is discussed in a separate chapter

dealing with the taxonomic affinities of the genus Hypengonoceras (see p. 361).

2. The origin of the Placenticeratidae is clearly in the Hoplitidae, but the exact

phylogeny of the group is not yet quite clear.

3. The role of sutural ontogeny in interpreting the phylogeny and its taxonomic

value is disputable.

4. The validity of the numerous generic names applied to the Placenticeratidae

has been dealt with by Kennedy & Wright (1983) and we can merely add some

more detail on the basis of the rich Zululand faunas.

5. The validity of the numerous species referred to the family, 130 of which are

listed in this study. This large number of specific names reflects, we believe, the

wide range of intraspecific variation in this group. It is necessary to try to define

how variable the species are and whether factors can be identified that influence

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

the variation. Without access to the original material no attempt is made at

compiling a full synonomy of all these ‘species’.

Starting at specific level, these problems can be discussed.

INTRASPECIFIC VARIATION

The fact that placenticeratid species are highly variable was already known

almost 150 years ago. Dujardin (1837: 232) in his description of Ammonites

polyopsis already commented ‘Cette ammonite varie tellement, que des échan-

tillons isolés pourraient étre pris pour des espéces distinctes, si l’on n’observait

quelquefois toutes les variations possibles sur les différents points d’un méme

échantillon’.

The most comprehensive discussion on the subject is surely provided by

Hyatt (1903), who, if one reads between the lines, was very close to the truth in

terms of current interpretation of variation in the group. Some of his comments

are well worth quoting:

‘The species of this genus (Placenticeras) could be readily distinguished if it

were not for the great range of form in the gerontic stage, which occurs in dwarfed

as well as in large specimens, and is continually mistaken for the ephebic stage.’

‘The species are all connected so closely by intermediate forms that distinct

lines are difficult to draw between contiguous species . . . are senile forms in the

phylum, or what I have named phylogerontic. They are not scaphitoid. .. .’

‘There is no real line between P. guadalupae, sancarlosense, and planum, nor

between newberryi and guadalupae, nor between guadalupae, sancarlosense,

syrtale, intercalare, and placenta, nor between intercalare, stantoni, pseudo-

placenta, and whitfieldi. As a matter of fact there is no real break, such as is

usually supposed to establish a species, between P. guadalupae and the extreme

form of whitfieldi.’

‘If, however, one admits that all American forms make up only one species,

it becomes illogical to separate the European forms from each other or the

American from them, and, consequently, all the forms of Placenticeras are one

species.’

CU Oa

A |

BONG 0 BME same,

_ Age © PCY Csr

Fig. 1. Comparison of suture lines to illustrate relative size of lateral lobes. A. Gissarites

kysylchense. B. Placenticeras placenta. (After Iljin 1958, fig. 2.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 245

Other comments on the wide intraspecific variation in placenticeratids are

found widely scattered in the literature, e.g. Paulcke (1907), Reeside (1927b),

Howarth (1965), Wolleben (1967), Summesberger (1979), Klinger & Kennedy

(1980b, 1980c), Kennedy & Wright (1983), and Kennedy (1984).

It remains, however, to define this variation, or at least to try to determine

which factors influence the gross morphology of the shell. Studies on the South

African material and others, e.g. Summesberger (1979), Kennedy & Wright

(1983), and Kennedy (1984), show that the following factors are of importance in

interpreting intraspecific variation:

1. Dimorphism.

2. Ontogenetic changes.

3. Differing rates of development giving individuals of the same size but at a

different point in ontogeny, different ornament; a result of size dimorphism and

of variation of adult size in individual dimorphs.

4. Variation in relative proportions and ornament, i.e. intraspecific variation per

se at the same ontogenetic stage.

5. Phylogenetic change.

6. Geographic variation.

2 L 2 U, J

Fig. 2. Sutural ontogeny of Metaplacenticeras pacificum (Smith, 1900). (After Smith 1900,

pls 27-28.)

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

DIMORPHISM, ONTOGENETIC CHANGES AND DIFFERING RATES OF DEVELOPMENT

These three factors are intimately linked and it is impossible to separate them

entirely in this discussion. Ontogenetic changes in placenticeratids are striking. In

most Placenticeras, the following can be observed:

(a) Juvenile stage. This includes the very early stage with the protoconch, nepionic

constriction, and subsequent early whorls (e.g. Hyatt 1903, pl. 43 (fig. 6)). The

protoconch is globular, wider than high. It is succeeded by depressed, reniform,

early whorls (Figs 9-10). Thereafter the whorls soon become compressed.

(b) Early phragmocone stage. Here the whorl section is distinctly compressed,

much higher than wide with maximum breadth at the umbilical shoulder (Fig. 11).

The flanks are smooth, save for fine sinuous, or sickle-shaped striae and growth

lines. The curve of the sickle may be slightly stronger than the haft, producing

feeble crescentic ribs or riblets on the outer half of the flanks (e.g. Hyatt 1903,

d J

(

E L Us U J

Fig. 3. Sutural ontogeny of Metaplacenticeras pacificum (Smith, 1900). (After Smith 1900,

pls 27-28.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 247

pl. 43 (figs 7-8)). The venter is concave to flat and smooth, bordered on either

side by entire, sharp ventrolateral edges. The umbilicus is narrow and funnel-

shaped with a sharp umbilical shoulder.

inflated and rounded, the umbilical shoulder becomes more rounded, and

umbilical tubercles may be connected by single or bifurcating ribs of varying

strength to weak or prominent lateral tubercles. Concomitantly, the venter

becomes tabulate, usually with distinct alternating clavi. Ventral clavi and

umbilical tubercles generally appear simultaneously (Figs 12-13).

Upper Lower

Albian Cenomanian

1 = 5 My

BBY Shy

=<.

ra)

nN.

ealh

-|I—

Nz N

lla

Anaplacenticeras

Hengestites

Semenovites

Cleoniceras s.l. Karamaiceras

SS Placenticeras s.

HOPLITIDAE PLACENTICERATIDAE

Fig. 4. Phylogeny and sutural development of Placenticeratidae from Hoplitidae. (After

Mirzoev 1967, fig. 5.)

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

k 1 2

J | ~ Soa

V oak (ier Leas

: h MY Uy y lvdvw | avdy Nvadl'p

q | / ll eo

ae: | |

vvlyd

pol

‘ Vv L U i i Size “Os

| aij Usyy vd. 5 dp

1 yin UJ 7 Jyd Jq

= |

Fig. 5. Sutural ontogeny. A. Anahoplites michalski. B. Kopetdagites grossouvrei. C. Besch-

tubeites beschtubensis. D. Placenticeras pitniakense. (After Michailova 1978, fig. 1.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 249

aia

Smm

| lI III IV V VI

—_+ —_____Y SSE

ventral lateral lobes

lobe

Fig. 6. Morphographic terminology applied to sutures in placenticeratids. A. Karamaites

grossouvrei (Semenov, 1899). B. Karamaites gaurdakense (Luppov, 1963). C. Karamaites

mediasiaticum (Luppov, 1963). (After Marcinowski 1980, fig. 13.)

(d) Late phragmocone and body chamber stage. Here the whorl section becomes

even more inflated and rounded, the umbilical tubercles (if present) may migrate

outward, in some up to midflank, and the venter becomes distinctly rounded and

eventually may be completely smooth on the body chamber with no ventral clavi,

if these were present initially. In addition, the umbilical seam egresses and part of

the body chamber may show slight scaphitoid uncoiling (e.g. Hyatt 1903, pl. 32;

herein Figs 12-13).

This generalized sequence of ontogenetic change can be recognized in most

species of Placenticeras, including the Zululand material, with minor deviations.

It is important to note that the rate and duration of each ontogenetic stage is

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

ms 5mm

IV V

5mm

Fig. 7. Variation in size of fourth and fifth lateral lobes. A. Karamaites kutuzovae (Ijin, 1975).

B. Proplacenticeras planum Ijin, 1975. C. Placenticeras luppovi Iljin, 1975.

(After Marcinowski 1980, fig. 14.)

extremely variable. This is especially true of the two middle growth stages. After

the juvenile stage, which, naturally, is present in all species, the shell may pass

through all three successive stages to maturity. In others, however, the smooth,

early phragmocone stage is retained to maturity, without a tuberculate stage, or

the smooth stage may be very short, resulting in forms with strong tubercles and

ribbing at very small diameters.

In summary, part of the difficulty in unravelling the systematics of Placen-

ticeratidae lies in the fact that ontogenetic changes occur at different rates in

different individuals of the same species, while others may omit later develop-

mental stages. As a result, assemblages can include adult forms with the same

ornament as juveniles, as well as individuals where late typically phragmocone

modifications already occur at very early growth stages of some forms. In rare

cases, an ontogenetic stage can even be omitted.

Dimorphism in Placenticeras, more specifically P. polyopsis (Dujardin,

1837), was demonstrated by Summesberger (1979) (as Stantonoceras depressum | |

(Hyatt)) and Kennedy & Wright (1983). Until quite recently, recognition of |

dimorphism was mainly restricted to pre-Cretaceous ammonites (e.g. Makowski |

1962; Callomon 1963, 1981), the notable exception being the Late Cretaceous

scaphitids (Cobban 1969). Only in recent years has dimorphism been recognized |

CRETACEOUS FAUNAS FROM SOUTH AFRICA Zo

MAASTRICHTIAN

Hoplitoplacenticeras

Metaplacenticeras

CAMPANIAN

SANTONIAN

CONIACIAN

TURONIAN

CENOMANIAN

Placenticeras

Hypengonoceras' Hengestites

ALBIAN (part)

Karamaites

Metaclavites

Fig. 8. Phylogeny of Placenticeratidae. (After Kennedy & Wright 1983, text-fig. 5.)

in Cretaceous forms and Kennedy & Wright (1985) now claim to have recognized

it in all but three Upper Cretaceous families. Dimorphism in Cretaceous taxa is

mainly limited to differences in relative adult size (see especially Callomon (1981:

269-270) on the Albian species described by Scholz (1979) as Pervinquieria

(Subschloenbachia) rostrata and Hysteroceras (Cantabrigites) cantabrigense). For

recent discussion see Kennedy & Wright (1985).

Unequivocal dimorphic pairs can only be recognized in adult individuals.

Criteria for recognizing adult specimens as applied mainly to Jurassic forms (cf.

Makowski 1962; Kennedy & Cobban 1976) include:

1. Slowing of growth, indicated by crowding and often interference of later septa

and/or simplification of the sutures.

2. Changes in ornament on the body chamber—either attenuation of the strong

ornament found on the phragmocone, or appearance of strong ornament on the

body chamber following weak ornament on the phragmocone. The former

commonly characterizes macroconchs, the latter microconchs.

3. Egression of the umbilical seam, leading to a slight scaphitoid uncoiling of the

body chamber.

4. Apertural modifications.

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Placenticeras kaffrarium Etheridge, 1904. Protoconch and early whorls. Note nepionic

constriction. NMB D941/7. x 25.

Fig. 10. Placenticeras kaffrarium Etheridge, 1904. Early ontogeny of whorls up to

‘P. umkwelanense’ stage. Unregistered specimen. Scale bar in mm.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 253

Some of these criteria are difficult to apply to placenticeratids. The septa in

most placenticeratids are already closely spaced in early parts of the phragmo-

cone, and interference of sutures is by no means restricted to the terminal stages

of growth. Apertural modifications are difficult to recognize in this family,

because complete apertures are rarely preserved. To our knowledge, the only

complete Placenticeras aperture was recorded by Vredenburg (1907); we have but

a single questionable example from Zululand (Fig. 14A).

Instead, egression of the umbilical seam and consequent scaphitoid uncoiling

of the body chamber, changes in the whorl section, and modifications of ornament

are much better indicators of maturity. Scaphitoid uncoiling of the body chamber

can be very obvious (e.g. Hyatt 1903, pl. 32), so much so that Chiplonkar &

Ghare (1977a) even erected a new genus Placentoscaphites within the family

Scaphitidae for what are in part mere Placenticeras adults with uncoiled body

chamber. This uncoiling of the body chamber is accompanied by rounding of the

venter and loss of ventral clavi. In addition, the umbilical wall and shoulder

flatten, the umbilical diameter increases, and the umbilical edge becomes rounded

towards and on the body chamber. In some forms with strong ornament, the

umbilical tubercles migrate away from the umbilical edge towards maturity and

weaken, or even disappear. Conversely, forms with smooth phragmocones may

suddenly develop weak umbilical and lateral ornament on the body chamber

(Fig. 46).

The manifestations of dimorphism in placenticeratids are complex, and not as

simple as implied by Kennedy & Wright (1983). The general trend seems to be

that in the early (Albian and Cenomanian) forms, such as ‘Karamaites’, dimor-

phism is mainly restricted to size, whereas later forms, e.g. Santonian Placen-

ticeras polyopsis (Dujardin), have strongly differentiated dimorphic pairs, with

microconchs having much stronger ornament at smaller diameters, whereas

macroconchs retain the smooth or weakly ornamented stage to much greater

diameters and are generally weaker ornamented at maturity than their micro-

conch counterpart (i.e. Stantonoceras vs Placenticeras). Partly because of this,

Kennedy & Wright (1983: 868) decided to retain Karamaites as a separate taxon,

indicating, however, that should subsequent investigations show Karamaites to

have strongly differentiated dimorphs, it should be reduced to a synonym of

Placenticeras.

Recent work by Seyed-Emami et al. (1984) suggests that some early (Middle

Cenomanian) placenticeratids are already strongly dimorphic. Karamaites gros-

souvrei (Semenov, 1899), which may reach diameters of up to half a metre, and

the very closely allied if not conspecific Karamaites mediasiaticum (Luppov,

1963), with weak lateral ornament and compressed whorl section, occur together

with the smaller, strongly ornamented Karamaites gaurdakense (Luppov, 1963).

Middle Coniacian Placenticeras kaffrarium from Zululand also show distinct

dimorphic pairs, macroconchs retaining weaker ornament to greater diameters

than the strongly ornamented microconchs. However, distinct dimorphic pairs

that differ only in terms of relative size also occur. Thus dimorphism in itself is a

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Placenticeras kaffrarium Etheridge, 1904. A-C. SAS Z1520. D-F. NMB D941/48.

Specimens showing compressed, smooth early phragmocone ‘umkwelanense’ stage. Note slight

falcoid swellings on flanks.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 255

Fig. 12. Placenticeras kaffrarium Etheridge, 1904. SAM-PCZ6194. Macroconch showing

distinct ontogenetic changes from ‘umkwelanense’ through ‘subkaffrarium’ to adult ‘kaffrarium’

stage. xX 0,7.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Placenticeras kaffrarium Etheridge, 1904. SAM-PCZ6194. Macroconch showing |

distinct ontogenetic changes from ‘umkwelanense’ through ‘subkaffrarium’ to adult ‘kaffrarium’ |

stage. X 0,7.

oO7

CRETACEOUS FAUNAS FROM SOUTH AFRICA

spaaouaovjdojdoyy *q

‘TX “YQouooo1IA “pleld AWN

peaiesaid ainjiode ayqissod yyim youosossrpy

OL61 “UOUsTTTOD rys4nmMoy

‘I80IZ SWS ‘PO6I “eSpuoyIg winuvsffoy spsaonuavjg “pT “314

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

difficult criterion to apply for generic separation and, on that basis alone,

Karamaites does not bear separation from Placenticeras.

There is thus a distinctive trend in the nature of dimorphism in placenti-

ceratids from size alone, to size and differences in strength of ornament.

Ornament-related dimorphism becomes obvious only when distinct lateral orna-

ment, especially lateral tubercles, developed—on present evidence, mainly

during the Turonian and Coniacian. At present it is not possible to pinpoint the

stage at which size related dimorphism gave way to ornament- and size-related

dimorphism. In any case, the transition is gradual and cannot be used as a basis

for generic and subgeneric division of the groups.

VARIATION IN RELATIVE PROPORTIONS

Intraspecific variation in the strict sense is difficult to separate satisfactorily

from the other variable factors, unless dealing with adult specimens. In immature

phragmocones one is never sure if differences in ornamentation are due to simple

intraspecific variation or because specimens are at a different ontogenetic stage.

In the Zululand specimens very conspicuous differences are found in the whorl

sections of specimens from the same restricted stratigraphic levels but with more

or less the same type of ornament. Thus the ratio of whorl breadth to whorl height |

appears to be in part independent of ontogenetic change, and part of true |

intraspecific variation. Intraspecific variation in the adult stage is very obvious, as

far as ornament is concerned, as 1s reflected by the multitude of names in current

use for specimens from the same restricted stratigraphic level.

PHYLOGENETIC CHANGE

Probably the most difficult factor to understand in the systematics of

Placenticeratidae is the role of phylogenetic change. Intraspecific variation and

the nature and rate of ontogenetic change are far more prominent, and com-

pletely overshadow morphological changes through time. As far as coiling is

concerned, Placenticeratidae are extremely conservative and are characterized by

a narrowly umbilicate shell form. Apart from slight modifications on the later part

of the phragmocone and on the body chamber, very little change can be detected

in the coiling of the shell. Only changes in ornament take place and these are very

subtle.

The earliest forms, e.g. Karamaites in the Upper Albian, already have

umbilical tubercles, ventral clavi, and falcoid or sickle-shaped ribs at some

ontogenetic stage. Initially the umbilical tubercles are the dominant element of

the ornament but, gradually, feeble lateral swellings appear on the flanks at the

point where the ribs bifurcate. Distinct swellings or tubercles develop mainly

during the Upper Cenomanian to Lower Turonian, but are well established by the

Upper Turonian. Appearance of lateral tubercles is also associated with the

development of stronger ribbing. Eventually lateral ornament becomes dominant

in the Coniacian, and distinct dimorphic pairs can be recognized on the basis of

ornament as well as disparate size in the Coniacian. During the Santonian the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 259

picture is less clear. In one branch of the placenticeratids there is a distinct trend

towards outwards migration of the tubercles—the umbilical tubercles migrate

away from the umbilical wall to a point near mid-flank, and the lateral tubercles

migrate close to the ventral tubercles. In addition, there appears to be a gradual

reduction in ribbing. This trend can be seen in the Santonian Placenticeras

polyopsis (Dujardin). This branch probably gave rise to the Upper Campanian to

Upper Maastrichtian Hoplitoplacenticeras. This genus displays as much variation

as all the other placenticeratids during the whole of the stratigraphic range

of the family. At the same time the genus Metaplacenticeras arose. It has approxi-

mately the same stratigraphic range as Hoplitoplacenticeras but is said to

differ in possessing a median keel at some stage of growth. This is to be discussed

below. However, Metaplacenticeras presumably arose from the same stock as

Hoplitoplacenticeras.

The other branch of placenticeratid evolution is the poorly known Maastrich-

tian genus ‘Gissarites’. This displays a very narrow venter and apparent reversal

to early Placenticeras ornament, consisting of conical umbilical tubercles, weak

lateral nodes, and very fine external clavi. The Lower Campanian ‘Diplacmo-

ceras’ bidorsatum perhaps occupies an intermediate position between Placen-

ticeras paraplanum and ‘Gissarites’ and is possibly the rootstock of ‘Gissarites’.

GEOGRAPHIC VARIATION

Another factor affecting variation may possibly be geographic distribution.

With the information at our disposal, it is difficult to separate the effects of

geographic isolation and possible development of subspecies and phenotypic

variation related to environmental control. A prerequisite for any comparisons of

this kind is precise stratigraphic control, based on taxa other than placenticer-

atids, unless we are to become involved in a circular argument. The only locality

where such correlation with Zululand is possible is Madagascar. Judging by the

Coniacian faunas from Betioky, illustrated by Collignon (19655), it seems that

placenticeratids are much less common there and, also, that coarsely ornamented

forms of Placenticeras kaffrarium are absent (or at least not illustrated). It is

impossible to determine whether this is due to genetic differences between what

are clearly populations of the same species, or a phenotypic response to different

environmental conditions.

SUTURAL ONTOGENY, PHYLOGENY AND SYSTEMATICS

Under the heading of intraspecific variation we mentioned that the Placenti-

ceratidae, with the exception of Hoplitoplacenticéras, show very little morpho-

logical change over their entire stratigraphic range from the Albian to the

Maastrichtian. In fact, some of the Placenticeras populations we have studied

show as much morphological variation at a single stratigraphic level as the whole

genus throughout its entire history. In part because of the relatively low

systematic value of ornament, various attempts have been made to base the

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Saddle incisione—Saddle division«—Intermed. type«—Lobe division

—?+=Lobe formation—?—>

eLO aie ar ae i gree o , an

te oan up du,

Be “ 56

a a a

Fig. 15. Principles of lobe formation after Wiedmann (1970).

systematics of the group on details of the adult suture. Inseparable from this are

attempts to use sutural ontogeny to trace the ancestry of the placenticeratids to

the Hoplitidae.

Details of the sutures used in placenticeratid systematics are as follows:

1. General outline of the lobes and saddles in the adult external suture.

2. Comparative ontogenetic studies of the sutures.

3. Number of lobes and saddles in the adult external suture.

The family Placenticeratidae is one of several groups of ammonites referred

to as ‘Pseudoceratites of the Cretaceous’ by Hyatt (1903) in his posthumous work,

in which “The complexity of the outlines of the lobes and saddles ... is a

retrogressive form that mimics to a certain extent primitive forms among

Goniatitinae and Ceratitinae’ (1903: 21), or, as defined by Casey (1978: 584)‘. . .

ammonites characterized by simplicity of the saddles of the suture-line though

with an increased number of elements’.

This type of suture line is often found associated with a narrowly umbilicate,

compressed shell form. In some of the Placenticeratidae, in the sense of Kennedy

& Wright (1983), the term pseudoceratitic is still applicable, but not in the

majority. In Hypengonoceras, which we do not regard as a placenticeratid, the

saddles in the external suture generally tend to be rounded and less divided than

the lobes, but not always. In all the other genera referred to the Placenticeratidae

by Kennedy & Wright (1983), both saddles and lobes are finely frilled and the

term ‘pseudoceratitic’ is a misnomer. Thus from this point of view, the family

Placenticeratidae—as interpreted by Wright (1957) and Kennedy & Wright

(1983)—is a heterogenous group. This is discussed further elsewhere (see p. 361).

Unfortunately, a discussion on the comparative sutural ontogenies of the

Placenticeratidae (or any other group for that matter) is very difficult, because of

lack of definition of individual elements and disparate uses of terminology.

Sutural terminology is either morphological and descriptive (‘morpho-

graphic’) or ontogenetic. Morphological terminology is useful for communicating

CRETACEOUS FAUNAS FROM SOUTH AFRICA 26%

Wedekind (1913) Ruzhentsev (1949b)

V (ventral)

O (omnilateral)

+U (umbilical s. str.)

Fig. 16. Comparison of sutural terminology of Wedekind (1913) and Ruzhentsev (1949). (After

Wiedmann & Kullmann 1981: 224.)

data concerning the adult suture line, but even here it is severely limited by lack

of precision. Casual reference to the number of lateral lobes, adventive and/or

auxiliary elements in the adult external suture line generally confuses rather than

clarifies.

Morphogenetic terminology, in which various elements of the suture can be

homologized in ontogenetic studies, appears to be the ideal. Unfortunately there

are difficulties which appear to be semantic and to a certain extent geopolitical.

Two contrasting types of terminology exist: that of Wedekind (1916) is generally

used by western palaeontologists, and that of Ruzhentsev (1949, 1957) is generally

used by Soviet palaeontologists (see Kullmann & Wiedmann 1970; Wiedmann &

Kullmann 1981; Fig. 16 herein). For reasons of clarity, summarized by Wiedmann

& Kullmann (1981), we prefer the terminology of Wedekind (1916) but, given

that most recent work on placenticeratid sutural ontogeny has been conducted by

authors employing Ruzhentsev’s terminology, both are given where possible, to

avoid confusion (see e.g. Casey 1978: 585).

The early sutural ontogeny is well known for various placenticeratids (see list

below), although differences of interpretation exist.

Smith 1900 Metaplacenticeras pacificum (Smith, 1900)

Hyatt 1903 Placenticeras pseudoplacenta Hyatt, 1903

Placenticeras whitfieldi Hyatt, 1903

Matsumoto 1953 Metaplacenticeras subtilstriatum (Jimbo, 1894)

Schindewolf 1967 Metaplacenticeras pacificum (Smith, 1900)

Mirzoev 1967 Karamaiceras kolbajense Sokolov, 1965

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Iljin 1975 Kopetdagites kopetdagensis Ijin, 1975

Mediasiceras beliakovae Ijin, 1975

Beschtubeites beschtubensis Ijin, 1975

Proplacenticeras pitniakense Iljin, 1975

Michailova 1978 = Karamaiceras kolbajense Sokolov, 1965

Turkmenites gaurdakensis Luppov, 1963

Kopetdagites grossouvrei (Semenov, 1899)

Kopetdagites sp.

Mediasiceras saggitalis Wjin, 1975

Beschtubeites beschtubensis Ijin, 1975

Beschtubeites kutusovae Ijin, 1975

Placenticeras bobkovae Ijin, 1975

Placenticeras pitniakense Iljin, 1975

Results of these investigations are unanimous in ascertaining that the primary

suture is quinquelobate: E L U, U, I (Wedekind); or V U U' I D (Ruzhentsev).

Very early in the ontogeny, the umbilical lobe U, (Ruzhentsev = I) divides into

two elements through the process of ‘Lobenspaltung’ (see Wiedmann 1970: 910,

fig. 1 IV) (Fig. 15), giving rise to U,,U,, of typical hoplitid affinities, resulting in

an adult sutural formula: E L U, U3; (U4 = S) U,,U;4 I (Wedekind).

Less easy to interpret (and reconcile) are: division of the lateral lobe L

(Ruzhentsev U); division of the saddle E/L (Ruzhentsev V/U); and, division of

the saddle L/U, (Ruzhentsev U,/U’).

According to Mirzoev (1967) and Michailova (1974, 1978), the lateral lobe L

(Ruzhentsev U) divides into two unequal parts very early in ontogeny. This bifid

nature of the lateral lobe is in direct contrast to the trifid division of the same lobe

in the Hoplitaceae. Soon afterwards a lobe arises in the lateral saddle E/L

(Ruzhentsev V/U), which is designated L by Mirzoev and Michailova, i.e., the

equivalent of A (adventive) of Wedekind. Furthermore, in Placenticeras s.s. a

new lobe arises in the saddle L/U, (Ruzhentsev—U,/U’), to which they apply the

symbol U'" (= new U')—in descriptive terms the fourth lateral lobe in the adult

suture (Fig. 5D).

The investigations of Mirzoev and Michailova are somewhat at variance with

those of Schindewolf (1967) and Kullmann & Wiedmann (1970). According to the

latter, the lateral lobe L is trifid, and no mention is made of a new lobe being

formed in the saddle L/U,. This disparity may in part be due to differences of

definition. Schindewolf (1967) had already pointed to the indiscriminate use of the

term ‘adventive lobe’. As per definition, an adventive lobe is a true lobe formed

in the external saddle (E/L) at an early ontogenetic stage. Thus reference to

‘adventitious elements’ in the adult placenticeratid suture (e.g. Wright 1957;

CRETACEOUS FAUNAS FROM SOUTH AFRICA 263

Kennedy & Wright 1983), when in fact a tripartite division of the lateral lobe or

a bipartite division of L, plus a deep incision on E/L is meant, is misleading, as

Schindewolf (1967) had already commented. More difficult to resolve is the

question whether the adventive lobe (Ruzhentsev—L) of Mirzoev and Michai-

lova is in fact a true adventive lobe as per definition, or merely a deep incision of

the saddle E/L. The time of formation seems to be crucial for definition.

According to Schindewolf (1967: 803 (725)) the umbilical lobes are formed at an

early ontogenetic stage—prior to incision of the saddles and the lobes. One would

expect the same of adventive lobes.

According to Michailova (1978) the ‘adventive’ lobe only arises from the end

of the second whorl onwards. Illustrations by Michailova (1978, fig. lv, g) also

appear to indicate that the ‘adventive’ lobe only really becomes prominent at a

stage that coincides with the incision (Zerschlitzung) of the other saddles and

lobes. This appears to favour the idea that we are here dealing with a large

incision in the E/L saddle, rather than a true adventive lobe—as also previously

suggested by Smith (1900), Reeside (1926: 2), Matsumoto (1953: 146) and

Schindewolf (1967: 744 (666)).

Definition of the incision of the lateral saddle E/L as an incision and not as

an adventive lobe is not merely a matter of semantics or terminology, but crucial

for determining phylogenetic relationships. True adventive lobes occur in the

family Engonoceratidae, but not in Placenticeratidae. Wright (1957) and pre-

viously Spath (1931) had sought to derive the Placenticeratidae from Engono-

ceratidae—a view now generally abandoned. Furthermore, arising from study of

the sutural ontogeny of the group is the problem of the systematic position of the

Fig. 17. Suture of Hypengonoceras warthi (Kossmat, 1895). (After Kossmat 1895, pl. 20 (6)

(fig. 8).)

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Albian genus Hypengonoceras, to be discussed below (see p. 361). Engono-

ceratidae can generally be separated quite easily from the Placenticeratidae by

their ceratitic suture lines with entire or little divided saddles (Fig. 17). In some

cases, however, such as the genus Parengonoceras Spath, 1924 (of Engonocera-

tidae), the sutures are finely incised, and morphological transitions to the genus

Hypengonoceras (of Placenticeratidae as interpreted by Kennedy & Wright

(1983)) occur, as demonstrated by Renz (1970). The only way to distinguish

between Parengonoceras and Hypengonoceras is by means of sutural ontogeny.

Unequivocal adventive lobes occur in Parengonoceras (Fig. 18), which clearly

places the genus in the family Engonoceratidae. As yet, the sutural ontogeny of

Hypengonoceras is unknown, and reference to the Placenticeratidae appears to be

based on stratigraphic rather than phylogenetic evidence. Details of this are

discussed elsewhere (p. 361). For the present it is sufficient to say that the

distinguishing feature between Engonoceratidae and Placenticeratidae appears to

be the presence of one or more adventive lobes in the former, as compared to a

mere, albeit large, incision of the lateral saddle E/L in the latter.

The lobe in the saddle L/U, (fourth lateral lobe), termed U'" by Michailova

(1974, 1978), has been used as a criterion for separating genera in the Placenti-

ceratidae. We lack material for detailed ontogenetic studies and can give no

verdict on the sutural status of the lobe—whether it in fact is a real (new) lobe or

merely an enlarged incision, similar to that in the external saddle; we suspect it to

be the latter. Whatever the case may be, the presence, or rather the size of this

‘lobe’ is regarded as being of systematic value by Michailova (1978) and

characteristic of the genus Placenticeras. In terms of descriptive terminology of

the adult suture, this is the fourth lateral lobe. As summarized by Marcinowski

(1980), the early (Albian—Cenomanian) placenticeratids referred to the genus

Karamaites also have this lobe, but it is consistently smaller than the fifth lateral

lobe (Figs 6-7). There is a progressive, phylogenetic increase in the size of this

‘lobe’ until in Placenticeras, in the Turonian, it exceeds the fifth lateral lobe in

size. In addition, Michailova (1978) claimed that the bases of the first three lateral

lobes (Ruzhentsev’s L, U*U') widen and are bulb-like at the base, but become

narrower at the neck.

An apparent final trend in sutural development of the Placenticeratidae is to

be found in the Maastrichtian genus ‘Gissarites’ Iljin, 1958 (Fig. 7A). Here the

sixth lateral lobe is larger than the fifth, in contrast to the situation in Placen-

ticeras, where the opposite holds true. In addition, the lateral branches of the

external lobe E (Ruzhentsev = V) are short and blunt in contrast to paw-shaped

in Placenticeras.

From the above it can be seen that details of the suture line are of rather

ambiguous value in the systematics of the Placenticeratidae, especially at generic

level.

Detailed studies of the sutural ontogeny, however, point unambiguously to

the origins of Placenticeratidae in Hoplitidae. Salfeld (1924) and Matsumoto

(1953) tried to trace the origins of Placenticeratidae back to the Phylloceratidae.

——

»-

CRETACEOUS FAUNAS FROM SOUTH AFRICA 265

Salfeld based his conclusions on incorrect sutural data from the type species of

Hypophylloceras, as shown subsequently by Wiedmann (1962: 248). Matsumoto

based his findings on supposed affinities of the juvenile shells of Metaplacenticeras

subtilstriatum and Phylloceras, and also claimed that the sutures of Metaplacenti-

ceras were somewhat phylloid—analogous to those of Neophylloceras. Also, the

fact that the siphuncle only reached its marginal position towards the end of the

second whorl was claimed to be a phylloceratid feature. Spath (1930) and Wright

(1957: L109, L390) tried to derive the Placenticeratidae from the Engono-

ceratidae, but the reasons for this are now seen as a combination of homoeo-

morphy and stratigraphic “‘neatness’ rather than actual relationship. It is now

generally accepted that the origin of Placenticeratidae lies in the Hoplitidae—a

theory dating back to Douvillé (1890), and followed by Kossmat (1897), Smith

(1900), De Grossouvre (1894), Pervinquiére (1907) and others.

Casey (1965: 461) suggested that early placenticeratids, such as Hengestites

Casey, 1960, Anaplacenticeras Ijin, 1959, and Karamaites Sokolov, 1965, arose

from the hoplitid Semenovites Glazunova, 1960. Schindewolf (1967) summarized

alleged phylogenetic relationships of Placenticeratidae and confirmed that the

origins must lie in the Hoplitidae. Mirzoev (1967) undertook detailed studies of

Cleoniceras, Semenovites and ‘Karamaiceras’, and substantiated the derivation of

Placenticeratidae from Hoplitidae via subfamily Semenovitinae Mirzoev, 1967,

the latter consisting of the two genera Semenovites Glazunova, 1960, and

Dm 10mm

Dm5,5mm

| DmSmm

e ‘ | / / of Dm 4mm

Chg tina a

: iy | Dm 2,3mm

b | aaa a Dm 2mm

a ee fr Dm1,2mm

Fig. 18. Sutural ontogeny of Parengonoceras discoides Renz. (After Renz 1970, fig. 2.)

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Metaclavites Casey, 1965 (see Fig. 4). It is significant that in the transition from

Hoplitidae to Placenticeratidae as demonstrated by Mirzoev, no significant

change in the form of the shell or in the degree of involution took place. Major

changes affected the suture line. The lateral lobe (L) (Ruzhentsev = U) changes

from asymmetrically trifid to bifid, and almost separated into two independent

lobes. Simultaneously, an incision arose in the first lateral saddle, separating it

almost as far as the base and the external lobe (Ruzhentsev = V) becomes

shallower. This all led to the typical ‘sagging’ suture line of Placenticeratidae.

Derivation of Placenticeratidae from Hoplitidae was further confirmed by the

detailed sutural studies of Michailova (1974, 1978), who derived ‘Karamaiceras’

from the Late Albian Anahoplites. In addition, she also listed the sutural

characteristics of the family as discussed above. These views were accepted by

Casey (1978).

Kennedy & Wright (1983) derived the mainstream of Placenticeratidae—

Karamaites and Placenticeras—from Metaclavites (Fig. 8) and _ tentatively

regarded Hypengonoceras and Hengestites as early, parallel offshoots derived

from Karamaites in the Albian. For reasons discussed below (p. 361), we prefer

to allocate Hypengonoceras to the family Engonoceratidae. This only leaves us

with the moot question of the systematic position of Hengestites. Unfortunately,

we have no additional material for meaningful discussion. As far as ontogenetic

development of ornament is concerned, Hengestites deviates distinctly from the

normal placenticeratid pattern. In Hengestites the ornamented stage with alter-

nating ventral clavi and lateral riblets precedes the smooth compressed stage with

entire venter; in Placenticeras the inner whorls are smooth with entire venter and

may be succeeded by an ornamented stage on the outer whorls, i.e. the complete

opposite. Removal of Hengestites from Placenticeratidae can only be seen as a

suggestion, pending detailed examination of the sutural ontogeny.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA Cuvier, 1797

Class CEPHALOPODA Cuvier, 1797

Order AMMONOIDEA Zittel, 1884

Suborder AMMONITINA Hyatt, 1889

Superfamily HOPLITACEAE Douvillé, 1890

Family Placenticeratidae Hyatt, 1900

(= Hypengonoceratidae Chiplonkar & Ghare, 1976;

Baghiceratinae Chiplonkar & Ghare, 1976)

Genus Placenticeras Meek, 1876

Placentocerus Meek, 1870 (nom. oblit.)

Diplacomoceras Hyatt, 1900

CRETACEOUS FAUNAS FROM SOUTH AFRICA 267

Diplacmoceras Hyatt, 1903 (illegitimate emendation of Diplacomoceras Hyatt, 1900)

Stantonoceras Johnson, 1903

Proplacenticeras Spath, 1926

Pseudoplacenticeras Spath, 1926

Anaplacenticeras Ijin, 1959

Gissarites Ijin, 1959

Karamaites Sokolov, 1961 (nom. nud.)

Parastantonoceras Collignon, 1965a

Karamaites Sokolov, 1965, in Casey, 1965

Karamaiceras Sokolov, 1967

Asiatostantonoceras Ijin, 1975

Turkmenites Ijin, 1975

Kopetdagites Iljin, 1975

Mediasiceras Iljin, 1975

Beschtubeites Ijin, 1975

Baghiceras Chiplonkar & Ghare, 1976

Malwiceras Chiplonkar & Ghare, 1976

Placentoscaphites Chiplonkar & Ghare, 1977a

Sancarlosia Chiplonkar & Ghare, 1978

Type species: Ammonites placenta DeKay, 1828, by subsequent designation

by Meek (1876: 462).

Diagnosis

Extremely variable, generally compressed and involute. Earliest stages with

rounded whorl section soon followed by typical compressed, high-whorled stage

with crater-like umbilical pit, flat smooth flanks and smooth flat venter with sharp

edges. Later stages may develop umbilical tubercles and generally ventral clavi,

which are demonstrably the base of a septate spine in some; lateral sickle-shaped

ribs with or without lateral thickening or tuberculation. Umbilical tubercles may

Or may not migrate away from umbilical edge towards maturity. At maturity,

ventral clavi generally disappear, the venter becomes rounded, the umbilical wall

slopes out gently and the umbilical seam egresses, leading to slight scaphitoid

uncoiling of the body chamber. Variation partially caused by different rates of

ontogenetic development. Strongly dimorphic; especially noticeable in forms with

lateral ornament. In early forms, dimorphs differ mainly in size. Suture charac-

teristically sagging, with saddle E/L deeply incised, and lateral lobe, L, arranged

obliquely in adapical direction, whereas umbilical lobes on flanks are arranged

more or less radially.

Discussion

The plethora of names listed in the generic synonomy bears testimony to the

confusion in placenticeratid systematics. Kennedy & Wright (1983) placed most of

these names in the synonymy of Placenticeras. They did, however, retain

Karamaites Sokolov because of lack of evidence of definite dimorphism in that

genus, as discussed above.

The earliest forms, ‘Karamaites’ of the Upper Albian to Middle Cenomanian,

are best known from Central Asia. They have ornament consisting of umbilical

tubercles, falcoid ribbing and alternating ventral clavi. This includes forms such as

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘Turkmenites’, ‘Kopetdagites’ and ‘Mediasiceras’. In the Upper Cenomanian

Kopetdagites, indications of later typical Placenticeras ornament start appearing.

Small tubercles or tubercle-like swellings develop on the flanks at the point of

bifurcation of the ribbing. This then leads to the typical ‘Proplacenticeras’ type of

ornament, shown best by P. kaffrarium, where strongly differentiated dimorphs

can be distinguished. In the Turonian to Coniacian ‘Proplacenticeras’, strong

lateral ornament develops and in the adult stage the umbilical tubercles may

migrate outwards, foreshadowing the Santonian Placenticeras polyopsis type of

ornament. Strongly ornamented microconchs correspond to Collignon’s (1965a)

genus Parastantonoceras. In later Placenticeras species, strongly ornamented

microconchs correspond to ‘Stantonoceras’ or ‘Asiatostantonoceras’. Placento-

scaphites clearly represents adult placenticeratids in which the body chamber

shows signs of uncoiling, and has nothing to do with the true scaphitids

whatsoever.

Of the other names included in the synonymy of Placenticeras, Sancarlosia is

an inflated, strongly ornamented form of Placenticeras, probably representing

microconchs, as may be Baghiceras and Malwiceras.

Gissarites (Fig. 1A) is a poorly known form from the Maastrichtian of

Central Asia. It has a very narrow venter and, in many respects, resembles the

early Placenticeras, with prominent umbilical tubercles but hardly any lateral

ornament. I[ljin (1958) maintained that the sixth lateral lobe is larger than the

fifth, by virtue of which Gissarites can be separated from Placenticeras. Due to the

great variation in this character, we do not believe that separation on details of the

suture is advisable. Another poorly known taxon that probably does not require

generic separation from Placenticeras, is the Lower Campanian Diplacomoceras.

This retains the entire venter to great diameters and, according to Hyatt (1903:

242), combines features of Engonoceratidae as far as ornament is concerned, but

has distinct placenticeratid sutures. ‘Diplacmoceras’ seems merely to be another

example of juvenile ornament being retained to the adult stage and can be derived

from Placenticeras paraplanum Wiedmann of the Upper Santonian (Kennedy

1986). ‘Pseudoplacenticeras’ is similar to ‘Diplacmoceras’; it is known only from

crushed specimens from a single locality (?Campanian) in Austria. Again,

separation seems unnecessary.

Occurrence

Upper Albian to Maastrichtian, world-wide.

Placenticeras kaffrarium Etheridge, 1904

Figs 9-14A, 19-20, 22-99

Placenticeras kaffrarium Etheridge, 1904: 89, pl. 3 (fig. 16). Besairié, 1930: 636, pl. 64

(fig. 3-3a), suture no. 8. Venzo, 1936: 107.

Placenticeras umkwelanensis Etheridge, 1904: 89, pl. 3 (figs 17-20).

Placenticeras whitfieldi (auct. non Hyatt): Boule, Lemoine & Thévenin, 1907: 48, pl. 12

(fig. S-Sa). Venzo, 1936: 107, pl. 11 (fig. 12).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 269

Placenticeras subkaffrarium Spath, 1921: 247, pl. 21 (fig. 2). Besairié, 1930, pl. 46 (figs 2-3).

Placenticeras cf. subkaffrarium Spath, 1921: 300.

Placenticeras whitefieldi |sic| Besairié, 1930, pl. 46 (fig. 1).

Placenticeras n. sp. aff. kaffrarium Etheridge: Venzo, 1936: 108, pl. 11 (fig. 13), text-fig. 3.

?Metaplacenticeras besairiei Collignon, 1936: 200, pl. 21 (figs 21-22).

Proplacenticeras memoriae-schloenbachi (Laube & Bruder) var. ambiloensis Collignon, 196Sa:

14, 16, pl. 381 (fig. 1646), pl. 382 (figs 1647-1648). Chiplonkar & Ghare, 1977c: 112,

fig. 6B-C.

Parastantonoceras murphyi Collignon, 1965a: 17, pl. 382 (fig. 1649).

Parastantonoceras besairiei (Collignon): Collignon, 1965a: 19, pl. 383 (fig. 1650).

Proplacenticeras stantoni Hyatt var. bolli Hyatt: Collignon, 1965a: 19, pl. 383 (fig. 1651).

Proplacenticeras stantoni Hyatt var. fortior Collignon 1965a: 19, pl. 383 (fig. 1652).

Proplacenticeras orbignyi (Geinitz): Collignon 1965a: 20, pl. 383 (figs 1653-1654).

Placenticeras reineckei Haughton, 1925: 271, pl. 13 (figs 4-5); 1926 (reprinted in French): 15,

pl. 2 (figs 4-5).

?Placenticeras merenskyi Haughton, 1930: 363, pl. 11 (figs 1-3).

Proplacenticeras aff. fritschi de Grossouvre var. eboroensis Collignon, 1965b: 38, pl. 430

(figs 1780-1781).

Proplacenticeras satriense Collignon 1965b: 40, pl. 431 (fig. 1782).

Proplacenticeras nov. sp. (Venzo) aff. kaffrarium Ether.: Collignon, 1965b: 40, pl. 431

(fig. 1783).

?Proplacenticeras merenskyi (Haughton): Klinger, 1977, fig. 7.

Placenticeras syrtale Morton var. tamulicum (Blanford) Kossmat: Boule, Lemoine & Thévenin,

1907: 47 (27), pl. 12 (figs 3-4).

?Proplacenticeras rampuraensis Chiplonkar & Ghare, 1977c: 109, figs 1-2, 6A, D.

?Proplacenticeras spathi Chiplonkar & Ghare, 1977c: 110, fig. 7E, F.

?Proplacenticeras fritschi (de Grossouvre): Chiplonkar & Ghare 1977c: 113, fig. 7B—C.

?Proplacenticeras stantoni (Hyatt): Chiplonkar & Ghare, 1977c: 114.

?Pseudoplacenticeras sp. cf. P. milleri (Hauer): Chiplonkar & Ghare, 1977c: 115, fig. 6E-F.

Proplacenticeras stantoni (Hyatt) var. bolli (Hyatt): Howarth, 1985: 84, fig. 7.

Type

The holotype (Fig. 19), by monotypy, is Etheridge’s original figured speci-

men (1904, pl. 3 (fig. 16)) from Umkwelane Hill, Zululand, housed in the Natal

Museum, Pietermaritzburg, NMP 355, type number T417.

Material

Etheridge’s syntypes of Placenticeras umkwelanense (1904, pl. 3

(figs 17-20)), from Umkwelane Hill, Zululand, housed in the Natal Museum,

Pietermaritzburg, NMP 453a—c, type number T418a—c. Lectotype here desig-

nated, NMP 453a (type number T418a) (Fig. 20A), the original of Etheridge’s

(1904, pl. 3 (fig. 17)); paralectotypes, NMP453b-c (type number T418b-—c)

(Fig. 20B—C). Spath’s (1921, pl. 21 (fig. 2)) holotype of Placenticeras subkaffra-

rium, SAM-5106 (Fig. 21). Spath’s Placenticeras cf. subkaffrarium (1921: 300),

SAM-—4957—4958 (Fig. 22). Several hundred specimens in the collections of the

South African Museum, Cape Town; Geological Survey, Pretoria; the British

Museum (Natural History), London; and University Museum, Oxford, from the

following localities.

Locality 10: railway cutting, 1,1 km north of Haig Halt, Mfolozi, imprecisely

located in most cases in the Coniacian.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

CRETACEOUS FAUNAS FROM SOUTH AFRICA 271

Fig. 20. Placenticeras kaffrarium Etheridge, 1904. A. NMP 453a, type number T418a. The

lectotype of Placenticeras umkwelanense Etheridge, 1904. B-—C. Paralectotypes of Placenticeras

umkwelanense, NMP 453b-c, type numbers T418b-c. D. SAM-—PCZ6190 from locality 60, the

basal Pterotrigonia bed, Skoenberg.

ANNALS OF THE SOUTH AFRICAN MUSEUM

272

Spath, 1921. SAM-5106. x 1.

tum

bkaffrar

tceras SU

The holotype of Placent

Fig. 20),

CRETACEOUS FAUNAS FROM SOUTH AFRICA 213

Locality 11: road cut, Mfolozi. Coniacian s.1. SAM—uncatalogued and unlocal-

ized material. .

Locality 13: hill slopes below Riverview compound, Coniacian II-III,

SAS H146/3.

Locality 16: small quarry east of track on Lot 71 13567 east of Riverview

sugarmill. Coniacian II or HI. SAM—uncatalogued material.

Locality 24: cuttings and excavations for new Nyalazi River bridge. Coniacia-

nlI-V. SAS A624.

Locality 25: cutting alongside road, 2,8 km ESE of Nyalazi River Trading Store.

Coniacian IT. SAM—uncatalogued and unlocalized material.

Locality 60: basal Pterotrigonia Bed, Skoenberg, and overlying silts. Coniacian I.

SAM-PCZ6190, PCZ6195, PCZ6200; NMB D917, D1267, D1269, D1271;

SAS Z201-Z205.

Locality 63: steep northern face of Skoenberg (Skoenberg East in Van Hoepen’s

catalogue), Coniacian I. NMB D941/1-51, D2846; SAS Z200.

Locality 71: degraded river cliffs on the north bank of the Munywana Creek north

of Skoenberg. Coniacian I. NMB D925/1-3, D1343-4; SAS Z252, Z401,

ZIG, ZV081, 7251, 2252/1.

Locality 72: degraded river cliffs and alluvial flats on the north side of the

Mzinene, 200-300 m east of the causeway across the river. Coniacian II? and

Ill. NMB D1148-D1150, D1151/1-15; SAS Z1028/1-3, A419, Z1125;

SAM-PCZ6201, Z6202.

Locality 92: bulldozer scrapings and adjacent hill slopes around the pumping

station at the southern end of the track leading south from the farm Panplaas.

Coniacian II and HI. SAM—PCZ6190, Z6192, Z8205; SAS Z907, Z1442a-b,

Z1520a—g, Z1603a—c, Z1063, Z1604, Z1065, 21681.

Locality 93: hill slopes extending on either side of the boundary fence of lots H101

and H102, Coniacian Il. SAM—PCZ6187—Z6188, 26193, Z6196—-6199,

Z6203-—Z6204, 26208, Z6215, Z6251—Z6255, 26258; SAM—93B; SAS Z77,

meee oe 07, Z510, 7512, 7519, 7678, 2680, 7681, 27682, 7686, 7688,

meee 297.120, 7880, 788/, 7888, 7916, Z917, 7919, 2920, 7926, 2927,

tm 2752, 2052, 7933, 2937, 2939, 7941, 2952, Z993, 7994, Z995,

m9, 21556,.2.1571.

Locality H196: on the west bank of the Hluhluwe River, at 32°19’30"E, 28°5'30"S.

Coniacian II. SAM—PCZ6497.

Locality 145: degraded bluffs on the eastern side of the Msunduzi, 3 km SW of the

farm Morrisvale. Coniacian Il. NMB D1184/1-26.

Over one hundred uncrushed specimens have been measured, including both

juvenile and adult, microconchs and macroconchs. Rather than list these

measurements in undigestible form, we have presented the data in graphic form

(Figures 51-55). Lists of measurements for statistical purposes are available upon

request from the Department of Invertebrate Palaeontology, South African

Museum.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 22. Placenticeras kaffrarium Etheridge, 1904. A-—C. SAM-—4958. The specimen described

by Spath (1921: 300) as Placenticeras cf. subkaffrarium nov. D-F. SAM-4957. The other

specimen described by Spath (1921: 300) as Placenticeras cf. subkaffrarium nov. Both xX 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 219

Fig. 23. Placenticeras kaffrarium Etheridge, 1904. Specimens showing different ontogenetic

changes. A-~B. NMB D917. x 1. C-D. SAM-PCZ6190. x 1,5. E-F. NMB D941/7. x 10.

ANNALS OF THE SOUTH AFRICAN MUSEUM

276

‘Tx ‘quswRUIO wntpdffoy, 9 addy 0} sadeys wniupiffoyqns, pue ,asuauvjamyuin, Ysnoy)

‘s]foym IOUT popuNos WOT sosuLYo daU9d0jUO dAISSId9NS SUIMOYsS YOUOSOINI, “OSTIG AWN ‘PO6L ‘28pusyig WNLADAL{DY SDAIINUIIV] g

0) q V

‘p7 “B14

On i)

‘Tx ‘a8eqs wniupvdf{oyqns, 0} ,asuauvjamyun,

passdidwios y8nory) ‘AjIvo popunol WoI sasueyo o1aU9d0jU0 SUIMOYsS uounseds “[STIG ANN ‘P06I ‘e8pueuI” wniuvaf{vy SpldIUIW]g “ST “BLA

CRETACEOUS FAUNAS FROM SOUTH AFRICA

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Description of the variation shown by the species is best accomplished in

terms of the the following factors.

1. Ontogenetic changes

For descriptive purposes the ontogeny in typical forms can be approximately

divided into four stages.

(a) Embryonic stage. This includes the very early stage with the protoconch and

subsequent early whorls. This can be seen in NMB D941/7 (Fig. 23E-F),

NMB D1150 (Fig. 24), SAS Z680, SAM-—PCZ6190 (Fig. 23C-D), NMB D1151

(Fig. 25). The protoconch is globular, as wide as high, followed by depressed,

reniform, early whorls. These soon become circular in cross-section at a diameter

of about 2 mm. Thereafter, the whorls soon become compressed, as in typical

large Placenticeras. The nepionic constriction can be seen in NMB D941/7

(Figs 9, 23E-F). Unfortunately most of these specimens have undergone second-

ary recrystallization, and details of the early sutural development are not clear,

nor can the position of the siphuncle be determined accurately.

(b) Early phragmocone stage (Figs 11, 23C-D). Here the form is at what may be

termed as the ‘umkwelanense’ stage, with very compressed whorls with maximum

breadth at the umbilical shoulder. The flanks are smooth, save for fine, falcoid or

sickle-shaped striae. The curve of the ‘sickle’ may be slightly stronger than the

haft, producing slight crescents on the ventral half of the flanks. The venter is

concave to flat and smooth, bordered on either side by entire, sharp, narrow

ventrolateral shoulders.

more inflated and rounded, the umbilical shoulder also becomes more rounded,

umbilical tubercles of varying strength, c. 7-8 per whorl develop, connected by

single or bifurcating ribs of varying strength to weak or prominent lateral

tubercles. Concomitantly, the venter becomes tuberculate with distinct alternat-

ing ventral clavi. Generally, as soon as the umbilical tubercles appear, the edges

of the venter become crenulate. This is the ‘subkaffrarium’ stage (Figs 26-28).

(d) Late phragmocone and body chamber. Here the whorl section becomes even

more inflated and rounded, the umbilical tubercles migrate outwards, weaken,

and the venter becomes distinctly rounded and eventually completely smooth on

the body chamber. Lateral ornament may also weaken considerably towards and

on the body chamber. The body chamber may also show slight egression of the

umbilical seam, resulting in scaphitoid uncoiling. This is the typical ‘kaffrariun’

stage (Figs 29-31).

With the exception of the embryonic stage, all these ontogenetic stages can

be observed in SAM-—PCZ6194 (Fig. 13), SAM-—93B/4 (Fig. 32), SAS Z1080

(Figs 33-34), and SAS Z1820 (Fig. 35).

However, part of the difficulty in unravelling the systematics of the genus

Placenticeras lies in the fact that these different ontogenetic changes occur at

‘T X ‘WUOUTeUIO [e19}e] JUSUTUIOId Jo YOR] INQ ‘IAR[O [eIWUDA puke SapoIOqnN} [eoT{Iquin

JUDUIWOId YUM a8e}S DNAUABOJUO wnDvaffoyqns, JesIdA, SuIMOYs UsUTIDNdSG ‘“66PY SVS ‘PO6I ‘Aspley wnuoiff[oy svsaouadvj[g ‘97 ‘314

‘

279

CRETACEOUS FAUNAS FROM SOUTH AFRICA

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

em D E

Fig. 27. Placenticeras kaffrarium Etheridge, 1904. A-B. SAM-PCZ6200. C-E. SAS Z682.

Two specimens with ‘subkaffrarium’ type of ornament to illustrate variation in whorl breadth and

outline. Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 281

Fig. 28. Placenticeras kaffrarium Etheridge, 1904. SAS Z1028/1. Specimen showing transition

from smooth ‘umkwelanense’ stage to ‘subkaffrarium’ stage with umbilical tubercles and ventral

clavi. X 1.

different rates, and stages are of different length in different individuals, or may

be omitted. This leads to the presence of adult individuals resembling—in terms

of ornament—the juveniles of others, while late phragmocone modifications

already occur in the very early developmental stages of some specimens (Fig. 36),

all at the same stratigraphic level (Figs 37-38). In rare cases an intermediate

ontogenetic stage can even be omitted. The best way of describing this variation

in ontogeny in combination with normal intraspecific variation is by illustrating

the different morphotypes. This is based mainly on the presence and/or stage of

appearance of the umbilical and lateral tubercles and crenulation of the venter.

2. Dimorphism

Distinct dimorphism can be distinguished in adults in these different morpho-

types (see p. 251 for criteria for recognizing maturity).

(a) In all adult specimens the venter becomes rounded and generally smooth on

the body chamber with no ventral clavi.

(b) In forms with strong ‘kaffrarium or ‘subkaffrarium’ type of ornament on the

phragmocone, umbilical tubercles migrate away from the umbilical wall and

weaken or disappear on the body chamber.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. Placenticeras kaffrarium Etheridge, 1904. SAS Z1520a. Specimen showing late

phragmocone and body chamber modifications. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 283

Fig. 30. Placenticeras kaffrarium Etheridge, 1904. NMB D1344. Specimen showing late

phragmocone and body chamber modifications. x 0,98.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 31. Placenticeras kaffrarium Etheridge, 1904. NMB D1344. Specimen showing late

phragmocone and body chamber modifications. X 0,75.

285

CRETACEOUS FAUNAS FROM SOUTH AFRICA

ive

ing SUCCESS

1904. SAM-93B/4. Specimen show

xa.

ontogenetic stages.

ium Etheridge

Fig. 32. Placenticeras kaffrar

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Placenticeras kaffrarium Etheridge, 1904. SAS Z1080. Macroconch showing successive

ontogenetic changes up to body chamber. xX 0,75.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 287

Fig. 34. Placenticeras kaffrarium Etheridge, 1904. SAS Z1080. Macroconch showing successive

ontogenetic changes up to body chamber. x 0,75.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Placenticeras kaffrarium Etheridge, 1904. SAS 1820. Microconch showing distinct

ontogenetic changes from ‘umkwelanense’ through ‘subkaffrarium’ to ‘kaffrarium’ stage. X 0,75.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 289

D e F

Fig. 36. Placenticeras kaffrarium Etheridge, 1904. A-—C. NMB D2845. D-F. NMB D941/48.

Both specimens at similar diameters to illustrate differing rates of ontogenetic change. The

upper specimen already has the mature ‘kaffrarium’ ornament, whereas the lower is still at the

‘umkwelanense’ stage. Both x 1,5.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

tat

= net,

nae

Fig. 37. Placenticeras kaffrarium Etheridge, 1904. SAS A419. Concretion from locality 72 to

illustrate the co-occurrence of ‘umkwelanense’ and ‘kaffrarium’ forms at the same stratigraphic

level. < i,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 291

Fig. 38. Placenticeras kaffrarium Etheridge, 1904. SAM, uncatalogued specimen. Concretion

from locality 72 to illustrate the co-occurrence of ‘umkwelanense’ and ‘kaffrarium’ forms at the

same stratigraphic level. Part of Peroniceras (Zuluiceras) at top left. Xx 0,5.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B Cc

Fig. 39. Placenticeras kaffrarium Etheridge, 1904. Whorl section of different specimens to

illustrate variation. A. NMB D1150, ‘kaffrarium’ type. B. NMB D1150, ‘sub-kaffrarium’ type.

C. NMB D1343, ‘umkwelanense’ type.

cone, weak umbilica | only, or umbilical and lateral ornament, appears on the

body chamber.

Dimorphism in the Zululand placenticeratids is restricted mainly to differ-

ences in size. In microconchs, which are more abundant in the present collection,

the last septum is formed at diameters between 110 and 120 mm, whereas

macroconchs show a wider range of maximum diameter, but generally form the

last septum at between 150 and 185 mm. The length of the body chamber is

between half and two-thirds of a whorl. Except for possibly one specimen,

SAS Z1081 (Fig. 14A), no complete aperture was found and apertural modifica-

tions, if present, are unknown.

A combination of differing rates of ontogeny and dimorphism allows

recognition of several morphotypes.

Type 1. In some specimens the shell never develops beyond the early smooth

‘umkwelanense’ stage and remains smooth throughout. At the end of the adult

phragmocone, and on the body chamber, however, the venter becomes distinctly

rounded, albeit very narrow in extreme cases, without an intermediate tubercu-

late stage. There is considerable variation in the whorl section, ranging from

extremely compressed, subtrigonal with flat flanks converging to a narrow venter,

e.g. SAM—PCZ6197/a (macroconch) (Figs 40-41) to broadly rounded ovoid, e.g.

SAM-PCZ6202 (macroconch) and NMB D1151/5 (microconch) (Figs 42—43)

(also Figs 44—-46B).

Type 2. In this form the greater part of the phragmocone remains smooth. Weak,

poorly developed umbilical tubercles only start appearing on the late adult

phragmocone or on the adult body chamber. Again, there is considerable

variation in the whorl section, as in type 1. NMB D941/33 (macroconch) (Fig. 47)

and SAS Z106Sc (microconch) (Fig. 48) are good examples (also Figs 49-50).

Type 3. The greater part of the phragmocone is smooth, and ornament only starts

appearing relatively late in ontogeny. On the adult body chamber, however,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 293

Fig. 40. Placenticeras kaffrarium Etheridge, 1904. SAM-—PCZ6197/a. Macroconch body

chamber and last few septa of specimen with type 1 ornament. Note the extremely compressed,

lanceolate whorl section and also the impression of the flat, entire venter of the inner

‘umkwelanense’ whorls. X 0,94.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41. Placenticeras kaffrarium Etheridge, 1904. SAM-—PCZ6197/a. Macroconch body

chamber and part of phragmocone with type 1 ornament. xX 0,94.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 295

Fig. 42. Placenticeras kaffrarium Etheridge, 1904. NMB D1151/5. Microconch with type 1

ornament. Note the adult features—slight scaphitoid uncoiling, outward slanting of the

umbilical wall and transition from flat to rounded venter. X 0,78.

ANNALS OF THE SOUTH AFRICAN MUSEUM

296

NMB D1151/5. Microconch with type 1

1904

kaffrarium Etheridge

ticeras

Fig. 43. Placen

ornament.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 297

Fig. 44. Placenticeras kaffrarium Etheridge, 1904. SAS Z1156. Macroconch with type 1

ornament. Note rounding of venter, outward-slanting umbilical wall and slight scaphitoid

uncoiling towards beginning of body chamber. xX 0,88.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 45. Placenticeras kaffrarium Etheridge, 1904. SAS Z1156. Macroconch with type 1

ornament. X 0,88.

‘IL x YOg ‘Joquieyo Apog dy} 0} dn

a8ejs ,asuaunjamyWn, YOOUS Zururej}oI “yUOUTeUIO | odA} YIM YOUOSOINR “EPEC GINN ‘A “Jequieyo Apog ay} UO puke a8e}s oUOCOUNse IY 93k] OY}

uo doyaaap Ajuo saposaqn) jeoriquin yoryM UT jUaWTRUIO Z odA} YIM YOUOSOIDIL “O89Z SVS “WV ‘PO6T ‘e8pHoyIA wnuvsffoy spsaoyuarv]d ‘Op B14

q V

299

CRETACEOUS FAUNAS FROM SOUTH AFRICA

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

oa ow,

Lore:

Fig. 47. Placenticeras kaffrarium Etheridge, 1904. NMB D941/33. Macroconch with type 2

ornament in which umbilical tubercles only start appearing on the late phragmocone stage or on

the body chamber. x 0,65.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 301

Fig. 48. Placenticeras kaffrarium Etheridge, 1904. SAS Z1065c. Microconch with type 2

ornament. X 0,88.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 49. Placenticeras kaffrarium Etheridge, 1904. NMB D941/15. Microconch with type 2

ornament. X l.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 303

Fig. 50. Placenticeras kaffrarium Etheridge, 1904. NMB D941/15. Microconch with type 2

ornament. xX 1.

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

DIAMETER (mm)

Fig. 51. Placenticeras kaffrarium Etheridge, 1904. Size distribution.

40 e e e

UMBILICAL DIAMETER (mm)

Oo 50 100 150 200 250

TOTAL DIAMETER (mm)

Fig. 52. Placenticeras kaffrarium Etheridge, 1904. Total diameter versus umbilical diameter.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 305

0,80

0,50

fo) 50 100 150 200 250

DIAMETER (mm)

Fig. 53. Placenticeras kaffrarium Etheridge, 1904. Total diameter versus Wb/Wh.

ornament becomes quite bold, with distinct umbilical and lateral ornament of

typical ‘kaffrarium’ type. The venter changes from flat and smooth to tuberculate

to rounded and smooth, or directly from flat and smooth to rounded and smooth

without an intermediate tuberculate stage. Examples are SAS Z1520g (macro-

conch) and NMB D1269 (macroconch, Fig. 56).

These three morphotypes are all basically variations on the ‘umkwelanense’

type of ornament, where the phragmocone remains predominantly smooth. In all

the remaining groups, lateral ornament appears at a relatively early ontogenetic

stage, and differences are found mainly in the relative strength of the umbilical

and lateral ornament, and, in consequence, the whorl section.

Type 4. Here umbilical tubercles and ventral clavi develop at a very early stage,

but no distinct lateral ribs or tubercles occur. The strength of the umbilical

tubercles varies considerably, and in some cases they are markedly spinose. The

umbilical tubercles generally weaken on the outer whorls and the flanks become

rounded. Simultaneously, the ventral clavi may disappear and the venter may be

flat and smooth as in ‘umkwelanense’ morphotypes—especially on internal

moulds—in association with weak umbilical tubercles, before becoming rounded.

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

UMBILICAL TUBERCLES

Fig. 54. Placenticeras kaffrarium Etheridge, 1904. Frequency of umbilical tubercles.

The whorl section is strongly subtrigonal with maximum whorl breadth at the site

of the umbilical tubercle, but breadth to height ratios vary widely. This is the

typical ‘subkaffrarium’ type of sculpture where umbilical ornament is dominant,

and the whorl section is distinctly subtrigonal. Typical examples are SAS Z201

(macroconch), SAS Z202 (macroconch) (Figs 57-58), NMB D941/2 (micro-

conch) (Fig. 59), NMBD1267 (microconch) (Figs 60-61), SAM-—PCZ6196

(microconch) (Figs 62-63), and NMB D925/3 (macroconch) (Figs 64-65) (see

also Figs 66-68).

Type 5. Here the whorl section is still distinctly subtrigonal but, in addition to

strong umbilical tubercles, strong lateral tubercles and strong ventral clavi occur,

all connected by prominent ribs. On the adult outer whorls the section becomes

rounded and ornament subdued. Examples are NMB D1271 (macroconch)

(Fig. 69), SAS Z1520a (macroconch) (Fig. 72), NMB D941/19 (?macroconch)

(Fig. 73); in some specimens, e.g. NMB D1269, D941/14 (Figs 70-71) fine, dense

ribbing occurs on the outer adult whorls.

Type 6. In this type the lateral tubercle can become much stronger than the

umbilical tubercle and the whorl section becomes polygonal at an early stage. In

relation to the other types, ornament on the adult body chamber is strong but

CRETACEOUS FAUNAS FROM SOUTH AFRICA 307

89 '10'11 12°13 '14'15 16 17 18 1920 21 = 25° 26-40 31-35 36-40

LATERAL TUBERCLES VENTRAL CLAVI

Fig. 55. Placenticeras kaffrarium Etheridge, 1904. A. Frequency of lateral tubercles. B. Fre-

quency of ventral clavi.

nevertheless weaker than on the phragmocone. This is the typical ‘kaffrariun’

type of ornament. Good examples are Z80 (microconch) (Figs 74-75), Z252

(macroconch) (Figs 76-77), Z1021 (macroconch) (Fig. 78), Z187 (microconch)

(Fig. 79) (see also Figs 80-81).

Type 7. A few specimens, e.g. PCZ6215 (Fig. 82) show a very strong increase in

ornament, passing rapidly from the smooth ‘umkwelanense’ to very robust

‘kaffrarium’ type of ornament (see also Figs 83-85).

3. Evolutionary changes

From the above description of the various morphological types encountered

in Zululand Coniacian representatives of Placenticeras, it is clear that the material

is extremely variable, and that we are dealing with but a single species.

Unfortunately no continuous exposures of the entire Coniacian Stage are

available in Zululand. However, from the preliminary stratigraphic framework

compiled by Kennedy & Klinger (1975), and partly revised for the Coniacian

Stage by Klinger & Kennedy (1984), Placenticeras occurs in the lower three

divisions of the Coniacian in Zululand, i.e. Coniacian I-III. In fact, Placenticeras

already occurs in the basal Pterotrigonia conglomerate of the St Lucia Formation.

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 56. Placenticeras kaffrarium Etheridge, 1904. NMB D1269. Part of body chamber of

macroconch with type 3 ornament. xX 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 309

Fig. 57. Placenticeras kaffrarium Etheridge, 1904. SAS Z202. Macroconch with type 4

ornament. X 0,72.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM ;

Fig. 58. Placenticeras kaffrarium Etheridge, 1904. SAS Z202. Macroconch with type 4

ornament. xX 0,72

’

CRETACEOUS FAUNAS FROM SOUTH AFRICA a1

Fig. 59. Placenticeras kaffrarium Etheridge, 1904. NMB D941/2. Microconch with type 4

ornament with umbilical spines weakening considerably on the outer whorls.

ANNALS OF THE SOUTH AFRICAN MUSEUM

312

Microconch with type 4

NMB D1267.

1 present at an ear

1904

ilical tubercles and ventral clav

dge

kaffrarium Etheri

lceras

Placent.

ornament with umb

Fig. 60.

ly stage, but no lateral

x 1

ornament.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 313

A B

Fig. 61. Placenticeras kaffrarium Etheridge, 1904. NMB D1267. Microconch with type 4

ornament. X 1,1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

314

ornament.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 315

A B

Fig. 63. Placenticeras kaffrarium Etheridge, 1904. SAM-PCZ6196. Microconch with type 4

ornament.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 64. Placenticeras kaffrarium Etheridge, 1904. NMB D925/3. Macroconch with type 4

ornament. Note the decrease in strength of the umbilical tubercles towards the end of the

phragmocone. X 0,85.

ST he eae 4 meted

CRETACEOUS FAUNAS FROM SOUTH AFRICA 317

Fig. 65. Placenticeras kaffrarium Etheridge, 1904. NMB D925/3. Macroconch with type 4

ornament. X 0,85.

ANNALS OF THE SOUTH AFRICAN MUSEUM

318

Fig. 66. Placenticeras kaffrarium Etheridge, 1904. NMB D1151. Macroconch with type 4

CUS.

ornament.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 319

Fig. 67. Placenticeras kaffrarium Etheridge, 1904. SAM-—PCZ6198. Impression of the inner

whorl of specimens with type 4 ornamentation to illustrate spinose nature of umbilical tubercles

and sigmoid lirae on the flanks. x 0,9.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM ;

Fig. 68. Placenticeras kaffrarium Etheridge, 1904. SAM-—PCZ6258. Impression of specimen

with type 4 ornament to illustrate extremely spinose umbilical tubercles on inner whorls and fine

lirae on flanks. xX 0,8.

Sak

ith type 5

i>)

1@)

so)

[a0]

< =

O ‘

ad —

[ae |

“ Ss

= Q

: a

a Ze

= nes

g Sx

f =

Z J 5

Zz, =e

: aE

i 6

iF)

Ras

Fig. 69. Placenticeras kaffrarium Ether

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 70. Placenticeras kaffrarium Etheridge, 1904. NMB D941/14. Macroconch with type 5

ornament and fine, dense ribbing on the outer whorls. ~X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 325

Fig. 71. Placenticeras kaffrarium Etheridge, 1904. NMB D941/14. Macroconch with type 5

ornament. Note the distinct ‘subkaffrarium’ inner whorls. X 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

324

Macroconch with type 5

SAS Z1520a

, 1904.

dge

ornament.

ium Ether

iceras kaffrar.

Placent

Pigs (72:

)

‘Tx ‘JugureuIo ¢ odA} YIM {YOUDOLDRY “6T/TP6 GINN ‘“PO6I ‘espueyI wnuvsfoy spsvoyuarvjd ‘¢L ‘314

323

CRETACEOUS FAUNAS FROM SOUTH AFRICA

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 74. Placenticeras kaffrarium Etheridge, 1904. SAS Z80. Microconch with type 6 ornament

with strong lateral ornament at an early stage, resulting in distinct polygonal whorl section. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 327

& B

Fig. 75. Placenticeras kaffrarium Etheridge, 1904. SAS Z80. Microconch with type 6 ornament

with strong lateral ornament at an early stage, resulting in distinct polygonal whorl section. xX 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

328

Macroconch with type 6

SAS 2252/2.

1904

Placenticeras kaffrarium Etheridge,

Fig. 76.

’

ornament

CRETACEOUS FAUNAS FROM SOUTH AFRICA

th type 6

SAS Z252/2. Macroconch wi

1904.

x 0,6.

idge,

kaffrarium Ether

lceras

Placent.

rip. 77.

ornament

oS)

wy)

co)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 78. Placenticeras kaffrarium Etheridge, 1904. SAS Z1021. Macroconch with type 6

ornament. xX 0,7.

CRETACEOUS FAUNAS FROM SOUTH AFRICA i io dl|

Fig. 79. Placenticeras kaffrarium Etheridge, 1904. SAS Z187. Microconch with typical type 6

ornament. Note scaphitoid uncoiling of the body chamber. xX 0,75.

ANNALS OF THE SOUTH AFRICAN MUSEUM

332

Placenticeras kaffrar

SAS Z1125. Microconch with type 6

ium Etheridge, 1904.

Fig. 80.

ornament.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 333

A ete

Fig. 81. Placenticeras kaffrarium Etheridge, 1904. NMB D1149/2. Microconch with type 6

ornament. X 0,80.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 82. Placenticeras kaffrarium Etheridge, 1904. SAS Z215. Microconch with type 7 orna-

ment. Note conical perforations on the bottom half of the shell. Possible signs of predation?

x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 335

Fig. 83. Placenticeras kaffrarium Etheridge, 1904. SAS Z890. Microconch with type 7 orna-

| ment with very strong, auricular lateral tubercles. X 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

ioe)

Microconch with type 7

SAS Z1181.

Etheridge, 1904.

lum

Placenticeras kaffrar

Fig. 84.

> Saal

ornament

CRETACEOUS FAUNAS FROM SOUTH AFRICA 337

A B

Fig. 85. Placenticeras kaffrarium Etheridge, 1904. NMBD1150. Microconch with type 7

ornament. Note the strong lateral ornament and polygonal whorl section. xX 1.

Fig. 86. Placenticeras kaffrarium Etheridge, 1904, uncatalogued. Whorl sections of three

Specimens, all from the same locality to illustrate variation in degree of inflation due to

intraspecific variation. X 1.

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 87. Placenticeras kaffrarium Etheridge, 1904. Whorl section of extremely inflated specimen

from the same locality as the specimens in Figure 86. X 1.

overlying Upper Cenomanian sediments at locality 60 at the Skoenberg, Zululand

(Fig. 20D). In the silts and sands overlying this basal conglomerate numerous

representatives of Placenticeras occur. The ‘umkwelanense’ and ‘subkaffrarium’

forms are abundant, the latter being very obvious in the assemblage by virtue of

the distinct subtrigonal whorl section and strong umbilical tuberculation. Forms

with lateral ornamentation are also known but none approach typical P. kaffra-

rium forms. In the slightly younger Placenticeras-bearing strata (Coniacian II)

along the lower reaches of the Mzinene River, at localities 71 and 72, and the

lower reaches of the Hluhluwe River, at localities 92 and 93, ‘umkwelanense’ and

‘kaffrarium’ forms dominate. In the exposures along the Msundusi River at

locality 145, the fauna seems similar to that of locality 60, but with elements of

‘kaffrarium’ also present. This indicates that the Zululand Placenticeras are part

CRETACEOUS FAUNAS FROM SOUTH AFRICA 339

Fig. 88. Placenticeras kaffrarium Etheridge, 1904. Whorl sections of two specimens, both from

the same locality as the specimens in Figures 86 and 87 to illustrate effects of intraspecific

variation on whorl section. X 1.

i. . oo

0 10

Fig. 89. Placenticeras kaffrarium Etheridge, 1904. SAS Z1442. Suture line of juvenile

specimen. Scale bar in mm.

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

1) a ie

0 10

Fig. 90. Placenticeras kaffrarium Etheridge, 1904. SAS A1446. Suture line of juvenile

specimen. Scale bar in mm.

Fig. 91. Placenticeras kaffrarium Etheridge, 1904. Unregistered specimen. Suture line. Scale

bar in mm.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 341

| i

I /

| I

| \

| 0 10

Fig. 92. Placenticeras kaffrarium Etheridge, 1904. Suture lines of SAS Z637 and NMB D941/4.

Scale bar in mm.

Py Ley Lp

0 10

Fig. 93. Placenticeras kaffrarium Etheridge, 1904. SAS Z939. Suture line. Scale bar in mm.

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

oy a Us

Ses ae

0 10

Fig. 94. Placenticeras kaffrarium Etheridge, 1904. Unregistered specimen. Suture line. Scale

bar in mm.

-_--—- oOo

ae ea i ee

CRETACEOUS FAUNAS FROM SOUTH AFRICA 343

Se a aa a aaa |

0 2

Fig. 96. Placenticeras kaffrarium Etheridge, 1904. NMB D941/7. Early suture. Scale bar

in mm.

0 2

Fig. 97. Placenticeras kaffrarium Etheridge, 1904. NMB D941/7. Early suture. Scale bar

in mm.

of a lineage in which early populations are dominated by ‘subkaffrarium’

morphotypes with distinct trigonal whorl section and strong umbilical tubercles,

succeeded by populations in which ‘kaffrarium’ forms, with polygonal whorl

section and lateral tubercles as strong or stronger than the umbilical ones

dominate. At both levels feebly ornamented ‘umkwelanense’ forms occur,

retaining the smooth early stage to varying diameters.

At best, subkaffrarium and kaffrarium could be separated as chronological

subspecies but given the presence of common morphotypes, even this is un-

necessary, although given a number of specimens it is possible to distinguish early

and late forms of the species. More important, the Zululand material shows

unequivocal evidence of progressive evolutionary change in a single lineage:

phyletic gradualism.

4. Intraspecific variation

Intraspecific variation per se is difficult to separate completely from the other

variable factors affecting differences in placenticeratid populations. Obvious

variation is reflected by reference to various morphotypes, but here differing rates

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

of ontogenetic development and, to a lesser degree, phylogenetic changes are also

involved. True intraspecific variation is best seen in different degrees of inflation

of the whorl section. A number of specimens from the same restricted strati-

graphic level (Figs 86-88) show the total spectrum of intraspecific variation in this

character—ranging from a compressed, flat-sided whorl section to an inflated,

trigonal whorl section with greatest width at the umbilical shoulder.

Unfortunately the material is not suitable for studying detailed sutural

ontogeny. Available sutures are shown in Figures 89-97 to show the variation.

Discussion

From the above description it is clear that all the Placenticeras from the

Coniacian of Zululand belong to a single extremely variable but phylogenetically

conservative species, for which we select the name Placenticeras kaffrarium. This

had already been suggested on earlier occasions (Klinger & Kennedy 1980b,

1980c), and is now confirmed. All the other names used for Zululand Placen-

ticeras, i.e. P. umkwelanense Etheridge, 1904, P. subkaffrarium Spath, 1921,

P. cf. subkaffrarium Spath, 1921, Placenticeras n. sp. aff. kaffrarium Venzo,

1936, merely represent different ontogenetic stages or intraspecific variants of

P. kaffrarium.

In view of the intraspecific variation, comparisons with material from other

regions are difficult. This is painfully obvious in the works of Boule et al. (1907),

Besairié (1930) and Venzo (1936); all identified feebly ornamented, Coniacian,

‘umkwelanense’ morphotypes with the smooth Campanian species Placenticeras

whitfieldi.

Forms similar to P. kaffrarium have been recorded from the Middle and

Upper Turonian of Madagascar under the names Proplacenticeras memoriae-

schloenbachi Laube & Bruder var. ambiloensis Collignon (1965a: 14, 16, pl. 381

(fig. 1646), pl. 382 (figs 1647-1648)), Parastantonoceras murphyi Collignon

(1965a: 17, pl. 382 (fig. 1649)), Proplacenticeras stantoni Hyatt var. fortior

Collignon (1965a: 19, pl. 383 (fig. 1652)) and Proplacenticeras orbignyi (Geinitz)

(Collignon 1965a: 20, pl. 383 (figs 1653-1654)). Nearly all these ‘species’ and

varieties can be matched in the material from the Coniacian of Zululand and we

regard them as synonyms, although the structure of Turonian Placenticeras

populations in this area remains unknown.

The Coniacian stratigraphy of Madagascar has recently been reviewed by the

authors (Klinger & Kennedy 1984), clarifying Collignon’s (19655) usage of the

terms Lower, Middle and Upper Coniacian. Collignon’s Lower and Middle

Coniacian zones of Peroniceras dravidicum and P. subtricarinatum, and of

Kossmaticeras theobaldi and Barroisiceras onilahyense, correspond approximately

to the second and third divisions of the Coniacian stage in Zululand, and the

Middle Coniacian of France, as defined by the authors.

Proplacenticeras aff. fritschi de Grossouvre var. eboroensis Collignon (1965b:

38, pl. 430 (figs 1780-1781)) from Collignon’s Middle Coniacian corresponds to

the typical Placenticeras kaffrarium (Type 2), with umbilical tubercles appearing

‘LX ‘ejosuy WoO ‘S76 ‘UOIYsNePY 1ayI0UI0L

spaouaovjg JO edhyojoy e4L “€ZS9-WIVS ‘PO6I ‘O8spluoyI wnuvsffoy spsaouuarvjid °86 ‘S14

345

CRETACEOUS FAUNAS FROM SOUTH AFRICA

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 99. Placenticeras ?kaffrarium Etheridge, 1904. The holotype of Placenticeras merenskyi

Haughton, 1930, from the Wanderfeld IV Beds near Bogenfels, South West Africa—Namibia.

ole

at a relatively late stage. Specimens such as Z680 from Zululand match this

variety perfectly.

Proplacenticeras sp. nov. (Venzo) aff. kaffrarium Etheridge (Collignon

1965b: 40, pl. 431 (fig. 1783)) appears to be a microconch of P. kaffrarium with

subdued outer flank ornament. Proplacenticeras satriense Collignon (19655: 40,

pl. 431 (fig. 1782)) closely matches Zululand P. kaffrarium (Type 4) macro-

conchs such as SAS Z202. Both these forms were recorded from the Upper

Coniacian of Madagascar, i.e. Middle Coniacian sensu Klinger & Kennedy 1983.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 347

Fig. 100. Placenticeras ?kaffrarium Etheridge, 1904. Holotype of Placenticeras merenskyi

Haughton, 1930, SAM-10569, from the Wanderfeld IV Beds near Bogenfels, South West

Africa—Namibia. x 1.

and must be viewed with some caution—especially as we do not have detailed

| descriptions of large populations. The effects of geographic isolation and perhaps

geographic subspecies development may merit valid taxonomic separation of

these forms.

: Comparisons with material from further afield than Madagascar are difficult

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 101. Placenticeras fritschi de Grossouvre, 1894. Cast of the lectotype (EMP collections);

the original of De Grossouvre (1894, pl. 5 (fig. 1)) from the Calcaires Durs de la Ribochére of

La Ribochére (Loir-et-Cher) France. xX 1.

The following named species belong to the group of Placenticeras kaffrarium:

Placenticeras reineckei Haughton (1925: 271, pl. 13 (figs 4-5)) (also 1926: 15, pl. 2

(figs 4-5)—French translation) (Fig. 98). Holotype by monotypy SAM-—6523

from south of the river Massango, Angola. In addition to the holotype we have

studied 10 specimens from San Nicolau, Angola (ex M. R. Cooper collection).

These specimens show the same range of ontogenetic and intraspecific variation

as P. kaffrarium. Haughton’s view that this species differs from P. subkaffrarium

CRETACEOUS FAUNAS FROM SOUTH AFRICA 349

Fig. 102. Placenticeras fritschi de Grossouvre, 1894. Cast of the lectotype (EMP collections);

the original of De Grossouvre (1894, pl. 5 (fig. 1)) from the Calcaires Durs de la Ribochére of

La Ribochére (Loir-et-Cher) France. X 1.

in having a smaller whorl breadth and greater number of ventral clavi is not

supported by the study of the additional material now available. The San Nicolau

locality was dated as Turonian—Lower Coniacian by Cooper (1978).

Placenticeras merenskyi Haughton (1930: 363, pl. 11 (figs 1-3)) (see also Klinger

1977, fig. 7) (Figs 99-100). Holotype, by monotypy SAM-—10569 from the Creta-

ceous (Cenomanian?) of Bogenfels, Namibia (see also McLachlan & McMillan

1979). This is a difficult species to interpret, being based on a single specimen in

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 103. Placenticeras fritschi de Grossouvre, 1894. FSR unregistered specimen from the

Calcaires Durs de la Ribochére of La Ribochére (Loir-et-Cher) France. X 1.

poor preservation. The flanks appear to be smooth and the venter flat and entire.

The umbilicus is partially occluded by matrix and encrusting ostreids but,

although probably wider than in most P. kaffrarium, is nevertheless within the

limits of the species. Umbilical ornament consists of about 12 small, pinched

tubercles, which is more than the average 7-8 in P. kaffrarium. Also, the

presence of umbilical tubercles in P. kaffrarium is normally associated with a

tuberculate venter. This places it closer to the Turonian material from Madagas-

car described by Collignon (1965a) as Proplacenticeras memoriaschloenbachi var.

ambiloensis which, according to present interpretation, corresponds to type 2 or

4 of P. kaffrarium.

Placenticeras memoriaschloenbachi Laube & Bruder (1887: 221, pl. 23 (fig. 1)).

Quite what this species actually is, remains problematic. The original highly

schematic figures show it to have 10 spirally elongated, umbilical tubercles, and a

narrow venter with sharp shoulders and no ventral clavi to a great diameter. This

CRETACEOUS FAUNAS FROM SOUTH AFRICA 35]

Fig. 104. Placenticeras fritschi de Grossouvre, 1894. SP unregistered specimen from Saujon, La

Pompiére (Charente-Maritime), France. x 1.

species is known from the Cenomanian to Turonian of west and Central Europe.

The combination of umbilical tubercles with a narrow, smooth, non-tuberculate

venter seem sufficient at present to separate it from the mainstream of

P. kaffrarium.

Placenticeras fritschi de Grossouvre (1894: 124, pl. 5 (figs 1-2), text-fig. 52)

(Figs 101-104) was recently reviewed by Kennedy (1984). The lectotype of the

species is the larger specimen figured by De Grossouvre (pl. 5 (fig. 1)). This

species has about ten small umbilical tubercles from an early stage, and twice as

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 105. Placenticeras semiornatum (d’Orbigny, 1850). A-B. Paralectotype. C-—D. Lecto-

type. Both from Touraine, France; horizon uncertain, but probably Santonian. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA S25

Fig. 106. Placenticeras semiornatum (d’Orbigny, 1850). MNHP 1986-27 from the Craie de

Villedieu of St Fraimbault (Sarthe) France. x 1.

many crescentic lateral ribs and an entire venter—apparently never developing

ventral clavi. Again, the combination of umbilical tubercles with non-tuberculate

venter may serve as a basis for separation from mainstream P. kaffrarium. Its

relationship to P. memoriaschloenbachi remains uncertain.

Placenticeras kharesmense Lahusen (1884: 134, pl. 2, pl. 3 (fig. 1)) (see also

Archanguelski 1916: 40, pl. 6 (fig. 5), pl. 7 (fig. 1)), originally described from the

Turonian of Turkestan, is similar to forms of P. kaffrarium with umbilical

tubercles only—i.e. ‘subkaffrarium’, and given more material may prove to be

conspecific, the name having priority over P. kaffrarium.

Placenticeras kysylcumense Archanguelski (1916: 45, pl. 7 (figs 4-7)) has an

inflated whorl section, strong umbilical and lateral tubercles and ventral clavi.

This is very difficult to distinguish from robust forms of P. kaffrarium and may be

conspecific. Kennedy (1984) suggested that it may be the microconch of

P. kharesmense.

Number of Specimens

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Placenticeras semiornatum (d’Orbigny, 1850), also recently reviewed by Kennedy

(1984) (Figs 105-106), can generally be distinguished from P. kaffrarium by

virtue of the absence of umbilical tubercles throughout (as far as can be

ascertained from the poor type material); it does, however, bear strong resem-

blance to smooth ‘umkwelanense’.

Beschtubeites beschtubensis Ijin (1975: 163, pl. 29 (figs 1-2), pl. 34 (fig. 7)) from

the Lower Turonian of the Amur River is a finely ornamented placenticeratid,

and may be distinguished from P. kaffrarium by virtue of the greater number of

lateral tubercles per whorl and ribs per whorl (25-29).

Beschtubeites kutuzovae Ijin (1975: 164, pl. 29 (fig. 3), pl. 34 (fig. 8)) is another

slightly coarser ribbed form with about 20—24 lateral tubercles per whorl.

Proplacenticeras kotzi Wjin (1975: 165, pl. 30 (figs 1-2), pl. 34 (fig. 9)), from the

Lower Coniacian of Central Asia, is similar to the forms of P. kaffrarium that

retain the smooth ‘umkwelanense’ stage up to the body chamber without

crenulation of the venter and only there develop a few feeble tubercles.

Proplacenticeras proplanum Iljin (1975: 166, pl. 31 (fig. 2), pl. 35 (fig. 10)), from

the Upper Coniacian of Central Asia, has 4-5 small umbilical tubercles and a

smooth venter, thus resembling Placenticeras fritschi de Grossouvre.

Indian material

Proplacenticeras rampuraensis Chiplonkar & Ghare (1977a: 109, figs 1-2,

6A, D) and Proplacenticeras spathi Chiplonkar & Ghare (1977a: 110, fig. 7E—-F),

from the Turonian Nodular Limestone at Rampura, also appear to be typical

P. kaffrarium. Proplacenticeras fritschi (de Grossouvre) of Chiplonkar & Ghare

(1977a: 113, fig. 7B-C), from the Turonian Limestone at Bagh, Mahakal,

Rampura, etc., is poorly preserved, but probably also belongs to P. kaffrarium.

Microconchs

Macroconchs

90 100 110 120 130 140 150 160 170 180 190 200 210 220 230

Total Diameter (mm)

Fig. 107. Histogram indicating frequency distribution of macroconchs and microconchs in

Placenticeras kaffrarium Etheridge, 1904.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 355

Proplacenticeras stantoni (Hyatt, 1903) Chiplonkar & Ghare (1977a: 114), from

the Turonian Nodular Limestone, north of Bagh Caves, Rampura and Mahakal,

was not figured and we cannot comment on it. Pseudoplacenticeras sp. cf.

P. milleri (von Hauer, 1866) (Chiplonkar & Ghare 1977a: 115, fig. 6E—-F) is from

the Turonian purple shales between Nimar Sandstone and Nodular Limestone at

Bagh caves. This differs from mainstream P. kaffrarium in only having feeble

ventral clavi and no lateral ornament.

All the specimens from the Narmada Valley Basin described by Chiplonkar

& Ghare (1977b) as Placentoscaphites are no more than adult placenticeratids,

some possibly deformed by compaction, with scaphitoid uncoiling of the body

chamber. Placentoscaphites helicus Chiplonkar & Ghare (1977b: 74, pl. 2

(figs 3—4)) with helical inner whorls seems to be either pathological or secondarily

deformed.

Occurrence

Turonian and Coniacian of Madagascar, Coniacian I-III of Zululand,

Alphard Group offshore Cape Province, Turonian of India?, questionably

Wanderfeld IV beds of Namibia, and “Turonian—Coniacian’ of Angola.

Genus Hoplitoplacenticeras Paulcke, 1907

Type species: Hoplites plasticus plasticus Paulcke, 1907 (p. 186) (ICZN

opinion 554, 1959; name no. 1629).

Discussion

In the most recent review of the Placenticeratidae, Kennedy & Wright (1983:

868) stated quite emphatically that “Iwo stocks (which) separated from Placenti-

ceras in the Santonian ... (Hoplitoplacenticeras and Metaplacenticeras) .. .

clearly deserve generic separation’. However, analysis of the type material of

German Hoplitoplacenticeras and Zululand material, as well as a review of the

literature, indicate that the intraspecific variation in these two groups is as great

as that found in normal Placenticeras, which not only clouds differences between

the two genera (Hoplitoplacenticeras and Metaplacenticeras) but also from the

mainstream Placenticeras.

Both genera were originally poorly defined. The combination Hoplites

Placenticeras was used by Paulcke (1907) in his description of an Upper

Cretaceous fauna from Cerro Cazador in southern Patagonia. According to

Paulcke this fauna could be traced back to Lower Cretaceous hoplitid ancestors

but, on the other hand, showed features that constituted transitions to Placenti-

ceras. On examining the fauna, Paulcke faced the question of where to place

it—in Hoplites or in Placenticeras—or whether to erect a new genus. Paulcke

favoured transitional genera, which exhibited features of both ancestral and

descendant genera. He suggested that, rather than to erect a meaningless new

name (‘bei denen sich kein Mensch etwas denken kann’), both names should be

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

used to indicate, from the nomenclature, from where and in which direction

development took place; that part of the morphology which was dominant in the

material should be emphasized (‘hervorheben’). If hoplitid features dominated

the name should be Hoplitoplacenticeras, whereas, if placenticeratid features

dominate, the name would be Hoplitoplacenticeras. Transitional forms between

Hoplites and Placenticeras would be designated Hoplitoplacenticeras. If neither

was dominant, neither part of the name would be italicized. This procedure was

contrary to the rules and the name Hoplitoplacenticeras was validated in 1959 in

ICZN opinion 554.

Wright (1957: L392) defined Hoplitoplacenticeras as ‘Rather evolute for

family; whorl section compressed to trapezoidal, venter flat; with prominent

variable coarse rounded or dense fine ribs, nearly straight, bearing 2 rows of

ventrolateral tubercles, of which outer row may be large and clavate; ribs cross

venter transversely and may have trace of siphonal tubercle. Genus is probably

too widely drawn’. Kennedy & Wright (1983: 870) repeated this diagnosis almost

word for word.

Kennedy (1986) provided a very detailed discussion of Hoplitoplacenticeras,

based on examination of Schliter’s (1867, 1871-1876) type material frony

Northern Germany. Apart from expanding on the generic diagnosis originally

provided by Wright (1957) and Kennedy & Wright (1983), Kennedy also

stabilized Hoplitoplacenticeras nomenclature by designating lectotypes for the

German species. Schliter referred the same specimens to different names on the

three occasions he discussed the group, and subsequent authors made type

designations that were ambiguous or invalid.

Examination of North German and Zululand material by both of us shows

that Hoplitoplacenticeras is as variable during the Upper Campanian to Upper

Maastrichtian as is Placenticeras during the Cenomanian to Maastrichtian.

As far as the shape of the whorl section is concerned (compressed or

inflated), the variation in Hoplitoplacenticeras matches that of Placenticeras—

compare e.g. H. costulosum (Schliiter, 1867) with H. dolbergense (Schliter,

1876). Ornament is as variable, ranging from virtually smooth in H. costulosum,

save for fine sigmoid striae and poorly developed umbilical tubercles, through

moderately ornate H. marroti (Coquand, 1859) with trituberculate ribbing, to

strongly ornamented H. dolbergense (Schliiter). The umbilical tubercles may

remain at the umbilical edge as in H. vari or H. costulosum. In H. lemfoerdense

an additional row of ventral tubercles develops. H. lemfoerdense thus has a set of

mid-lateral, inner and outer ventrolateral and ventral tubercles. This peculiar

arrangement of tubercles is also shown by the Upper Maastrichtian H. lafresnay-

anum (d’Orbigny, 1841), recently revised by Kennedy (1986). For this distinctive

group Kennedy (1986) has proposed the subgenus H. (Lemfoerdiceras), type

species H. lemfoerdense (Schliter, 1872).

The only apparent difference between Placenticeras and Hoplitoplacenti-

ceras, as far as ornament is concerned, seems to be the positioning of the external

clavi. In Placenticeras they always alternate on either side of the venter, whereas

CRETACEOUS FAUNAS FROM SOUTH AFRICA a5)

in Hoplitoplacenticeras they correspond in all species except H. coesfieldiense

schlueteri Michailov, 1951. (But Cobban (pers. comm.) says there are American

specimens of P. intercalare that have matched clavi and alternate clavi on the

same individual.) As far as this feature is concerned, Placenticeras polyopsis

appears transitional to Hoplitoplacenticeras in the adult stage—the position of the

clavi changing from alternating to only slightly offset. Another apparent differ-

ence seems to be that a smooth Placenticeras juvenile stage has never been found

in Hoplitoplacenticeras. Also, as far as we know, fine, thread-like ribbing has

never been observed in adult Placenticeras.

According to the diagnosis given by Wright (1957), there may be traces of

a siphonal row of tubercles. The only species with distinct siphonal tubercles

are Hoplitoplacenticeras awadi (Hassan 1971: 71, pl. 2 (figs 8-9)) and

H. kambysis (Quaas 1902: 309, pl. 29 (figs 8-11)) from the Maastrichtian of

Egypt. The latter species is in fact a scaphitid, according to Dr Z. Lewy (pers.

comm. 1985). Kennedy (1986) regarded both as scaphitine homoeomorphs of

Hoplitoplacenticeras.

As far as the suture lines are concerned, we have to rely mainly on the work

of Paulcke (1907). Fortunately the Patagonian material allowed him to examine

the sutures in detail through all stages of growth. What is of interest here is that

the sutures change according to ontogeny and morphology.

The sutures of Patagonian Hoplitoplacenticeras are not in the least pseudo-

ceratitic, neither in the juvenile nor in the adult stage (Fig. 110). They are

ammonitic throughout. In the juvenile stage the lateral lobe (L) is distinctly trifid;

the saddle E/L is broad and from a very early stage already shows a prominent

incision. The adult sutures vary considerably and, according to Paulcke, the

variation is largely determined by the whorl section and ornament. In those forms

with inflated whorl section and strong ornament, e.g. H. p. hauthali Paulcke, the

lateral lobe remains more or less symmetrically trifid and independent, and the

first lateral saddle (E/L) entire. In forms where placenticeratid features dominate,

1.e., typically compressed whorl section and involute coiling, e.g. H. p. laevis

Paulcke, the lateral lobe becomes remarkably asymmetrical until the ventral

prong of the lobe is almost as long as the median one, thus giving the lobe a nearly

bifid appearance, and the incision in the first lateral saddle (E/L) becomes very

prominent and nearly as deep as the prongs of the lateral lobe. Also, the saddles

and lobes tend to become constricted near their bases.

To summarize, Hoplitoplacenticeras can be distinguished from Placenticeras

mainly by virtue of the ventral tubercles corresponding and an apparent lack of a

smooth juvenile stage.

Distinction between Hoplitoplacenticeras and Metaplacenticeras in terms of

the type species is easy. Metaplacenticeras was introduced by Spath (1926: 79) in

cryptic manner ‘Metaplacenticeras gen. nov., proposed for Placenticeras pacificum

J. P. Smith (“Development and Phylogeny of Placenticeras”’. . . .) This last is

characterized by its falcoid ribbing and has a suture-line distinct from that of the

typical Placenticeras placenta (DeKay).’

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

The most comprehensive discussion on Metaplacenticeras was given by

Reeside (1926). As far as ornament is concerned, the type species has a tricarinate

venter to a diameter of 100 mm; in the adult stage the venter is narrow and flat

or slightly concave, bordered by finely nodose keels.

According to Reeside’s interpretation of the genus, it included species such

as Placenticeras californicum Anderson, P. sanctaemonicae Waring ‘and probably

also some of the forms included by Paulcke under the names Hoplites plasticus-

costatus and H. plasticus-laevis, from the Senonian of Patagonia’ (Reeside

1926: 2). Apart from the central keel, which may, in some cases, be only be

weakly developed, allocation of species such as Placenticeras californicum to

either genus seems feasible. Both genera have overlapping stratigraphic ranges in

the Upper Campanian in Japan (Matsumoto 1982a, 19826), but in North America

Hoplitoplacenticeras seems to occur slightly earlier than Metaplacenticeras. For

the present it seems advisable to retain the two genera, though fully aware of the

range of overlap.

Matsumoto (1984: 20-21) recently discussed the genus Metaplacenticeras and

alleged that a true adventive lobe (A) was present in the suture. This is totally at

variance with views held by Schindewolf (see above, p. 263). In addition,

Matsumoto doubted if both Metaplacenticeras and Hoplitoplacenticeras belonged

to Placenticeratidae—tentatively suggesting a pseudoschloenbachid origin for

Hoplitoplacenticeras.

Occurrence

Hoplitoplacenticeras occurs world-wide in the Upper Campanian, and ranges

to the Upper Maastrichtian in western Europe. It has been reported from

Germany (Schliiter 1867, 1872-1876; Giers 1964; Schmid & Ernst 1975), France

(De Grossouvre 1894), European Russia (Michailov 1951), Central Asia

(Atabekian & Khakhimov 1976), Don basin (Naidin 1974), Poland (Nowak

1909), Sweden (Odum 1953), Spain (Basse 1931), Israel (Chavan 1947), Libya

(Maxia 1943), Madagascar (Basse 1931; Collignon 1970), Egypt (Mahmoud

1955), Angola (Howarth 1965; SAM collections), Zululand (Kennedy & Klinger

1975), Natal (Kennedy & Klinger 1973), Patagonia (Paulcke 1907), Wyoming

(Cobban 1963), Texas (Young 1963), British Columbia (Usher 1952; Ward 1978),

and Japan (Matsumoto 1982a, 1982b, 1984).

Hoplitoplacenticeras howarthi Collignon, 1970

Figs 108-109

Hoplitoplacenticeras howarthi Collignon, 1970: 80, pl. 639 (fig. 2351).

Hoplitoplacenticeras plasticum plasticum Paulcke: Kennedy & Klinger, 1973: 102, pl. 5

(fig. 4a—e).

Type

Holotype, by original designation, the original of Collignon (1970: 80, pl. 639

(fig. 2351)) from the Upper Campanian of Mokotibe (Antsalova), Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 359

Fig. 108. Hoplitoplacenticeras howarthi Collignon, 1970. SAM-PCZ6586. Macroconch. x 1.

Material

SAS Z646, NMB D1314, SAM-—PCZ6586-—6587, all from locality 110, cliff

and foreshore section at the south-western tip of the Nibela Peninsula, St Lucia

Formation, Campanian III. University of Natal specimens 6556, from Somtseu

Road, Durban.

Dimensions

D Wb Wh Wb/Wh Gs Ut Vt

PCZ6586 107 39 (36,4) 49 (45,8) 0,79 22 (20,6) it —

D1314 70 — 29 (41,4) — 20 (28,6) 8 —

Description

The available specimens show virtually all the stages of ornamentation except

those of the early embryonic stage. On the inner whorls of Z646 (Fig. 109) the

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 109. Hoplitoplacenticeras howarthi Collignon, 1970. SAS Z646. Microconch. x 1.

very early, poorly developed Placenticeras-like ornament can be seen, soon to be

followed by the very coarse ornament of the greater part of the phragmocone.

This consists of low, single and bifurcating ribs. They arise on the umbilical wall,

giving rise to obliquely elongated tubercles from which they either bifurcate or

cross the flanks undivided. The ribs are rather low and sinuous on the flanks and

some show incipient looping. Near the ventrolateral shoulder they give rise to a

ventrolateral clavus, curve slightly forward and end in outer ventral clavi. The

ventrolateral and ventral clavi correspond on either side of the venter. This type

of ornament is present in all four specimens, but that of D1314 (Fig. 14B) is most

prominent with near horn-like ventrolateral tubercles on the outer whorl.

PCZ6586 (Fig. 108) shows the adult ornament of the body chamber. Here the

prominent lateral and ventral ornament of the phragmocone is replaced by fine,

thread-like lirae, and the venter becomes more or less rounded. Part of the suture

is exposed in PCZ6587.

Discussion

The Zululand material closely matches the Madagascan holotype at the same

diameter. Given the wide variation in placenticeratids, we doubt that the seven

other species described from the same locality by Collignon are anything more

than intraspecific variants of a single species.

The small specimen from the subsurface deposits of Durban originally

described as H. plasticum plasticum (Kennedy & Klinger 1973: 102, pl. 5

(figs 4a—e)) matches the Zululand material and is best referred to H. howarthi.

Occurrence

Upper Campanian of Zululand, Natal and Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 361

Fig. 110. Suture ontogeny of Hoplitoplacenticeras. Composite drawing. (After Paulcke 1907,

text-figs 10a, 5, 10b-c, 8, 12, 25, bottom to top.)

THE SYSTEMATIC POSITION OF THE GENUS HYPENGONOCERAS

Genus Hypengonoceras Spath, 1922

Type species. Placenticeras warthi Kossmat (1895: 176, pl. 20 (fig. 8))

(= Ammonites Orbignyanus Stoliczka (non Geinitz) (1864: 92, pl. 48 (fig. 2))

from the Lower Utatur Group (Albian) of southern India, by the original

designation of Spath (1922: 112).

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Narrowly umbilicate, shell compressed; flanks generally little rounded.

Ornament consists of low falcoid ribs, often very weak and only visible under

oblique lighting. Umbilical tubercles may be present but are never very promi-

nent. Venter flat to concave, with alternating ventral clavi at least at some stage.

Dimorphism pronounced, but mainly restricted to differences in size. Macro-

conchs may reach diameters up to half a metre; microconchs up to 100 mm.

Suture variable; in typical forms some saddles are little divided and ‘pincer-like’

with bifid folioles; in others as incised as in normal placenticeratid pattern.

Discussion

The genus Hypengonoceras was introduced in characteristically brief style by

Spath (1922: 112) as follows: ‘The Engonoceratidae include Hypengonoceras,

nov., proposed for the Albian ‘“‘Placenticeras’’ (type: Pl. warthi, Kossmat,

‘“Untersuch. tb. d. Siidind. Kreidef.,” loc. cit., 1895 p. 80, pl. xx, fig. 8; and

STOLICZKA, loc. cit., p. 92, pl. xlviii, fig. 2) = group IV of Placenticeras in

VREDENBURG (‘“‘Amm. of the Bagh Beds,” Rec. Geol. Surv. India,

vol. xxxvi, pt. 2, 1907, p. 120). They are clearly derivatives of Hoplitida, and

have no direct relationship either with the Mammitid Placenticeras or with

Hoplitoplacenticeras (so-called ‘‘Hoplites’’) of the Senonian.’

Correct interpretation of the genus, as well as its systematic position, is —

disputed.

In terms of Vredenburg’s (1907) original diagnosis of his ‘Section IV’ (of

Placenticeras), for which Spath created the genus Hypengonoceras, only the type

species belongs here. This section is ‘Compressed; smooth, outermost subsidiary

saddle split into two portions by a deep adventitious lobe, so that the external

saddle appears divided into four portions instead of three; the suture has

numerous inflections, but is very slightly frilled, the summit of the saddles being

almost devoid of marginals with the exception of a deep narrow median notch’

(Vredenburg 1907: 119).

Adkins (1928: 260) referred Hypengonoceras to the family Engonocerati-

dae, but Spath later (1931: 340) referred the genus to the Placenticeratidae

without stating any reasons. Roman (1938: 869) put the genus in the family

Placenticeratidae.

Casey (1960: 208) stated of Hypengonoceras ‘With its simple, pincer-like

endings to the saddles, Hypengonoceras shows great resemblance to the Engono-

ceratids Knemiceras and Parengonoceras of Lower and Middle Albian age,

though it exhibits the more complex fragmentation of external saddle character-

istic of the Placenticeratidae . . . nothing is known, however, to bridge the gap

between Hypengonoceras and the rather diverse forms that comprise the Upper

Cretaceous Placenticeratidae. . . . It is probable that both Hypengonoceras and

Hengestites are forerunners rather than direct ancestors of the Upper Cretaceous

Placenticeratidae.’” Even Schindewolf (1967: 744) casually referred Hypengono-

CRETACEOUS FAUNAS FROM SOUTH AFRICA 363

ceras to the Placenticeratidae in suggesting that the sutural ontogeny of the genus

probably is similar to that of Metaplacenticeras.

Kossmat’s (1895: 170 (74)) observation that A. warthi differs from the other

placenticeratids in that the external saddle is subdivided into four instead of three

auxiliary saddles which slope gently down to the first lateral lobe seems to have

received little subsequent attention.

Up to that stage, Hypengonoceras was interpreted in terms of the type

species H. warthi, known from the Upper Albian of southern India (Stoliczka

1865; Kossmat 1895) and possibly Madagascar (Boule et al. 1907) only, and the

diagnosis provided by Vredenburg (1907), Wright (1957) and Casey (1960)

regarding the pincer-like saddles and generally simple suture sufficed. However,

since then, several other species have been referred to Hypengonoceras that blur

the apparent simple image of the genus. Hypengonoceras decaryi Collignon (1963:

128, pl. 291 (fig. 1269)) is known from the Upper Albian, Zone of Pervinquieria

inflata of Madagascar, and was also recorded from Mozambique by Forster (1975:

216, pl. 10 (fig. 7), text-fig. 64) and is now known from Zululand (see below).

This is a very large species; the diameter of the wholly septate holotype is

240 mm. Ornament consists of low, rib-like swellings, and, according to Forster,

8 to 10 prorsiradiate umbilical bullae per whorl. It retains the clavate venter to a

very large diameter, before eventually becoming smooth. What is noteworthy, is

that the saddles in the suture line are by no means as pincer-like as in the type

species (Fig. 17). In fact, were it not known that the species occurs in the Albian,

it could be regarded as a normal ‘Proplacenticeras’. Collignon (1963: 126, pl. 290

(fig. 1268)) in fact described another species from the same locality and strati-

graphic level (Mont Raynaud, Diego-Suarez) as Proplacenticeras rerati. The

species is based on a similarly large holotype, still septate at 240 mm diameter,

with low ribbing originating from weak, conical umbilical tubercles. The venter

also shows the same transition from tuberculate with long clavi to smooth. The

suture is very little incised. Apart from differences in umbilical diameter and

ornament between the two species, they are well within the range of intraspecific

variation in placenticeratids. They are probably conspecific or at least congeneric.

Three more species were described by Collignon (1966) from the Upper

Albian Zone of Dipoloceras cristatum of Tarfaya. Hypengonoceras chouberti

Collignon (1966: 17, pl. 4 (figs 1-1la, 2—2a), pl. 5 (fig. 127)) is another enormous

species. The largest fragment figured by Collignon (1966, pl. 5) has a whorl height

of 137 mm. The inner whorls of the species are coarsely ornamented, but it

becomes smooth on the outer whorls. Hypengonoceras fauremuretae Collignon

(1966: 18, pl. 6 (fig. 1-1a, 2—2a, 3-3a, 4~4a)) is virtually smooth throughout as

compared to H. chouberti and has well-developed ventral carinae with tiny clavi.

The sutures, however, especially the lobes are typically rounded and splayed like

the ‘fingers of a glove’. Hypengonoceras tarfayense Collignon (1966: 19, pl. 7

(fig. 1-1a)) is another gigantic species with measured maximum diameter (still

septate) of 236 mm. It has delicate ventral clavi which eventually disappear on the

phragmocone, resulting in a smooth, rounded venter. Lateral ornament consists

364 ANNALS OF THE SOUTH AFRICAN MUSEUM

of fine, dense costules only. The suture line is poorly preserved, but seems to have

rather massive rectangular saddles and narrow lobes.

Arias & Wiedmann (1977: 9, fig. 5) described Hypengonoceras ibericum

from the Albian Utrillas Beds of south-eastern Spain. This is very strongly

ornamented form with 10 prorsiradiate to sinusoidal ribs, each originating at a

small umbilical tubercle, bearing a lateral tubercle and ending in a ventrolateral

tubercle, at which point they bifurcate, and eventually terminate in external clavi

on the venter. Quadituberculate ornament is hitherto unknown in Hypengono-

ceras—and we doubt if H. ibericum is in fact correctly referred to Hypengono-

ceras, and suggest it is probably a Knemiceras. Hypengonoceras ibericum apart,

Hypengonoceras is now no longer as simple to interpret as initially conceived.

Two questions have to be answered: Is Hypengonoceras a member of the

Engonoceratidae or of the Placenticeratidae? If the latter, what is its relationship

with the early placenticeratids ‘Karamaites’ and Hengestites?

Engonoceratidae and Placenticeratidae are generally distinguished by the

simplified, ceratitic saddles of Engonoceratidae, as compared to finely incised

saddles and lobes of Placenticeratidae. As far as this criterion is concerned

Hypengonoceras is transitional between Engonoceratidae and Placenticeratidae

—but then so is Parengonoceras Spath, 1924, which has always unquestionably

been regarded as a member of the Engonoceratidae. Schindewolf (1967: 745) has

argued that a reduction of a placenticeratid suture to that of an engonoceratid is

possible, but that the reverse (if Hypengonoceras were to be referred to

Placenticeratidae) is less likely, though possible. Thus, as far as the general

outline of the saddles is concerned, Hypengonoceras could be referred to either

family, though the majority of known specimens would seem better placed in

Engonoceratidae.

As far as details of the suture line are concerned, the first significant (and up

to now only) attempt at trying to establish the systematic position of Hypengono-

ceras, was by Renz (1970: 1028) in an introduction to the genus Parengonoceras.

Freely translated Renz stated that ‘The suture lines seem to be of significant

importance in distinguishing between Parengonoceras and Hypengonoceras.

Reference is made to two illustrations of external sutures: the one of the holotype,

and a second of a fragment from the Cenomanian of Mont Raynaud in

Madagascar (Boule et al. 1907, text-fig. 26, p. 46). The external saddle is

characteristic, in being clearly narrower than in Parengonoceras, and is divided

into two halves by a deep incision, which could correspond to a third adventive

lobe. The question of whether we are here dealing with real adventive lobes or

only with subdivisions (Teilprodukte) of the lateral lobe, will be significant for

allocation of this group to Engonoceratidae or Placenticeratidae.’ The importance

of seeking real grounds for separating Hypengonoceras from Parengonoceras

becomes even more obvious in Renz’s (1970) description of the Andean repre-

sentatives of Parengonoceras. Parengonoceras discoides Renz, 1970, is described

as being externally similar to Hypengonoceras fauremuretae Collignon, 1963

(Renz 1970: 1036); Parengonoceras barbacoense Renz, 1970, was initially (Renz

CRETACEOUS FAUNAS FROM SOUTH AFRICA 365

1968: 626) identified as a Hypengonoceras, and later (1970: 1044) compared with

Hypengonoceras warthi as far as the narrow bifid saddles are concerned, and with

Hypengonoceras decaryi Collignon and H. fauremuretae Collignon as far as the

lack of ornament.

Despite the (superficial?) similarity between Parengonoceras and Hypen-

gonoceras, there was never any doubt about the systematic placing of the former

genus. Again we suspect that this systematic grouping had been influenced more

by stratigraphy than natural affinities. Being of Lower to Middle Albian age,

Parengonoceras was more naturally placed in the predominantly Lower Cre-

taceous Engonoceratidae, rather than the predominantly Upper Cretaceous

Placenticeratidae. As far as the sutural ontogeny is concerned, there is little doubt

that Parengonoceras has two true adventive lobes, and that the lateral lobe

remains simple (Fig. 18). The lateral lobe (L) and the adjacent adventive lobe

(A,) are nearly equal in size, and slightly deeper than the flanking U, and A,

lobes, giving the ventral part of the external suture a bow-like sagging appear-

ance, quite unlike the typical sagging zig-zag outline of typical Placenticeratidae.

In addition, the lobes are splayed and digitate—like the fingers of a glove. No

detailed ontogenetic studies have as yet been undertaken on Hypengonoceras,

neither is the available material from Zululand suitably preserved for such

studies. As far as the general outline of the external suture of H. warthi is

concerned, however, apart from the width of the lateral saddle, it shows far

greater similarity to Parengonoceras than to Placenticeras. Also, the relative

proportions of the umbilical lobes, U, and U; (‘fourth and fifth lateral lobes’) is

similar to that of later (Turonian and younger) ‘Proplacenticeras’ rather than that

of early “Karamaites’. Thus on these grounds, Hypengonoceras can be clearly

ruled out from the direct line of ancestry of Placenticeras. Casey (1960: 208) had

already suggested that Hypengonoceras and Hengestites be regarded as forerun-

ners, rather than direct ancestors of the Upper Cretaceous Placenticeratidae.

Casey (1960: 208), however, reiterated the accepted view of Hypengonoceras

bearing strong resemblance to Engonoceratidae, and considered derivation of

Hypengonoceras from Parengonoceras or some allied member of the Engono-

ceratidae probable. Recent data (e.g. Mirzoev 1967; Michailova 1974, 1978) have

unequivocally shown the origin of early Placenticeras (‘Karamaites’) to lie in the

Hoplitidae, and not in the Engonoceratidae. These data would then suggest that

Hypengonoceras would be better treated as a late Upper Albian member of the

Engonoceratidae, rather than as an early placenticeratid of unknown affinities, as

tentatively suggested in Kennedy & Wright’s (1983, text-fig. 5) phylogenetic

diagram.

Occurrence

Albian of India (Stoliczka 1863-1865; Kossmat 1895), Madagascar (Collig-

non 1963; Boule et al. 1907), Mozambique (Forster 1975), Zululand (Kennedy &

Klinger 1975), Israel (Lewy 1981), Morocco (Collignon 1966), and Saghalin

(Matsumoto 1942).

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hypengonoceras decaryi Collignon, 1963

Figs 111-126

Placenticeras Warthi Boule, Lemoine & Thévenin, 1907 [non Kossmat]: 26 (46), pl: 12. (3)

(fig. 6—6a).

Hypengonoceras decaryi Collignon, 1963: 128, pl. 291 (fig. 1269). Forster, 1975: 216, pl. 10

(fig. 7), text-fig. 64.

Type

Holotype, by original designation, is the specimen figured by Collignon

(1963, pl. 291 (fig. 1269)) from locality 92-5, Mont Raynaud (Diego Suarez),

Madagascar, Upper Albian, Pervinquieria inflata Zone.

Material

Nine specimens, SAM-—PCZ6272-4, Z6277a—b, Z6278-6279, BMNH-

C81264—-81265, all from locality 179, sisal fields north of the Msundusi River

around the pumping station 2 100 m SSW of Ndumu Store, Ndumu, northern

Zululand. Recent excavations for a new road have exposed tens of metres of this

section. Mzinene Formation, Albian IV.

Dimensions

Specimen D Wb Wh Wb/Wh U

PCZ6273 240,0 70,0 (29,2) 141,0 (58,7) 0,50 12 (5,0)

PCZ6272 288,0 73,0°(26;0). 157,0 64,5} 0,48 11,0 (3,8)

PCZ6274 at 316,0 85,0 (26,9) 185,0 (58,4) 0,46 —

at 88,0 24,5 (27,8) 53,0 (60,2) 0,46 —

at Pe 15,0 (26,3) 31,5 G48) 0,48 —

at 37,0 11,0 (29,7) 24,0 (64,9) 0,46 —

at DF 7,3 (32,4) 11,4 G0) 0,64 —

Holotype 240 80 (33) 135 (56) 0,59 27 (11)

Forster (1975) 110 28 (25) 57 (52) 0,59 8 (7)

BMNH-C81264 — 184,0 — 101,0 (55,0) — 9,0 (5,0)

PCZ6277a e129 — — — —

PCZ6277b c. 160 — — — ee

Description

Markedly dimorphic; macroconchs still septate at diameters between 200 and

nearly 300 mm, and microconchs septate to a diameter of 100 mm. Macroconchs

include the holotype, PCZ6262-—6274, and C81264, whereas microconchs include

the specimen figured by Forster (1975, pl. 10 (fig. 7), text-fig. 64) and

PCZ6277a-b.

Very involute, with a tiny deep umbilicus comprising on average 5 per cent

of the total diameter. The umbilical seam is characteristically undercut on the

mould, with a rounded, convex umbilical wall, even on the body chamber.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 367

Fig. 111. Hypengonoceras decaryi Collignon, 1963. SAM-PCZ6272. Macroconch. Note

persistence of ventral clavi to relatively large diameter. x 0,65.

ANNALS OF THE SOUTH AFRICAN MUSEUM

368

PCZ6272. Macroconch. X 0,65.

, 1963. SAM-

ignon

Hypengonoceras decaryi Colli

Fig. 112

369

CRETACEOUS FAUNAS FROM SOUTH AFRICA

«1;

Macroconch.

PCZ6273

1963. SAM-—

i Collignon

Fig. 113. Hypengonoceras decary

ANNALS OF THE SOUTH AFRICAN MUSEUM

370

371

57.

x 0

icroconch

PCZ6277

, 1963. SAM-

ignon

CRETACEOUS FAUNAS FROM SOUTH AFRICA

i Coll

Fig. 115. Hypengonoceras decary

372 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 116. Hypengonoceras decaryi Collignon, 1963. SAM-—PCZ6274. Macroconch. xX 0,56.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 373

Fig. 117. Hypengonoceras decaryi Collignon, 1963. SAM-—PCZ6274. Macroconch. x 0,56.

ANNALS OF THE SOUTH AFRICAN MUSEUM

374

ree ha

, 1963. SAM-—PCZ6276. Macroconch. xX 0,9.

ignon

Hypengonoceras decaryi Coll

Fig. 118.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 375

Fig. 119. Hypengonoceras decaryi Collignon, 1963. SAM-—PCZ6376. Macroconch. x 0,9.

ANNALS OF THE SOUTH AFRICAN MUSEUM

376

, 1963. SAM-—PCZ6275. Microconch.

Hypengonoceras decaryi Collignon

Fig. 120

i>)

ie)

(S)

—

faa] .

< LV a)

g 3

fs >

< a

2) (7p)

a .

= (oe)

O &

~% 5 onl

fy, Lal

y :

2 &

=) =

jo)

= S

n >

eo)

iy <

Fig. 121. Hypengonoceras decary

378 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 122. Hypengonoceras decaryi Collignon, 1963. BMNH-C81264. Macroconch. x 0,9.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 379

Fig. 123. Hypengonoceras decaryi Collignon, 1963. BMNH-C81264. Macroconch. x 0,9.

380 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 124. Hypengonoceras decaryi Collignon, 1963. SAM-—PCZ6278. Inner whorls of macro-

conch showing position of umbilical tubercle. x 1.

Fig. 125. Hypengonoceras decaryi Collignon, 1963. BMNH, unregistered specimen. Suture

line. Scale bar in mm.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 381

Fig. 126. Hypengonoceras decaryi Collignon, 1963. SAM-PCZ6273. Suture line. x 1.

The whorls are compressed, tapering to a very narrow, flattened venter.

Ontogenetic change in whorl section is best seen in PCZ6278 (Fig. 124) and

PCZ6274 (Figs 116-117). During the ontogeny the maximum whorl breadth

migrates from near the umbilical edge to near the dorsal third of the flanks.

The venter is narrow, initially rounded in the very early (embryonic) stages

of growth, but then becomes concave and eventually flat on the greater part of the

phragmocone, and finally slightly rounded on the body chamber.

At first glance the species appears completely devoid of lateral ornament,

save fine, prorsiradiate flexuous striae. However, closer examination under

oblique illumination shows obscure, low folds on the flanks of the phragmocone,

especially on PCZ6272 (Figs 111-112). Examination of the inner whorls of

PCZ6278 (Fig. 124) shows the impression of about four low, rounded umbilical

tubercles per half whorl on the umbilical plug. The umbilical tubercles may

possibly be associated with stronger ribbing, but we lack material with shell to

substantiate this. This stage with umbilical tubercles persists to a diameter of

about 75 mm in PCZ6278.

The body chamber is at least half a whorl in length. The sutures are extremely

variable. Unfortunately the early sutural development could not be determined,

but the adult suture varies from specimen to specimen at the same diameters. The

lobes in PCZ6273 (Fig. 113) are rounded, with only minute central incisions—vir-

tually ceratitic—whereas the sutures of PCZ6272 (Fig. 111) are highly denticulate

and though perhaps not typically placenticeratid, certainly not ‘pincer-like

Hypengonoceratid’.

382 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 127. Suture line of Hypengonoceras chouberti Collignon, 1966. (After Collignon 1966,

pl. 5.)

Discussion

The Zululand material differs from Collignon’s (1963) holotype in generally

having a much smaller umbilicus, but is similar in having a very compressed,

trigonal whorl section and virtually smooth flanks. Given the range of intraspecific

variation in other ammonites, these differences are rather trivial. The species is of

great interest in demonstrating dimorphism in Hypengonoceras.

In view of the considerable intraspecific variation, comparison with other

described species is difficult: H. warthi (Kossmat) has low, broad, falcoid ribs at

diameters comparable to the Zululand material; H. chouberti Collignon is

smooth, more compressed, with well-developed keels on the ventral shoulders

bearing tiny clavi; H. tarfayense Collignon is a smooth, slender species, with a

very narrow venter and tiny umbilicus and an ornament of fine dense costules on

the flank. As far as is known, none of these species has umbilical tubercles at any

stage of growth, thus differing from the Zululand material.

Occurrence

Upper Albian of Zululand, Mozambique and Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 383

SPECULATIONS ON INTRASPECIFIC VARIATION IN

PLACENTICERATIDAE

Description of the Zululand faunas and work on large collections of

placenticeratids from other areas has clearly demonstrated the considerable

extent of intraspecific variation (e.g. Hyatt 1903; Reeside 1927b; Wolleben 1967;

Kennedy & Wright 1983). This raises the question why this intraspecific variation

is so widespread in the Placenticeratidae and, for that matter, most of the other

known Cretaceous narrowly umbilicate, compressed taxa, e.g. Engonoceratidae,

Pseudoschloenbachia (Collignon 1969; unpublished data Zululand), Eulopho-

ceras (unpublished data Zululand), Submortoniceras (Klinger & Kennedy 1980a),

etc.

Arnould-Saget (1956) attempted to explain the extreme intraspecific varia-

tion in Knemiceras from southern Tunisia. Here both ornament and suture lines

are very variable. According to her, the variations are individual and irrational;

their systematic study is sterile from an evolutionary point of view, and variation

is ascribed to palaeobiological causes. The Tunisian Knemiceras lived in a stable

environment in the neritic zone at the margins of the old Saharan mainland,

without important detritus deposits and without violent pertubation. These

optimum conditions did not create any need for adaptation; the life potential of

Knemiceras wasted itself in a useless and directionless polymorphism, but also

underlines the mode of life of these animals—more benthonic than neritic! Thus

Arnould-Saget.

In Zululand, Placenticeras kaffrarium occurs in a transgressive environment,

following regression and/or non-deposition and erosion during the Upper Ceno-

manian and Turonian. Associated ammonite faunas consist mainly of hetero-

morphs, e.g. Bostrychoceras, Scaphites, Allocrioceras, abundant Baculites, Pseud-

oxybeloceras, and evolute forms of Peroniceras—P. (P.) lepeei and P. (P.)

tridorsatum, and ornate Forresteria. Apart from the absence of complete aper-

tures in the placenticeratids, preservation of all these forms is superb, often with

the original nacre present. This all is indicative of a near-shore, protected

environment with absence of strong currents and wave action but, nevertheless,

normal salinity.

As we have demonstrated, coiling in most placenticeratids remains virtually

unchanged throughout their evolutionary history from the Cenomanian to the

Maastrichtian, 1.e. narrowly umbilicate compressed. With rare exceptions (e.g.

the Jurassic Amaltheidae—Howarth 1958; Kennedy 1977) oxyconic coiling seems

to be a unidirectional evolutionary ‘cul-de-sac’ (e.g. Submortoniceras—Klinger &

Kennedy 1980a; Peroniceras (Zuluites)—Klinger & Kennedy 1984). The only

noticeable, albeit slow and gradual trend in most of the Placenticeratidae, is the

acquisition of lateral ornament at the expense of umbilical ornament.

384 ANNALS OF THE SOUTH AFRICAN MUSEUM

Instead of regarding the variation in Placenticeras as useless and directionless

and coupled to a semi-benthonic mode of life, as suggested by Arnould-Saget

(1956), Chamberlain & Westermann (1976) suggested that the ornamentation in

ammonite shell morphotypes like those of Placenticeras may in fact have been

related to an active, nektonic mode of life, and to have been of hydrodynamic

significance. There is thus a distinct relationship between lateral ornament and

shell diameter, especially in compressed forms. To minimize drag, compressed

species with adult diameters of about 100 cm should become progressively

smoother during growth. The juveniles would be favoured if they had ornamented

shells. In constrast, the opposite trend would be of advantage in species with adult

diameters of 10 to 20 cm. This is what is seen in the most common form of

dimorphism in Placenticeras. The macroconchs are generally much more weakly

ornamented than the microconchs on the outer whorls. This does not, however,

explain in functional terms why some forms retain the smooth ‘umkwelanense’-

type of ornament up to the adult stage, whereas others are already ornate at small

diameters. It is also interesting to note that, according to Chamberlain &

Westermann (1976), all drag-producing ornament is located on the flanks.

Ornament on the umbilicus has little drag-producing effect. This may be perhaps

why umbilical ornament is gradually diminished in some placenticeratids, or tends

to migrate outwards during growth, thus producing extra drag.

LIST OF DESCRIBED SPECIES REFERRED TO PLACENTICERATIDAE

In view of the wide intraspecific variation shown by placenticeratid species,

detailed synonomies are difficult to compile, except where large populations are

available. We merely list species described. Lectotypes are designated where

necessary. These are indicated by *.

Genus Placenticeras

Placenticeras aktaschensis (Iljin, 1975) (p. 159, pl. 32 (figs 2-3), pl. 33 (fig. 4)).

Cenomanian of Central Asia.

Placenticeras ambai (Chiplonkar & Ghare, 1976) (p. 3, pl. 1 (figs 1, 4), text-

figs 2-3). Presumably Upper Albian to Cenomanian of India.

Placenticeras asiaticum (Iljin, 1975) (p. 156, pl. 31 (fig. 2a—b), pl. 33 (fig. 2)).

Cenomanian of Central Asia.

Placenticeras baghensis (Chiplonkar & Ghare, 1976) (p. 4, pl. 1 (figs 3-6)).

Presumably Upper Albian to Cenomanian of India.

Placenticeras beliakovae (Iljin, 1975) (p. 160, pl. 33 (fig. 1), pl. 34 (fig. 5)).

Cenomanian of Central Asia.

Placenticeras benningi Stephenson, 1956 (p. 247, pl. 44 (figs 21, 22), pl. 45

(figs 7-11)).

Placenticeras besairiei (Collignon, 1936) (Collignon 196556: 19, pl. 383

(fig. 1650)). Middle Turonian of Madagascar. (? = Placenticeras kaffrarium)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 385

Placenticeras beschtubensis (Ijin, 1975) (p. 163, pl. 29 (figs 1-2), pl. 39 (fig. 7)).

Turonian of Central Asia.

Placenticeras bidorsatum (Roemer, 1841) (p. 88, pl. 13 (fig. 8)). Lower Cam-

panian of Germany.

Placenticeras blanfordi (Chiplonkar & Ghare, 1977a) (p. 73, pl. 1 (fig. 3)).

Cenomanian—Turonian of India.

Placenticeras bobkovae Iljin, 1975 (p. 169, pl. 31 (fig. 1), pl. 35 (fig. 12)). San-

tonian of Central Asia.

Placenticeras canaliculatum Hyatt, 1903 (p. 243). Lower Campanian of Germany.

(Objective synonym of P. bidorsatum Roemer, 1841, fide Kennedy 1986).

Placenticeras carteri (Chiplonkar & Ghare, 1977a) (p. 70, pl. 2 (fig. 1), text-

fig. 1). Cenomanian—Turonian of India.

Placenticeras colquitti Wolleben, 1967 (p. 1164, pl. 150 (figs 1-4), text-fig. 8e).

Santonian of Texas.

Placenticeras crassatum Hyatt, 1903 (p. 24). Lower Senonian of Germany.

(= Placenticeras polyopsis (Dujardin))

Placenticeras cumminsi Cragin, 1893 (p. 237). Late Cenomanian, New Mexico

(Cobban 1983a), Texas.

Placenticeras dangerfieldi (Chiplonkar & Ghare, 1977a) (p. 69, pl. 1 (fig. 2)).

Cenomanian—Turonian of India.

Placenticeras fourtaui (Chiplonkar & Ghare 1977a) (p. 73, pl. 1 (fig. 1)). Ceno-

manian—Turonian of India.

Placenticeras fritschi de Grossouvre, 1894 (p. 124, pl. 5 (figs 1-2), text-fig. 52).

Coniacian of France, Czechoslovakia (Fritsch & Schloenbach 1872), and

India (Chiplonkar & Ghare 1977a).

Placenticeras gaurdakense (Luppov, 1963) (p. 144, pl. 1 (fig 1)). Middle Ceno-

manian of Central Asia.

Placenticeras gissarensis (Iljin, 1975) (p. 155, pl. 31 (fig. 1la—b), pl. 33 (fig. 1)).

Cenomanian of Central Asia.

Placenticeras grossouvrei Hyatt, 1903 (p. 237). Santonian of France. (= Placen-

ticeras polyopsis (Dujardin))

Placenticeras grossouvrei Semenov, 1899 (p. 97, pl. 2 (fig. 5)). Cenomanian of

Transcaspia. (non Placenticeras grossouvrei Hyatt, 1903)

Placenticeras guadalupae (Roemer, 1852) (p. 32, pl. 2 (fig. 1)). Campanian of

Germany (Riedel 1937), New Mexico, Texas, U.S. Western Interior (Ree-

side 1927b). (= Placenticeras syrtale (Morton, 1834))

Placenticeras helicus (Chiplonkar & Ghare, 1977a) (p. 74, pl. 2 (figs 3-4)).

Cenomanian—Turonian of India.

Placenticeras hispanicum (Mas & Wiedmann, 1980) (p. 267, figs 4-5). Middle

Cenomanian of Spain.

Placenticeras hyatti Diener, 1925 (p. 185). Santonian of France. (= Placenticeras

polyopsis (Dujardin, 1837))

Placenticeras iljini (Khakimov, 1976) (in Atabekian & Khakimov 1976: 93, pl. 10

(fig. 1)). Lower Campanian of Central Asia.

386 ANNALS OF THE SOUTH AFRICAN MUSEUM

Placenticeras incisum Hyatt, 1903 (p. 238). Santonian of France. (= Placenticeras

polyopsis (Dujardin))

Placenticeras intercalare (Meek & Hayden, 1860) (p. 117). Campanian of North

America and possibly Germany (Riedel 1937).

Placenticeras\ intermedium Johnson, 1903 (p. 206, pl. 8 (fig. 27)). Upper San-

tonian-lower Campanian of New Mexico. (= Placenticeras syrtale

(Morton))

Placenticeras kaffrarium Etheridge, 1904 (p. 89, pl. 3 (fig. 16)). Coniacian of

Zululand, offshore Alphard Group, Madagascar, Angola, ?South West

Africa—Namibia, and probably India.

Placenticeras keatingi (Chiplonkar & Ghare, 1977a) (p. 71, pl. 2 (fig. 2), text-

fig. 2). Cenomanian—Turonian of India.

Placenticeras kharesmense (Lahusen, 1884) (Archanguelski 1916: 40, pl. 6

(fig. 5), pl. 7 (fig. 1)). Turonian of Turkestan.

Placenticeras kolbajense (Sokolov, 1967) (p. 138, text-fig. p. 139). Upper Albian

of Central Asia.

Placenticeras kossmati (Chiplonkar & Ghare, 1979) (p. 131, pl. 1 (figs 1-3), pl. :

(fig. 1)). Upper Turonian—Coniacian of India.

Placenticeras kotzi (Iljin, 1975) (p. 165, pl. 30 (figs 1-2), pl. 39 (fig. 9)). Conia-

cian of Central Asia.

Placenticeras kutuzovae (Iljin, 1975) (p. 164, pl. 29 (fig. 3), pl. 39 (fig. 8)).

Turonian of Central Asia.

Placenticeras kysylchense (Ijin, 1958) (p. 728, figs 1a—b, 2d—e). Maastrichtian of

Uzbekistan.

Placenticeras kysylcumense Archanguelski, 1916 (p. 45, pl. 7 (figs 4-7), text-

fig. 16). Turonian of Turkestan.

Non Placenticeras liardense Whiteaves, 1889 (p. 150, pl. 20 (figs 1-2)). Upper

Cretaceous of Canada. (= Paragastoplites—fide Imlay 1961: 63)

Placenticeras luppovi Iljin, 1975 (p. 170, pl. 32 (fig. 1), pl. 35 (fig. 13)). San-

tonian of Central Asia.

Placenticeras maherndli Summesberger, 1979 (p. 155, pl. 14 (figs 58-61), pl. 15

(figs 62-66), text-figs 40-47). Upper Santonian of Austria.

Placenticeras meeki Boehm, 1898 (p. 200). Upper Senonian of North America

(Hyatt 1903; Reeside 1927a).

Placenticeras meeki var. tuberculata Hyatt, 1903 (p. 232, pl. 47 (fig. 5)). Upper

Senonian of Dakota.

Placenticeras memoriaschloenbachi Laube & Bruder, 1887 (p. 221, pl. 23

(fig. 1)). Cenomanian—Turonian of Bohemia and Saxony, France (Kennedy

et al. 1981).

Placenticeras memoriaeschloenbachi Laube & Bruder var. ambiloensis Collignon,

1965 (Collignon 1965a: 14, 16, pl. 381 (figs 1646), pl. 382 (1647-8)). Turo-

nian of Madagascar and India (Chiplonkar & Ghare 1977a). (= Placenticeras

kaffrarium Etheridge, 1904)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 387

Placenticeras merenskyi Haughton, 1930 (p. 363, pl. 11 (figs 1-3)). Cenomanian?

of South West Africa—Namibia.

Placenticeras milleri von Hauer, 1866 (p.5, pl. 2 (figs 1-2)). Coniacian—

_ Santonian of northern Alps and Gosau Beds, and tentatively India (Chip-

lonkar & Ghare 1977a).

Placenticeras mintoi Vredenburg, 1907 (p. 111, pls 14-15). Cenomanian of Bagh,

India (Chiplonkar & Ghare 1977a). (= ?Knemiceras)

Placenticeras murphyi (Collignon, 1965a) (p. 17, pl. 382 (fig. 1649)). Upper

Turonian of Madagascar. (= ?Placenticeras kaffrarium Etheridge)

Placenticeras newberryi Hyatt, 1903 (p. 203, pl. 31 (figs 3-5)). Upper Cretaceous,

New Mexico, Western Interior (Reeside 1927a). (= Placenticeras syrtale

(Morton, 1834))

Placenticeras orbignyanum (Geinitz, 1849) (p. 114, pl. 4 (fig. 1)). Coniacian of

N. Germany, Czechoslovakia and Madagascar. (= ?Placenticeras fritschi

de Grossouvre)

Placenticeras ornatus (Chiplonkar & Ghare, 1977a) (p. 72, pl. 1 (fig. 5)). Ceno-

manian—Turonian of India.

Placenticeras paraplanum Wiedmann, 1978 (p. 666, pl. 1 (figs 3-4), text-fig. 2a).

Santonian of the Gosau Beds, Austria, France (Amedro & Hancock 1985;

Kennedy 1987).

Placenticeras patagonicum Leanza, 1967 (p. 16, pl. 1 (figs 1-2), pl. 2 (fig. 1), pl. 3

(figs 1-3), pl. 4 (figs 2-3), pl. 9 (figs 1-5), pl. 10 (fig. 3)). Allegedly Lower

Campanian of Patagonia. (Turonian—pers. observation H.C.K.)

Non Placenticeras perezianum (Whiteaves, 1876) (p. 19, pl. 2 (fig. 1)). Lower

Cretaceous British Columbia, Canada. (= Cleoniceras (Grycia)—fide

McLearn 1972: 59)

Placenticeras pitniakense Wljin, 1975 (p. 167, pl. 32 (fig. 2), pl. 35 (fig. 11)).

Coniacian of Central Asia.

Placenticeras placenta (DeKay, 1828) (p. 278, pl. 5 (fig. 2)). Upper Cretaceous,

New Jersey, Alabama, U.S.A., Turkestan (Archanguelski 1916).

Placenticeras planum Hyatt, 1903 (p. 202, pl. 33 (figs 2—4)). Upper Cretaceous,

Texas, U.S.A. (Reeside 1927a), and tentatively India (Chiplonkar & Ghare

1977a). (= Placenticeras syrtale (Morton, 1834))

Placenticeras polyopsis (Dujardin, 1837) (p. 232, pl. 17 (fig. 12)). Santonian of

France, Austria, Germany, and possibly Alabama, U.S.A.

Placenticeras polyopsis amudariense (Iljin, 1975) (p. 171, pl. 32 (fig. 3), pl. 35

(fig. 14)). Santonian of Central Asia. (? = Placenticeras polyopsis s.s.)

Placenticeras proplanum (Iljin, 1975) (p. 166, pl. 37 (fig. 2), pl. 35 (fig. 10)).

Coniacian of Central Asia.

Placenticeras pseudocostatum Johnson, 1903 (p. 137, pl. 10 (fig. 29a). pl. 11

(fig. 29b—c)). Upper Santonian—Lower Campanian, New Mexico. (= Placen-

ticeras syrtale (Morton, 1834))

Placenticeras pseudorbignyanum Hyatt, 1903 (p. 242). Santonian of northern

Germany.

388 ANNALS OF THE SOUTH AFRICAN MUSEUM

Placenticeras pseudoplacenta Hyatt, 1903 (p. 216, pl. 43 (figs 3-11), pl. 44).

Upper Cretaceous of Utah, Minnesota, U.S.A. (Cobban 1983b—as Placen-

ticeras cumminsi Cragin, 1893), ?Palestine (Taubenhaus 1920), Oregon

(Reeside 1927a).

Placenticeras pseudoplacenta var. occidentalis Hyatt, 1903 (p. 217 (pars), pl. 45

(fig. 2 only)). Cenomanian of Texas, U.S.A. (= Placenticeras cumminsi

Cragin, 1893)

Placenticeras radiatum Riedel, 1937 (p. 219, pl. 16 (figs 1-2)). Santonian of

Germany. (= Placenticeras polyopsis (Dujardin, 1837))

Placenticeras rampuraensis (Chiplonkar & Ghare, 1977c) (p. 109, figs 1-2, 6A,

D). Turonian of India.

Placenticeras reineckei Haughton, 1925 (p. 271, pl. 13 (figs 4-5)). Upper Turo-

nian of Angola. (= ?Placenticeras kaffrarium Etheridge, 1904)

Placenticeras ribourianus (d’Orbigny, 1850) (p. 213). Santonian of France.

(= Placenticeras polyopsis (Dujardin, 1837))

Placenticeras rotundatum Johnson, 1903 (p. 135, pl. 9 (figs 28a—b)). Upper

Santonian—Campanian of New Mexico. (? = Placenticeras syrtale (Morton,

1834))

Placenticeras rooneyi Wolleben, 1967 (p. 1164, pl. 150 (figs 6-7), pl. 151

(figs 3-4) pl. 152 (figs 33-34), text-fig. 7d-g). Upper Santonian—Lower

Campanian of Texas and New Mexico, U.S.A.

Placenticeras saggitalis (Iljin, 1975) (p. 161, pl. 33 (fig. 2), pl. 34 (fig. 6)).

Cenomanian of Central Asia.

Placenticeras sancarlosense Hyatt, 1903 (p. 200, pl. 30 (figs 1-3), pl. 31

(figs 1-2)). Upper Santonian—Lower Campanian of Texas, and Western

Interior, U.S.A (Reeside 1927a). (= Placenticeras syrtale (Morton, 1834))

Placenticeras sancarlosense var. pseudosyrtale Hyatt, 1903 (p. 200, pl. 32, pl. 33

(fig. 1)). Upper Santonian—Lower Campanian of Texas, New Mexico, and

Western Interior, U.S.A (Reeside 1927a).

Placenticeras sanctacrucense Leanza, 1967 (p. 15, pl. 8 (fig. 2), pl. 4 (fig. 1)).

Allegedly Lower Campanian of Patagonia (Turonian—pers. observation

Be Ky

Placenticeras satriense (Collignon, 19656) (p. 40, pl. 431 (fig. 1782)). Upper

Coniacian of Madagascar. (= ?Placenticeras kaffrarium Etheridge)

Placenticeras schliiteri Hyatt, 1903 (p. 239). Santonian of northern Germany.

Placenticeras semiornatum (d’Orbigny, 1850) (p. 212). Upper Coniacian—

Santonian of France (Kennedy 1984).

Placenticeras simonyi Kossmat, 1907 (p. 49, pl. 4 (fig. 1)). Cenomanian of

Sokotra.

Placenticeras spathi (Chiplonkar & Ghare 1977c) (p. 110, fig. 7E-F). Turonian of

India.

Placenticeras spillmanni Hyatt, 1903 (p. 233, pl. 47 (figs 6-8)). Upper Cretaceous

of New Jersey and Mississippi, U.S.A.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 389

Placenticeras stantoni Hyatt, 1903 (p. 214). Upper Cretaceous of Utah and

Minnesota, U.S.A. (Cobban 1983b), and ?India. (= Placenticeras cumminsi

Cragin)

Placenticeras stantoni var. bolli Hyatt, 1903 (p. 214, pl. 40 (figs 3-7), pls 41-42,

43 (figs 1-2)). Upper Cretaceous of Texas, U.S.A., ?Upper Turonian of

Madagascar and ?India (Chiplonkar & Ghare 1977a). (= Placenticeras

cumminsi Cragin) (Chiplonkar & Ghare 1977a)

Placenticeras stantoni var. fortior (Collignon, 1965a) (p. 19, pl. 383 (fig. 1652)).

Turonian of Madagascar. (? = Placenticeras kaffrarium Etheridge)

Placenticeras stoliczkai (Chiplonkar & Ghare, 1979) (p. 131, pl. 2 (fig. 2), pl. 3

(figs 1-2)). Turonian—Coniacian of India.

Placenticeras subkaffrarium Spath, 1921 (p. 247, pl. 21 (fig. 2)). Coniacian of

Zululand and Madagascar. (= Placenticeras kaffrarium Etheridge)

Non Placenticeras subplanatum Taubenhaus, 1920 (p. 40, pl. 7 (fig. 2), pl. 8

(fig. 4)). Upper Cretaceous of Palestine. (Not a placenticeratid— Coilopo-

ceras ?)

Placenticeras syrtale (Morton, 1834) (p. 40, pl. 16 (fig. 4)). Upper Santonian

—Lower Campanian of U.S. Gulf Coast and Rocky Mountain areas (Reeside

19274).

Placenticeras syrtale adkinsi Wolleben, 1967 (p. 1164, pl. 151 (figs 8-9), pl. 152

(figs 1-2, 5-8), text-fig. 8g). Upper Santonian—Lower Campanian of Texas

and New Mexico.

Placenticeras syrtale costata Riedel, 1931 (p. 696, pl. 79 (fig. 2)). Santonian of

Germany. (= Placenticeras polyopsis (Dujardin))

Placenticeras syrtale var. halei Hyatt, 1903 (p. 206, pl. 27 (figs 16-17), pl. 28

(figs 3-6)). Upper Cretaceous of Alabama, U.S.A.

Placenticeras tamulicum (Blanford, 1862) (p. 118). Upper Turonian—Coniacian of

India and Madagascar.

Placenticeras telifer (Morton, 1834) (p. 38, pl. 2 (fig. 7)). Senonian of New Jersey,

U.S.A.

Placenticeras vancouverense (Meek, 1876) (p. 370, pl. 6 (fig. 1)). Upper Creta-

ceous of British Columbia, Canada and Sucia Island, U.S.A. (= ?Hoplito-

placenticeras)

Placenticeras viedmaense Leanza, 1967 (p. 12, pl. 4 (figs 1-3), pl. 8 (fig. 1), pl. 10

(fig. le)). Allegedly Lower Campanian of Patagonia (Turonian—pers. obs.

H.C.K.).

Placenticeras vredenburgi (Sarkar, 1966) (p. 144, pl. 11 (figs 1-2)). Turonian—

Coniacian of India (Chiplonkar & Ghare 1977a).

Placenticeras washbournei Leanza, 1967 (p. 17, pl. 5 (fig. 1), pl. 10 (fig. 4)).

Allegedly Lower Campanian, Patagonia (= Turonian—pers. obs. H.C.K.).

Placenticeras whitfieldi Hyatt, 1903 (p. 222, pl. 45 (figs 3-16), pls 46, 47

(figs 1-4)). Upper Cretaceous of Nebraska, South Dakota and Colorado,

U.S.A. (= Placenticeras meeki Béhm)

390 ANNALS OF THE SOUTH AFRICAN MUSEUM

Placenticeras ?yakounense (Whiteaves, 1900) (p. 280, pl. 36 (figs 1, la—b)).

Upper Cretaceous of Canada. (= ?Hoplitoplacenticeras)

Placenticeras n. sp. aff. kaffrarium Etheridge: Venzo, 1936 (p. 108 (50), pl. 11 (7)

(fig. 13)). Coniacian of Zululand and Madagascar (= Placenticeras kaffrar-

ium Etheridge)

Non Placenticeras zehariense (Collignon, 1966) (p. 33, pl. 18 (figs 5-9)). Turo-

nian of Morocco. (= ?Hoplitoides—fide Cobban & Hook 1980)

Genus and subgenus Hoplitoplacenticeras Paulcke, 1907

* Hoplitoplacenticeras antokazoense Collignon, 1970 (pp. 76, 80, pl. 638 (fig. 76),

pl. 639 (fig. 2354))—the !atter herein designated lectotype. Upper Cam-

panian of Madagascar.

Non Hoplitoplacenticeras awadi Hassan, 1971 (p. 71, pl. 2 (figs 8-9)). Lower

Maastrichtian of Egypt (= scaphitid).

*Hoplitoplacenticeras besairiei Collignon, 1970 (p.77, pl. 638 (figs

2349-50))—figure 2349 herein designated lectotype. Upper Campanian of

Madagascar. i

Hoplitoplacenticeras coesfeldiense var. schliiteri Michailov, 1951 (p. 82, pl. 15

(figs 60-61)). Upper Campanian of Don Basin, U.S.S.R., Westphalia,

F.G.R., and Wyoming, U.S.A. (Cobban 1963).

Hoplitoplacenticeras coesfeldiense (Schliiter, 1867) (p. 14, pl. 1 (figs 1, 4-5)).

Upper Campanian of Germany (Giers 1964), European Russia (Michailov

1951), Central Asia (Atabekian & Khakimov 1976), Caucasus, Crimea

(Naidin & Shimanskij 1959), and Ukraine (Naidin 1974), Sweden (Odum

1953), and France (Kennedy 1986).

Hoplitoplacenticeras costulosum (Schliter, 1867) (p. 17, pl. 2 (figs 2-4)). Upper

Campanian of Germany and, tentatively, Madagascar.

Hoplitoplacenticeras dolbergense (Schliter, 1876) (p. 159, pl. 44 (figs 1-4)).

Upper Campanian of Germany (Giers 1964; Schmid & Ernst 1975), tenta-

tively Madagascar (Collignon 1971), and France (Kennedy 1986).

Hoplitoplacenticeras fugen Matsumoto, 1984 (p. 25, pl. 8 (fig. 5)). Upper Cam-

panian of Hokkaido.

Hoplitoplacenticeras gosseleti (de Grossouvre, 1894) (p. 116, pl. 36 (fig. 1)).

Upper Cretaceous of France.

Hoplitoplacenticeras howarthi Collignon, 1970 (p. 80, pl. 639 (fig. 2351)). Upper

Campanian of Madagascar.

Non Hoplitoplacenticeras kambysis (Quaas, 1902) (p. 309, pl. 29 (figs 8-11)—as

?Scaphites kambysis). Lower Maastrichtian of Egypt. (It is a scaphitid.)

Hoplitoplacenticeras marroti (Coquand, 1859) (p. 995). (De Grossouvre 1894:

118, pl. 8 (fig. 3), pl. 9 (figs 2-3)). Upper Campanian of France?, Angola,

Germany (Howarth 1965: 391, pl. 12 (fig. 3), pl. 13 (fig. 3)) and Madagascar

(Collignon 1970: 80, pl. 639 (fig. 2353)).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 391

Hoplitoplacenticeras monju Matsumoto, 1982a (p. 249, figs 1-2). Upper Cam-

panian of Hokkaido. |

Hoplitoplacenticeras praematura (Imkeller, 1901) (p. 58, fig. 1). Upper Cam-

panian of Germany.

Hoplitoplacenticeras rarecostatum Khakimov, 1976 (in Atabekian & Khakimov

1976: 88, pl. 10 (fig. 5)). Upper Campanian of Central Asia.

Hoplitoplacenticeras rejaudryi (de Grossouvre, 1894) (p. 78, pl. 7 (figs 1-5),

pl. 14 (figs 6-75)). Upper Campanian of France.

Hoplitoplacenticeras trangahyense Collignon, 1970 (p. 76, pl. 638 (fig. 2347)).

Upper Campanian of Madagascar.

Hoplitoplacenticeras vancouverense (Meek, 1861) (Usher 1952: 93, pl. 25

(figs 1-2), pl. 31 (figs 21-22)). Upper Campanian of British Columbia.

Non Hoplitoplacenticeras yakounensis (Whiteaves, 1900) (p. 280, pl. 36 (fig. 1,

la—b)). Upper Campanian of Canada. (= Anahoplites—fide McLearn 1972:

57)

Subgenus Lemfoerdiceras Kennedy, 1986

Hoplitoplacenticeras (L.) lafresnayanum (d’Orbigny, 1842) (p. 326, pl. 97

(figs 3-5)). Upper Maastrichtian of France.

Hoplitoplacenticeras (L.) lemfoerdense (Schliter, 1872) (p. 160, pl. 19 (figs 1-2),

pl. 44 (figs 8-9)). Upper Campanian of Germany.

Genus Metaplacenticeras Spath, 1926

Metaplacenticeras? bowersi Anderson, 1958 (p. 255, pl. 70 (figs 3-4)). Upper

Campanian of California.

Metaplacenticeras californicum (Anderson, 1902) (p. 78, pl. 8 (figs 173-177)).

Upper Campanian of California (Anderson 1958: 254, pl. 36 (fig. 1)).

Metaplacenticeras pacificum (Smith, 1900) (p. 207, pls 26-28). Upper Campanian

of California (Anderson 1902: 79, pl. 8 (figs 162-164), pl. 9 (fig. 180); 1958:

254, pl. 37 (figs 1-4)).

Metaplacenticeras sanctaemonicae (Waring, 1917) (p. 70, pl. 9 (figs 20-21)).

Upper Campanian of California (Anderson 1958: 255).

Metaplacenticeras subtilstriatum (Jimbo, 1894) (p. 171, pl. 17 (fig. 1)). Upper

Campanian of Hokkaido.

ACKNOWLEDGEMENTS

Financial aid from the Foundation for Research Development, South Africa,

to Klinger, and the Trustees of the Sir Henry Strakosh Bequest, the Royal Society

and the Natural Environment Research Council to Kennedy, is gratefully

acknowledged.

We thank Sally Dove and Jacque Blaeske for technical assistance and Sandra

Saven for typing various drafts of the manuscript.

392 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

ADKINS, W. S. 1928. Handbook of Texas Cretaceous fossils. University of Texas Bulletin 2838:

1-385, pls 1-37.

AmeprRO, F. & Hancock, J. M. 1985. Les Ammonites de l’autoroute ‘l)Aquitane’, France

(Turonien et Santonien). Cretaceous Research 6 (1): 15-32.

ANDERSON, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proceedings of the California

Academy of Sciences (Geology) 2: 1-154, pls 2-5.

ANDERSON, F. M. 1958. Upper Cretaceous of the Pacific coast. Memoir of the Geological

Society of America 71: 1-378, pls 1-75.

ARCHANGUELSKY, A. D. 1916. Les mollusques du Crétacé supérieur du Turkestan. Mémoires

du comité Géologique (n.s.) 152: 1-57, 8 pls.

Arias, C. & WIEDMANN, J. 1977. Ammoniten und Alter der Utrillas-Schichten (Mittelkreide)

in der Ostlichen Provinz Albacete, SE Spanien. Neues Jahrbuch ftir Geologie und

Paldontologie Monatshefte 1977 (1): 1-14.

ARNOULD-SAGET, S. 1956. Contribution a l’étude des Engonoceratidae (les couches 4 Knemi-

ceras du sud Tunisien). Annales des mines et de la géologie Tunis 20: 1-47, pls 1-12.

ATABEKIAN, A. A. & KHAKIMOV, F. CH. 1976. (Campanian and Maastrichtian ammonites from

Central Asia.) Akademia Nauk Tadchikskoi SSR Institut geologii 1976: 1-146, pls 1-13. (In

Russian. )

Basse, E. 1931. Monographie paléontologique du crétacé de la Province de Maintirano

Madagascar. Gouvernement général de Madagascar et dépendances. Tananarive: Service

des Mines. :

Besairi£, H. 1930. Les rapports du Crétacé Malgache avec le Crétacé de |’Afrique australe.

Bulletin de la Societé Géologique de France (4) 30: 613-643, pls 64-67.

BLANForD, H. F. 1862. On the Cretaceous and other rocks of the South Coast and Trichinopoly

districts, Madras. Memoirs of the Geological Survey of India 4: 1-200.

BoeuM, J. 1898. Uber Ammonites pedernalis v. Buch. Zeitschrift der Deutschen Geologischen

Gesellschaft 1898: 183-201.

Bou te, M., Lemoine, P. & THEVENIN, A. 1906-7. Paléontologie de Madagascar. Annales de

Paléontologie 1 (1906): 173(1)-192(20), pls 14(1)-20(7); 2 (1907): 1(21)-56(76),

pls 1(8)-8(15).

CaLLomon, J. H. 1963. Sexual dimorphism in Jurassic ammonites. Transactions of the Leicester

literary and philosophical Society 57: 21-56.

CaLLomon, J. H. 1981. Dimorphism in ammonoids. Jn: House, M. R. & SENIor, J. R. eds. The

Ammonoidea. Systematics Association. Special Volume 18: 257-273.

Casey, R. 1960. Hengestites, a new genus of Gault ammonite. Palaeontology 2: 200-209, pl.

oo:

Casey, R. 1965. The Ammonoidea of the Lower Greensand. VI. Palaeontographical Society

(Monograph): 399-546, pls 67-90.

Casey, R. 1978. The Ammonoidea of the Lower Greensand. IX. Palaeontographical Society

(Monograph): 583-632, pls 98-100.

CHAMBERLAIN, J. A. & WESTERMANN, G. E. G. 1976. Hydrodynamic properties of cephalopod

shell ornament. Paleobiology 2: 316-331.

CHAVAN, A. 1947. La faune campanienne de Mont des Oliviers d’apres les matériaux

Vignal-Massée. Journal de Conchyliologie 87: 125-197, pls 2-4.

CHIPLONKAR, G. W. & GHARE, M. A. 1976. Palaeontology of the Bagh Beds. Part VII:

Ammonoidea. Bulletin of Earth Sciences 1976 (4, 5): 1-10, pls 1-2.

CHIPLONKAR, G. W. & GHARE, M. A. 1977a. Palaeontology of the Bagh Beds. Part X.

Scaphitidae. The Proceedings of the Indian Academy of Sciences 85B (2): 67—76, pls 1-2.

CHIPLONKAR, G. W. & GHARE, M. A. 1977b. Comments on Proplacenticeras stantoni (Hyatt)

and its variety bolli (Hyatt). Journal of the University of Poona, Science and Technology 50:

221-226, 1 pl.

CHIPLONKAR, G. W. & GHARE, M. A. 1977c. Ammonoidea from the Bagh Beds of Narmada

Valley. Indian Journal of Earth Sciences 4: 109-116, figs 1-7.

CHIPLONKAR, G. W. & GuHarRE, M. A. 1978. Taxonomic comments on Placenticeratidae with

diagnosis of a new genus Sancarlosia. Biovigyanam 4: 75-80.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 393

CHIPLONKAR, G. W. & GuHarE, M. A. 1979. Placenticeratid ammonoids from Trichinopoly

Group, South India with comments on biozonation. Geological Survey of India. Miscel-

laneous publications 45: 129-136, pls 1-3, 1 table.

CospBaN, W. A. 1963. Occurrence of the Late Cretaceous ammonite Hoplitoplacenticeras in

Wyoming. Professional Papers. United States Geological Survey 475—C: C60-—Cé2.

Coppan, W. A. 1969. The Late Cretaceous ammonites Scaphites leei Reeside and Scaphites

hippocrepis (DeKay) in the Western Interior of the United States. Professional Papers.

United States Geological Survey 619: 1-29, pls 1-5.

CosBaANn, W. A. 1983a. Mid-Cretaceous (Turonian) ammonite fauna from Fence Lake area of

west-central New Mexico. Memoirs. New Mexico Bureau of Mines and Mineral Resources

41: iv + 5-50, pls 1-14.

CopBan, W. A. 1983b. Molluscan fossil record from the northeastern part of the Upper

Cretaceous seaway, Western Interior. In: CoBBAN, W. A. & MEREWETHER, E. A. eds.

Stratigraphy and paleontology of Mid-Cretaceous rocks in Minnesota and contiguous areas.

Professional Papers. United States Geological Survey 1253: 1-25, pls 1-15.

CopBan, W. A. & Hook, S. C. 1980. The Upper Cretaceous (Turonian) ammonite family

Coilopoceratidae Hyatt in the Western Interior of the United States. Professional Papers.

United States Geological Survey 1192: iv + 1-28, pls 1-21.

CoLLIGNON, M. 1936. Fossiles du Turonien supérieur d’Antanitiloky. Jn: BeEsarri£é, H.

Recherches géologique a Madagascar, Prémiere Suite, La géologie du Nord-Ouest.

Mémoires de l’academie Malgache 21: 200-202, pl. 21.

CoLLIGNON, M. 1963. Atlas des fossiles caractéristiques de Madagascar (Ammonites). X

(Albien). Tananarive: Service Géologique. xv + 184 pp., pls 241-317.

CoLLIGNOoN, M. 1964. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XI.

(Cénomanien). Tananarive: Service Géologique. iv + 152 pp., pls 318-375.

CoLLIGNON, M. 1965a. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XII

(Turonien). Tananarive: Service Géologique. iv + 82 pp., pls 376-413.

CoLLIGNON, M. 1965b. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XIII

(Coniacien). Tananarive: Service Géologique. vii + 88 pp., pls 414-454.

CoLLIGNOoN, M. 1966. Les cephalopodes crétacés du Bassin cétier de Tarfaya. Notes et

Mémoires Service Géologique Maroc 175: 10-148, pls 1-35.

CoLLIGNON, M. 1969. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XV

(Campanien Inférieur). Tananarive: Service Géologique. xi + 216 pp., pls 514-606.

CoLLIGNON, M. 1970. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XVI

(Campanien moyen) & (Campanien supérieur). Tananarive: Service Géologique. iv +

82 pp., pls 607-639.

CoLLIGNoN, M. 1971. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XVII

(Maestrichtien). Tananarive: Service Géologique. iv + 44 pp., pls 640-658.

Cooper, M. R. 1978. Uppermost Cenomanian—basal Turonian ammonites from Salinas,

Angola. Annals of the South African Museum 75 (5): 51-152.

CoquanpD, H. 1859. Synopsis des animaux et des végétaux fossiles observés dans la formation

basal crétacée du Sud-Ouest de la France. Bulletin de la Societé géologique de France (2)

16: 945-1023.

CRAGIN, F. W. 1893. A contribution to the invertebrate paleontology of the Texas Cretaceous.

Texas Geological Survey, 4th Annual Report, part 2: 139-246, pls 24-46.

DeKay, J. E. 1828. Report on several fossil multilocular shells from the State of Delaware etc.

Annals of the Lyceum of Natural History of New York 2: 273-279, pl. 5 (figs 2-5 only).

Diener, C. 1925. Ammonoidea neocretacea. Fossilium Catalogus (1: Animalia) 29: 1-244.

DovuviLL£, H. 1890. Sur la classification des Cératites de la Craie. Bulletin de la Societé

géologique de France (3) 18: 273-292.

DusarDIN, F. 1837. Mémoire sur les couches du sol en Touraine et description des coquilles de

la craie et des Faluns. Mémoires de la Societe géologique France 2: 211-311, pls 15-20.

ETHERIDGE, R. 1904. Cretaceous fossils of Natal. 1. The Umkwelane Hill Deposit. Report of

the Geological Survey of Natal and Zululand 1: 71-93, pls 1-3.

FORSTER, R. 1975. Die geologische Entwicklung von Siid-Mozambique seit der Unterkreide und

die Ammoniten-Fauna von Unterkreide und Cenoman. Geologisches Jahrbuch (B) 12:

3-324, pls 1-17, 81 text-figs.

394 ANNALS OF THE SOUTH AFRICAN MUSEUM

FritscH, A & SCHLOENBACH, U. 1872. Cephalopoden der béhmischen Kreideformation. Prague:

A. Fritsch. 52 pp., 16 pls.

Geinitz, H. B. 1849-50. Das Quadersandsteingebirge oder Kreidegebirge in Deutschland. 293

pp., 12 pls. Freiberg: Craz & Gerlach.

Giers, R. 1964. Die Grossfauna der Mukronatenkreide (unteres Obercampan) im 6stlichen

Miinsterland. Fortschritte der Geologie Rheinland und Westfalen 7: 213-294, pls 1-8.

Giazunova, A. E. 1960. (About a new Albian genus of ammonites from Caspia.) Informatssb.

Palaeontolog. i stratigr. Vses i.i. Geol. in-t. 35: 92-94. (In Russian.)

GrossouvrE, A. DE 1894. Rechérches sur la craie supérieure, 2: paléontologie. Les ammonites

de la craie supérieure. Mémoires pour servir a l’explication de la carte géologique détaillée

de la France. 264 pp., 39 pls.

Hassan, M. Y. 1971. New molluscan fauna from the Maastrichtian of Kharga Oasis, South

Western Desert of Egypt. Proceedings of the Egyptian Academy of Sciences 23: 65-75, pls

1-2.

Hauer, F. von 1866. Uber die Cephalopoden der Gosauschichten. Beitrdge zur Paldontologie

Osterreichs 1: 7-14, pls 2-4.

HaucurTon, S. H. 1925. Notes on some Cretaceous fossils from Angola (Cephalopoda and

Echinoidea). Annals of the South African Museum 22 (1): 263-288, pls 12-15.

HauGuHtTon, S. H. 1926. Notes sur quelques Fossiles crétacés de Angola. Comunicagdes da

Comissao do Servico Geoldgico de Portugal 15: 5-32, pls 1-4.

HauGuton, S.H. 1930. On the occurrence of Upper Cretaceous marine fossils near Bogenfels.

S.W. Africa. Transactions of the Royal Society of South Africa 18: 361-365, pl. 11.

HowartTH, M. K. 1958. A monograph of the ammonites of the Liassic Family Amaltheidae in

Britain. Palaeontographical Society (Monograph): 1-53, 10 pls.

HowartTH, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bulletin of the

British Museum of Natural History (Geology) 10: 337-412, pls 1-13.

HowartH, M. K. 1985. Cenomanian and Turonian ammonites from the Novo Redondo area,

Angola. Bulletin of the British Museum (Natural History) (Geology) 39: 73-105.

Hyatt, A. 1900. Cephalopoda. Jn: ZitreL, K. A. von. ed. Textbook of Palaeontology:

p. 502-604. Translated by: Eastman, C. R. [1896-1900]. London and New York:

Macmillan.

Hyatr, A. 1903. Pseudoceratites of the Cretaceous. Monograph. United States Geological

Survey 44: 1-351, 47 pls.

Inmn, V. D. 1958. (New genus of ammonites from south western Uzbekistan.) Dokladi

Akademii Nauk SSSR 121: 727-729, 1 fig. (In Russian.)

Inn, V. D. 1959. (Stratigraphy of the Upper Cretaceous deposits of West Uzbekistan and

adjacent regions of Turkmenia.) Doklady Trud¥ VNIGNI 1959: 181-222, pls 1-8. (In

Russian.)

Inn, V. D. 1975. (Ammonites of the family Placenticeratidae Hyatt from the Upper

Cretaceous sediments of Central Asia.) Trudy¥ vsesoyuznyi Nauchno—Issledovatel’ skii

Geologorazvedochny Neftyanoi Institut (VNIGNI) 171: 154-174, pls 26-34. (In Russian. )

IMLtay, R. W. 1961. Characteristic Lower Cretaceous megafossils from northern Alaska.

Professional Papers. United States Geological Survey 335: 1-74, pls 1-20.

IMKELLER, H. 1901. Die Kreidebildungen und ihre Fauna am Stallauer Eck und Enzenauer

Kopf bei Télz. Palaeontographica 48: 1-64, pls 1-3.

Jimpo, K. 1894. Beitrage zur Kenntniss der Fauna der Kreide-formation von Hokkaido.

Geologisch Palaeontologische Abhandlungen (Neue Folge) 2: 149-194, pls 17-25.

JOHNSON, D. W. 1903. The geology of the Cerillos Hills, New Mexico. Contributions of the

Department of Geology Columbia University 10: 1-221, pls A—U, 1-14.

KENNEDY, W. J. 1977. Ammonite evolution. Jn: HaLLaM, A. ed. Patterns of Evolution:

251-304. Amsterdam: Elsevier.

KENNEDY, W. J. 1984. Systematic palaeontology and stratigraphic distribution of the ammonite

faunas of the French Coniacian. Special Papers in Palaeontology 31: 1-160, pls 1-33,

42 figs.

KENNEDY, W. J. 1986. Campanian and Maastrichtian ammonites from northern Aquitaine,

France. Special Papers in Palaeontology 36: 1-145, pls 1-23.

KENNEDY, W. J. 1987. Ammonites from the type Santonian and adjacent parts of northern

Aquitaine, western France. Palaeontology 30: 765-782, pls 80-82.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 395

KENNEDY, W. J. & CoBBAN, W. A. 1976. Aspects of ammonite biology, biogeography, and

biostratigraphy. Special Papers in Palaeontology 17: 1-94, pls 1-11, 24 text-figs.

KENNEDY, W. J. & KLINGER, H.C. 1973. IN: KENNEDY, W. J., KAUFFMAN, E. G. & KLINGER,

H. C. Upper Cretaceous invertebrate faunas from Durban, South Africa. Transactions of

the Geological Society of South Africa 76 (2): 95-111, pls 1-6.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bulletin of the British Museum (Natural History)

(Geology) 25: 263-315, 1 pl., 12 text-figs.

KENNEDY, W. J., JUIGNET, P. & Hancock, J. M. 1981. Upper Cenomanian ammonites from

Anjou and the Vendée, Western France. Palaeontology 24: 25-84, pls 3-17.

KENNEDY, W. J. & Wricut, C. W. 1983. Ammonites polyopsis Dujardin, 1837, and the

Cretaceous ammonite family Placenticeratidae Hyatt, 1900. Palaeontology 26: 855-873,

pls 85-87.

KENNEDY, W. J. & WricuT, C. W. 1985. Evolutionary patterns in Late Cretaceous ammonites.

Special Papers in Palaeontology 33: 131-143.

KLINGER, H. C. 1977. Cretaceous deposits near Bogenfels, South West Africa. Annals of the

South African Museum 73 (3): 81-92.

Kuincer, H. C. & Kennepy, W. J. 1980a. Cretaceous faunas from Zululand and Natal, South

Africa. The ammonite subfamily Texanitinae Collignon, 1948. Annals of the South African

Museum 80: 1-357.

KLINGER, H. C. & KENNEDY, W. J. 19806. The Umzamba Formation at its type section,

Umzamba Estuary (Pondoland, Transkei), the ammonite content and palaeogeographical

distribution. Annals of the South African Museum 81 (6): 207-222.

KLINGER, H. C. & KENNEDY, W. J. 1980c. In: KLINGER, H. C., KAUFFMAN, E. G. & KENNEDY,

W. J. Upper Cretaceous ammonites and inoceramids from the offshore Alphard Group of

South Africa. Annals of the South African Museum 82 (7): 293-320, 10 figs.

KuinGer, H. C. & KENNEDY, W. J. 1984. Cretaceous faunas from Zululand and Natal, South

Africa. The ammonite subfamily Peroniceratinae Hyatt, 1900. Annals of the South African

Museum 92 (3): 113-294.

KossmaT, F. 1907. Geologie der Inseln Sok6tra, Sémha und Abd el Kiri. Denkschriften der

kaiserlichen Akademie der Wissenschaften, mathematisch-naturwissenschafliche Klasse 71

(1): 1-62, pls 1-5.

KossmaT, F. 1895-1898. Untersuchungen tber die siidindische Kreideformation. Beitrdge zur

Palaeontologie Oesterreich-Ungarns und des Orients 9 (1895): 97-203 (1-107), pls 15-25

(1-11); 11 (1897): 1-46 (108-153), pls 1-8 (12-19); 11 (1898): 89-152 (154-217), pls 14-19

(20-25).

KULLMANN, J. & WIEDMANN, J. 1970. Significance of sutures in phylogeny of Ammonoidea.

Paleontological contributions of the University of Kansas 47: 1-32.

LAHUSEN, J. 1884. (Description of two ammonite species from the Amudara Valley.) In:

Romanowsky, G. (Material on the Geology of Turkestan): 133-135. St Petersburg:

Academic Press. (In Russian.)

LausE, G. C. & BRupER, G. 1887. Ammoniten der bohmischen Kreide. Palaeontographica 33:

217-239, pls 23-29.

LEANZA, A. F. 1967. Description de la fauna de Placenticeras del Cretacico superior de

Patagonia austral con consideraciones acerca de sun posicion estratigrafica. Boletin de la

Academia Nacional de Ciencias, Cordoba 46: 7-28, pls 1-10.

Lewy, Z. 1981. A Late Albian Hypengonoceras (Ammonoidea) from the ‘Bentonite-Bed’ at

Makhtesh Ramon, Southern Israel. Israel Journal of Earth-Sciences 30: 35-38, 4 figs.

Luprov, N. P. 1963. (New Cenomanian and Lower Turonian ammonites of the genus

Placenticeras from Central Asia.) Trudy VSEGEI (n.s.) 109: 135-160. (In Russian.)

Maumoup, I. G. E. 1955. Etudes paléontologiques sur la faune crétacique du Massif du

Moghara. Publications de l’institut du desert d’ Egypte 8: 1-103, pls 1-19.

Makowsk!, H. 1962. Problem of sexual dimorphism in ammonites. Palaeontologia Polonica

12: 1-92, pls 1-20.

MARCINOWSKI, R. 1980. Cenomanian ammonites from German Democratic Republic, Poland

and the Soviet Union. Acta geologica polonica 30: 215-325, pls 1-20.

396 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mas, J. R. & WIEDMANN, J. 1980. Ammoniten und Alter der Mittelkreide-Transgression in der

westlichen Provinz Valencia, Spanien. Neues Jahrbuch fiir Geologie und Paldontologie,

Abhandlungen 159: 256-272.

Matsumoto, T. 1942. Fundamentals in the Cretaceous stratigraphy of Japan. 1. Memoirs of the

Faculty of Science, Kyushu Imperial University (D) Geology 1: 129-280, pls 5-20.

Matsumoto, T. 1953. The ontogeny of Metaplacenticeras subtilstriatum (Jimbo). Japanese

Journal of Geology and Geography 23: 139-150, pl. 13.

Matsumoto, T. 1982a. Note on Hoplitoplacenticeras from Hokkaido. Proceedings of the

Japanese Academy 58B: 249-252, figs 1-2.

Matsumoto, T. 19826. A stratigraphical restudy at the type locality of Metaplacenticeras

subtilstriatum (Jimbo) (Cretaceous Ammonite). Proceedings of the Japanese Academy

58B: 253-255.

Matsumoto, T. 1984. Some ammonites from the Campanian (Upper Cretaceous) of northern

Hokkaido. Palaeontological Society of Japan Special papers 27: 1-93, pls 1-31.

Maxis, C. 1943. Ammoniti maestrichtiane della Tripolitania. Societe geologica italiana

Bolletino 61: 469-487, pl. 8.

McLacuian, I. R. & McMILLAN, I. K. 1979. Microfaunal biostratigraphy, chronostratigraphy

and history of Mesozoic and Cenozoic deposits on the coastal margin of South Africa.

Geokongress 77: Special publication. Geological Society of South Africa 6: 161-181.

McLearn, F. H. 1972. Ammonoids of the Lower Cretaceous Sandstone Member of the Haida

Formation, Skidegate Inlet, Queen Charlotte Islands, western British Columbia. Bulletin.

Geological Survey of Canada 188: 1-78, 45 pls.

MEEK, F. B. 1861. Descriptions of new Cretaceous fossils collected by the North-Western

Boundary Commission, on Vancouver and Sucia islands. Proceedings of the Academy of

Natural Sciences of Philadelphia 1861 (10): 314-318.

MEEK, F. B. 1870. A preliminary list of fossils collected by Dr. Hayden in Colorado, New

Mexico and California, with brief descriptions of a few of the new species. Proceedings of

the American Philosophical Society 11 (1869-1870): 425-431.

MEEK, F. B. 1876. A report on the invertebrate Cretaceous and Tertiary fossils of the upper

Missouri country. /n: HAyDEN, F. V. Report of the United States Geological Survey of the

Territories 9: lxiv, 1-629, 45 pls.

MEEK, F. B. & HAYDEN, F. V. 1860. Descriptions of new Lower Silurian (Primordial), Jurassic,

Cretaceous and Tertiary fossils, collected in Nebraska Territory with some remarks on the

rocks from which they were obtained. Proceedings of the Philadelphia Academy of Sciences

13: 415-447.

MicnalLov, N. P. 1951. (Upper Cretaceous ammonites from southern European Russia and

their significance for stratigraphy.) Akademia Nauk U.S.S.R. Trudy Instituta geologicheski

Nauk 129: 1-145, pls 1-19. (In Russian.)

MicuaiLova, I. A. 1974. (The relationship between early Cretaceous and late Cretaceous

Hoplitaceae.) Review. Bulgarian Geological Society 35: 117-132. (In Russian.)

MicuaiLova, I. A. 1978. (Characteristics of the morphogenesis of the family Placenticeratidae

Hyatt.) Doklady Akademii Nauk SSSR 242: 207-210. (In Russian.)

Mirzoev, G. G. 1967. (On the interrelationship of the families Hoplitidae and Placenti-

ceratidae.) Paleontologicheskii Zhurnal (for 1967): 56-70, pls 3—4. (In Russian.)

Morton, S. G. 1834. Synopsis of the organic remains of the Cretaceous groups of the United

States. Illustrated by nineteen plates to which is added an appendix containing a tabular

view of the Tertiary fossils discovered in America. Philadelphia: Key & Biddle. 88 pp.,

18 pls.

Narpin, D. P. 1974. Ammonoidea. Jn: KrimuHo ts, G. JA. ed. (Atlas of Upper Cretaceous faunas

of Donbassa.) NEDRA: 158-195, pls 53-70. (In Russian.)

Narpin, D. P. & SHIMANSKU, V. N. 1959. Golovonogie mollyuski. Jn: Moskvin, M. M. ed.

Atlas verk-ehnemelovij fauny severnogo Kavkaza i Kryma: 166-220, pls 1-23. Moskva:

VNIGAZ.

Nowak, J. 1909. (On some cephalopods and the character of the fauna of the Carpathian

Campanian.) Kosmos 34: 765-787. (In Polish with German summary.)

@pum, H. 1953. De geologiska resultaten fran Borrningorna vid Hollviken. V. The macro-

fossils of the Upper Cretaceous. Sveriges geologiska undersokning (C), Arsbok 46: 3-37,

pls 1-4.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 397

Orsicny, A. D’. 1840-2. Paléontologie francaise: Terrains Crétacés. I. Céphalopodes. Paris:

Victor Masson.

Orpicny, A. D’. 1850. Prodréme de Paléontologie stratigraphique universelle des animaux

mollusques et rayonnés 2: 1-428. Paris: Masson.

PauLcxe, W. W. 1907. Die Cephalopoden der oberen Kreide Siidpatagoniens. Berichte der

Naturforschenden Gesellschaft zu Freiburg i.B. 15: 167-248, pls 10-19.

PERVINQUIERE, L. 1907. Etudes de paléontologie tunisienne. I, Céphalopodes des terrains

secondaires. Carte Géologique Tunisie: 1-438, pls 1-27.

Quaas, A. 1902. Beitrag zur Kenntnis der Fauna der obersten Kreidebildungen in der

Libyschen Wiiste (overwegischichten und Blattertone). Palaeontographica 30: 153-336,

pls 20-33.

REESIDE, J. B. 1926. A comparison of the genera Metaplacenticeras Spath and Placenticeras

Meek. Professional Papers. United States Geological Survey 147—A: 1-2, pls 1-2.

REESIDE, J. B. 1927a. Cephalopods from the lower part of the Cody Shale of Oregon Basin,

Wyoming. Professional Papers. United States Geological Survey 150—A: 1-10, pls 1-8.

REESIDE, J. B. 1927b. The cephalopods of the Eagle Sandstone and related formations in the

Western Interior of the United States. Professional Papers. United States Geological

Survey 151: 1-40, pls 1-45.

RENz, O. 1968. Uber die Untergattungen Venezoliceras Spath und Laraiceras n. subgen. der

Gattung Oxytropidoceras Stieler (Ammonoidea) aus den venezolanischen Anden. Eclogae

geologicae Helvetiae 61: 615-655, pls 1-13.

RENz, O. 1970. Uber die Gattung Parengonoceras Spath, Knemiceras Boehm und Neophlyc-

ticeras Spath (Ammonoidea) aus den Anden Venezuelas. Eclogae geologicae Helvetiae 63:

1021-1057, pls.

RreDeEL, I. 1931. Zur Stratigraphie und Faziesbildung im Oberemscher am Siidrand des Beckens

von Minster. Jahrbuch der preussischen geologischen Landesanstalt. Bergakademie 51:

605-713, pls 72-79.

RIEDEL, I. 1937. Die Salzbergmergel und ihre aquivalente in Westfalen. Jahrbuch der

preussischen geologischen Landesanstalt. Bergakademie 58: 207-229, pl. 16.

RoeMER, F. A. 1840-41. Die Versteinerungen des norddeutschen Kreidegebirges. 145 pp., 16 pls.

Hahn’schen Hofbuchandlung (p. 1-48, pls 1-7 (1840); p. 49-145, pls 8-16 (1841)).

Roemer, F. A. 1852. Die Kreidebildungen von Texas und ihre organischen Einschlisse. Bonn.

RomMAN, F. 1938. Les ammonites jurassiques et crétacées. 554 pp., 53 pls, 496 figs. Paris:

Masson.

RUZHENTSEV, V. E. 1949. (Fundamental types of evolutionary changes in the suture lines of

Upper Palaeozoic ammonites.) Trudy Palaeontologischskii Institut 20: 183-198. (In

Russian. )

RUZHENTSEV, V. E. 1957. (Phylogenetic system of Palaeozoic ammonites.) Byulleten’ Mos-

kovskogo Obschestva Isytatele. Prirody. Otdel Geologicheski 32: 49-64. (In Russian.)

SALFELD, H. 1924. Die Bedeutung der Konservativstamme fiir die Stammesentwicklung der

Ammonoideen. Leipzig: Salfeld. 16 pp., 16 pls.

SARKAR, S. S. 1966. On Placenticeras mintoi Vredenburg, 1908 from the Bagh Beds (Creta-

ceous) of India. Paldontologische Zeitschrift 40: 144-146.

SCHINDEWOLF, O. H. 1967. Studien zur Stammesgeschichte der Ammoniten. VI. Akademie

der Wissenschaften und der Literatur. Abhandlungen der Mathematisch-

naturwissenschaftlichen Klasse 8: 722 (643)—808 (130).

SCHLUTER, C. 1867. Beitrag zur Kenntniss der jitingsten Ammoneen Norddeutschlands. Bonn:

A. Henry. 36 pp., 6 pls.

SCHLUTER, C. 1871-1876. Cephalopoden der oberen deutschen Kreide. Palaeontographica 21

(1871): 1-24, pls 1-8; 21 (1872): 25-120, pls 9-35; 24 (1876): 1-144 (121-264) + x,

pls 36-55.

ScHMID, F. & Ernst, G. 1975. Ammoniten aus dem Campan der Lehrter Westmulde und ihre

stratigraphische Bedeutung. 1. Teil: Scaphites, Bostrychoceras und Hoplitoplacenticeras.

Berichte der Naturhistorischen Gesellschaft 119: 315-359, pls 1-4.

ScHOLZ, G. 1979. Die Ammoniten des Vracon (Oberalb, dispar-Zone) des Bakony-Gebirges

(Westungarn) und eine Revision der wichtigsten Vracon-arten der West-Mediterranen

Faunenprovinz. Palaeontographica A 165: 1-136, pls 1-30.

398 ANNALS OF THE SOUTH AFRICAN MUSEUM

SEMENOV, W. P. 1899. (The fauna of the Cretaceous Deposits of Mangyschlak and some other

localities in the Transcaspian Province.) Travaux de la Societe Imperial de St. Petersbourg

28 (Section Géologie et Minéralogie): 1-178, pls 1-5. (In Russian.)

SEYED-EMAMI, K., Forster, R. & MojTeHEDIA, A. 1984. Ammoniten aus dem mittleren

Cenoman von Nordost-Iran (Koppeh-Dagh). Neues Jahrbuch fiir Geologie und Paldonto-

logie, Monatshefte 1984 (3): 159-172, 6 figs.

SmitH, J. P. 1900. The development and phylogeny of Placenticeras. Proceedings of the

California Academy of Sciences (3. Geology) 1: 181-232, pls 25-28.

SoxoLov, M. I. 1961. (Karamaites—a new ammonite genus from the Vraconnian deposits of

eastern Mangyschlak.) Byulleten’ Moskovskogo Obschchestva Ispytatelei prirody. Otdel

Geologischeski 36: 152. (In Russian.)

SoxoLov, M. I. 1967. (Karamaiceras gen. nov. from the Vraconnian deposits of Eastern

Mangyshlak.) Paleontologischeskii Zhurnal (for 1967): 138-139. (In Russian.)

SpaTH, L. F. 1921. On Cretaceous Cephalopoda from Zululand. Annals of the South African

Museum 12 (7): 217-321, pls 19-26.

SpaTH, L. F. 1922. On Cretaceous Ammonoidea from Angola, collected by Professor J. W.

Gregory, D.Sc., F.R.S. Transactions of the Royal Society of Edinburgh 53: 91-160, 4 pls.

SpaTtH, L. F. 1924. On a new ammonite (Engonoceras iris, sp. n.) from the Gault of

Folkestone. Annals and Magazine of Natural History (9) 14: 504-508.

SpaTH, L. F. 1926. On new ammonites from the English Chalk. Geological Magazine 63:

77-83.

SpATH, L. F. 1930. The Lower Cretaceous ammonoidea with notes on Albian Cephalopoda

from Hazara. The fossil fauna of the Samana Range and some neighbouring areas. V.

Palaeontologia Indica (n.s.) 15: 50-66, pls 7-9.

SpATH, L. F. 1931. A monograph of the Ammonoidea of the Gault. VIII. Palaeontographical

Society (Monograph): 313-378, pls 31-36.

STEPHENSON, L. W. 1956. Fossils from the Eutaw Formation, Chattahoochee River region,

Alabama-—Georgia. Professional Papers. United States Geological Survey 274J: 227-250,

pls 38-45.

STOLICZKA, F. 1863-66. The fossil cephalopoda of southern India. Ammonitidae with revision

of the Nautilidae etc. Memoirs of the Geological Survey of India (1), Palaeontologica indica

3 (1863) (10): 414-56, pls 26-31; (1864) (2-5): 57-106, pls 32-54; (1865) (66-9): 107-154,

pls 55-80; (1866) (10-13): 155-216, pls 81-94.

SUMMESBERGER, H. 1979. Eine obersantone Ammonitenfauna aus dem Becken von Gosau

(Oberosterreich). Annalen des Naturhistorischen Museum, Wien 82: 109-176, pls 1-15.

TAUBENHAUS, H. 1920. Die Ammoneen der Kreideformation Palastinas und Syriens. Zeitschrift

des deutsch-Paldstina Vereins 63: 1-58, pls 1-10.

UsHer, J. L. 1952. Ammonite faunas of the Upper Cretaceous rocks of Vancouver Island,

British Columbia. Bulletin. Geological Survey of Canada 21: i-v, 1-182, pls 1-30.

VENZO, S. 1936. Cefalopodi del Cretaceo medio-superiore dello Zululand. Palaeontographia

italica 36: 1-33, pls 5(1)—-12(8).

VREDENBURG, E. W. 1907. The ammonites of the Bagh Beds. Records of the Geological Survey

of India 36: 109-125, pls 14-17.

Warp, P. D. 1978. Revision to the stratigraphy and biochronology of the Upper Cretaceous

Nanaimo Group, British Columbia and Washington State. Canadian Journal of Earth

Sciences 15: 405-423.

Warinc, C. A. 1917. Stratigraphic and faunal relations of the Martinez to the Chico and Tejon

of Southern California. Proceedings of the California Academy of Sciences (4) 7: 41-124,

pls 7-116.

WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentralblatt fiir Mineralogie,

Geologie und Paldontologie (B) (for 1916) 8: 185-195.

WHITEAVES, J. F. 1876. On some invertebrates from the coal-bearing rocks of the Queen

Charlotte Islands. Jn: WuiTEAvES, J. F. Geological Survey of Canada. Mesozoic Fossils 1:

1-92, pls 1-10.

WuitTEAVES, J. F. 1889. On some invertebrates from the coal-bearing rocks of the Queen

Charlotte Islands. Geological Survey of Canada. Mesozoic Fossils 1: 1-92, pls 1-100.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 399

WuitEAvES, J. F. 1900. On some additional or imperfectly understood fossils from the

Cretaceous rocks of the Queen Charlotte Islands, with a revised list of the species from

these rocks. Geological Survey of Canada. Mesozoic Fossils 1 (4): 263-307, pls 33-39.

WIEDMANN, J. 1962. Die systematische Stellung von Hypophylloceras Salfeld. Neues Jahrbuch

. fiir Geologie und Paldontologie, Abhandlungen 115: 243-262.

WIEDMANN, J. 1970. Probleme der Lobenterminologie. Eclogae Geologicae Helvetiae 63:

909-922.

WIEDMANN, J. 1978. Eine palaogeographisch interessante Ammonitenfaunula aus der alpinen

Gosau. Eclogae Geologicae Helvetiae 71: 663-675, pls 1-2.

WIEDMANN, J. & KULLMANN, J. 1981. Ammonoid sutures in ontogeny and phylogeny. In:

House, M. R. & SENior, J. R. eds. The Ammonoidea. Systematics Association. Special

Volume 18: 215-255.

WOLLEBEN, J. A. 1967. Senonian (Cretaceous) Mollusca from Trans-Pecos Texas and north-

eastern Chihuahua, Mexico. Journal of Paleontology 41: 1150-1165, pls 147-152.

WriGcuT, C. W. 1957. Cephalopoda, Ammonoidea. Jn: Moore, R. C. ed. Treatise on

invertebrate paleontology. Part L, Mollusca 4. xxii + 490 pp. New York and Lawrence:

Geological Society of America and University of Kansas Press.

YounG, K. 1963. Upper Cretaceous ammonites from the Gulf Coast of the United States.

Publications. University of Texas 6304: 1-373, pls 1-82, text-figs 7-34.

APPENDIX

ORIGINAL DIAGNOSES OF GENERA HERE PLACED IN THE

SYNONYMY OF PLACENTICERAS

Karamaiceras Sokolov, 1967: 138

Name of the genus from the mouth of the Karamai to the east of Mangyshlak.

Type species. K. kolbajense sp. nov., Upper Albian, zone of Stoliczkaia

dispar (Vraconnian); East Mangyshlak.

Diagnosis

Shell involute, discoidal, formed by high, strongly convoluted whorls. Whorl

section in the species is a high, narrow triangle with truncated apex and weakly

convex flanks. Umbilicus is rather narrow, deep, funnel-like with steep walls and

a sharp edge. Ornament is represented by compressed, weakly s-shaped, curved

ribs, umbilical and marginal tubercles. Suture line consists of shallow bulb-shaped

lobes and low, wide, rounded saddles. Ventral lobe is short, the umbilical (one)

is sharply asymmetric. The base of the latter has a kind of wide sinus with two

relatively short branches; the inner one of which is longer than the outer. The

second lobe which divides the first lateral saddle attains the length of the outer

branch of the umbilical lobe.

Species composition. Type species.

Comparison

The new genus is distinguished from the closest genus Placenticeras by the

development of the suture line.

(Free translation from original Russian. )

400 ANNALS OF THE SOUTH AFRICAN MUSEUM

Turkmenites Ijin, 1975: 154-155

Type species. Placenticeras gaurdakensis Luppov, 1963, from the Lower

Cenomanian of Central Asia (Luppov 1963: 144, pl. 1 (figs 1, 2)).

Diagnosis

Shells are compressed with narrow, funnel-shaped umbilicus, smooth venter

and slightly curved flanks. The umbilical tubercles are mammiform, pointed on

the shell and blunt on internal casts. The external tubercles are elongated

longitudinally.

The ribs are always sickle-shaped, the most prominent are on the outer part

of the flanks. The surface of the shell is covered with fine sickle-shaped striae. The

lobes are strongly separated by means of asymmetrical bases of the first three and

by a wide, denticulated base at the posterior. The first lobe is always slanted

towards the second. The fourth lobe is very small, smaller by far than the fifth

lobe. The transformation of the first umbilical lobe comes about by means of

formation at its base, and later on the inner sides of the lobes and incisions, along

with their subsequent complete separation and transformation into independent

lobes. On the first whorl, in the base of the first umbilical lobe, two notches arise

of which the outer one lags somewhat in development. On the second whorl a

notch arises on the inner flank of the lobe. At this particular stage, a notch arises

in the outer parts of the saddle V/U close to its top, from which a secondary lobe

develops.

Comparison

Of the genus Semenovites Glazunova (Glazunova 1960) from the sediments

of the Upper Albian, characterized by a well-developed first lobe and the

complete separation of the third lobe.

(Free translation from original Russian.)

Kopetdagites Ijin, 1975: 157

Type species. Kopetdagites kopetdagensis gen. et sp. n. from the sediments of

the Upper Cenomanian of Turkmenia.

Diagnosis

Shells high, from compressed to inflated whorl section. Venter is smooth. On

early whorls venter is smooth or crenulated, on middle and adult by means of two

rows of highly crenulated tubercles, oblique in the side of the flanks and

projecting over the ventral sides. The umbilicus is narrow and funnel-shaped. The

umbilical tubercles are always high, mammiform. The flanks are covered with

sickle-shaped ribs, bifurcating on to the outer half of the flank. On the point of

division of the ribs, tubercle-like swellings may arise.

The suture line at a diameter of 40 mm and upwards on the flanks is relatively

deep with long side branches and has ventral lobes and up to 9 lobes alternating

with the saddles. The first lobe is short, slanted towards the second lobe. The

CRETACEOUS FAUNAS FROM SOUTH AFRICA 401

fourth lobe is small and weakly developed. The fifth lobe is significantly larger

than the fourth and somewhat larger than the sixth. The first saddle is always

lower than the second. On the inner side are up to six lobes, of which the dorsal

is the most prominent. The transformation of the first umbilical lobe takes place

by formation, at the beginning, on its exterior and subsequently on the inner side

of the notches (branching or dendritic) with their complete separation and

transformation into independent lobes later on. On the second whorl, a notch

develops in the outer segment of saddle V/U out of which a further secondary lobe

develops.

Comparison

Of Turkmenites gen. n. characterized by the presence of bifurcating ribs.

(Free translation from original Russian.)

Mediasiceras Ijin, 1975: 159-160

Type species. Mediasiceras beliakovae Iljin gen. et sp. n. from the sediments

of the Lower Cenomanian of the south-western spurs of the Ghissarski Range.

Diagnosis

Shells discoidal with high arrow-shaped whorl section, narrow, funnel-shaped

umbilicus and tapering to the centre, smooth, narrow, concave on the shells, and

ventral side, along the middle of which a low ridge is discernible. The flanks are

slightly convex near the middle. Ornament consists of sharp, weak, short-curved

ribs on the outer part of the flanks; parts of fine, sickle-shaped striae just begin to

appear from the suture and the small tubercles along the edge.

The external suture consists of short, with strongly separated, lateral

extensions, ventral lobes, and 8-9 strongly separated lobes, alternating with bifid

blunt saddles. The dorsal lobes are peak-shaped.

The distinctly expressed slant of the first lobe (from the ventral lobe) toward

the second and the third lobe is a characteristic feature of the suture line of the

genus.

Comparison

Of the genus Proplacenticeras Spath (1926) characterized by the structure of

the suture line, having a distinctly expressed slant of the first lobe towards the

second lobe, and an asymmetrical branching base of the second and third lobes.

Of the genus Anaplacenticeras Ijin (1959) characterized by the presence of a ridge

on the ventral side, the presence of umbilical tubercles, a funnel-shaped form of

the umbilicus on the shells, and the formation of the suture line, having wider, low

saddles and lobes with individual bases, situated on well-defined stems.

(Free translation from original Russian. )

402 ANNALS OF THE SOUTH AFRICAN MUSEUM

Beschtubeites Ijin, 1975: 162-163

Type species. Beschtubeites beschtubensis gen. et sp. n. from the sediments of

the Lower Turonian of the lower reaches of the Amudar River.

Diagnosis

Shells high, from compressed to slightly inflated whorl sections. The umbili-

cus is narrow and funnel-shaped. The venter is smooth and flat, or slightly convex.

Umbilical and marginal tubercles are always present in adult specimens. The

flanks are covered with sickle-shaped ribs. In places, bifurcation usually produces

tubercles or tubercle-like swellings. The suture line of the adult specimens consists

of broad, relatively shallow ventral lobes and 8—9 lobes alternating with saddles.

The first lobe is parallel or slightly slanted towards the second lobe. The fourth

lobe is somewhat smaller, rarely equal to the fifth lobe. The fifth lobe is equal or

somewhat larger than the sixth. The first saddle is the highest. The development

of the first umbilical lobe results through separation of its four incisions and

transformation of them into independent lobes (morphologically first four lobes).

In the region of the umbilicus on the second whorl, an umbilical lobe arises which

shifts to the external side. The lobes originating in the area of the umbilicus,

either undergo double division or remain undivided and shift on to the external or

internal side.

Comparison

Of Kopetdagites gen. n., is characterized by the structure of the suture line,

of which the first lobe at least may have a very slight slant towards the second

lobe, but at the same time the base of the lobe is not displaced towards the ventral

side and the first saddle is always higher than the second.

(Free translation from original Russian. )

Diplacomoceras Hyatt, 1900 (1903, p. 242)

‘This genus resembles Engonoceras quite closely in its involute compressed

volutions and channelled venter bordered by continuous ridges, and in the

tuberculation of the sides it is sufficiently near to be included in that genus if the

sutures were unknown. These are, however, well known, thanks to Schliiter’s fine

figures, and they appear to be similar to those of Placenticeras. This genus

therefore combines the external characters of Engonoceratidae with the sutural

lines of Placenticeratidae, and perhaps ought to be placed in a separate family.

The young, however, are unknown, and it is perhaps better to wait until their

development places their affinities on a firmer basis. It is also possible that they

may be accounted for as members of the Placenticeratidae that were arrested in

their development, retaining the neanic condition of the venter and lateral zones

in their later stages, but not arrested in their sutures, which approximate to those

of Placenticeras.’

CRETACEOUS FAUNAS FROM SOUTH AFRICA 403

Stantonoceras Johnson, 1903: 136

‘Shell large in typical species; descended from the same ancestor as Placen-

ticeras; having stages of growth corresponding to ancestral types which possessed,

in successive generations, keels that were very narrowly flat, channelled, and

alternately nodose; but so far advanced at maturity as to bear little or no

resemblance to the typical Placenticeras; outer whorls broadly rounded or

subquadrate in cross-section; moderately embracing; definite ventral keel nearly

or quite obsolete; umbilical angle nearly or quite imperceptible; surface orna-

mented with more or less prominent nodes, elongated nodes or pseudocostae;

septa similar to those of Placenticeras, but most simple and less crowded.

Although somewhat less accelerated than the type of this genus, I am of the

opinion that Ammonites guadaloupa Roemer should be included here. It is of

course one of the transition types, and its generic reference must be more or less

a matter of judgment. But it has advanced so far from the original stock, and has

so many features in common with this genus, that it seems to me impossible to

classify it elsewhere.’

Proplacenticeras Spath, 1926: 79

‘... is of Turonian-Coniacian age and differs in suture-line as well as in

whorl shape and ornamentation.’

Pseudoplacenticeras Spath, 1926: 79

‘This differs from Diplacmoceras Hyatt, by absence of ventral groove, inner

row of tubercles, and less compressed young stage.’

Anaplacenticeras Iljin, 1959: 201

Type species. Anaplacenticeras turkmenense sp. nov. from the Cenomanian

deposits of Kopet-Dagh.

Diagnosis

Shell flat, with high whorl section, deep, step-like umbilicus. Umbilical wall

steep, umbilical edge is sharp. Siphonal side is narrow, slightly concave on the

shells and weakly convex in the middle, falcate ribs. On the umbilical edge are

infrequent, very small tubercles.

Suture line

The siphonal lobe is weakly developed, very short. The first lobe is deep. The

second and third have a common base, are massive and strongly incised. The

fourth lobe is short, significantly smaller than the fifth. Saddles are narrow, low.

The third saddle, subdividing the second and third lobes, is very small and low.

Comparison

According to the shell form, the formation of the umbilicus and the

ornamentation, Anaplacenticeras gen. n. suggests the Albian Cleoniceras cleon

404 ANNALS OF THE SOUTH AFRICAN MUSEUM

Orb., Cl. mangyschlakense Luppov, but the type of suture line and whorl section

of these genera is clearly distinct.

In Anaplacenticeras gen. n. the siphonal side is flat as distinct from the

pointed or strongly rounded one of the genus Cleoniceras Parona & Bonarelli.

The suture line of Anaplacenticeras gen. n. has more massive lobes and

narrower saddles. In Cleoniceras Parona & Bonarelli the ratios are the reverse.

Anaplacenticeras gen. n. is distinguished from the smooth representatives of

the genus Placenticeras Meek (Pl. kharesmense Lah., Pl. arkhangelskii Iljin) by a

stepped form of umbilicus, very small umbilical tubercles, and a distinct umbilical

edge. The last two characteristics connect the genus with the genus Proplacen-

ticeras Spath, but the suture lines in these genera are different.

(Free translation from original Russian.)

Gissarites Ijin, 1959: 727

Diagnosis of the genus

Shell discoidal, involute with narrow umbilicus and wedge-shaped whorl

section. Maximum width is located at the umbilical edge or on the lower third of

the flanks. On internal casts the siphonal side is pointed in the form of a flat keeél;

in shelly preservation it is compressed, narrow, not expanding with the growth of

the whorls and along the edges of it, minute elongated tubercles are located in

zig-zag fashion. The flanks are smooth. Along the umbilical edge a series of high

tubercles is located. Approximately at mid-flank there are weak bulges. The

thickness of the shell differs in different parts of the shell. It reaches maximum

thickness in the region of the umbilicus on the umbilical edge, and along the edges

of the siphonal side. In consequence of this, the umbilicus is more open in the

middle and has steep, high walls, while on the shells the umbilicus is narrower

with relatively sloping walls.

The suture line is of the placenticeratid type, and consists of bulb-shaped

lobes and divided, rounded saddles. The siphonal lobe is wide, shallow, with

short, blunt lateral branches. The lobes on the flanks are well developed. The

deepest of them is the third lobe; the sixth lobe is longer and wider than the fifth

and only a little less deep than the fourth. The saddles are rounded, and one and

a half to two times wider than the lobes. The first saddle is trifid with a

significantly developed middle branch. The remaining saddles are bifid.

Comparison

In contrast to the nearest genus, Placenticeras (Meek, 1876, p. 462),

Gissarites gen. nov. has a tapered siphonal side at the centres and is very narrow;

the siphonal side on the shells not expanding with the growth of the whorls. The

suture line of the new genus is distinguished from the suture line of the genus

Placenticeras by an inverse ratio of the depth of the fifth and sixth lobes. In the

genus Placenticeras the fifth lobe is larger than the sixth; in the new genus the sixth

lobe is larger than the fifth; the lateral branches of the siphonal lobe are short and

blunt in contrast to the paw-shaped ones of Placenticeras.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 405

Species composition and distribution

Two species in the genus are known: Gissarites kysylchense sp. nov. and

Gissarites tagamense sp. nov. They occur in the Maastrichtian deposits of the

south-western spurs of the Ghissarski Range. In the present article the first of the

named species is described, which is taken as the type of genus.

(Free translation from original Russian.)

Parastantonoceras Collignon, 19655: 17

The generic characters are very close to those of Stantonoceras of the

Santonian-Campanian: there are two rows of tubercles, one umbilical of strong

pointed tubercles, one large, blunt, flat and latero-external disappearing on the

body chamber; the external region bears strong, elongate tubercles in two parallel

rows and alternating. But here: the umbilical tubercles are at the summit of the

umbilical wall, the latero-external tubercles on the second third of the flanks and

the tubercles limiting the external region are very strong instead of being small.

On the other hand, the suture shows a reduced number of elements, the saddles

much larger and, as a result, fewer, widely separated lobes.

(Free translation from the original French.)

Asiatostantonoceras Ijin, 1975: 172

Type species. Stantonoceras tagamense I\jin (1959) from the sediments of the

Upper Santonian of the south-western spurs of the Ghissarski Range.

Diagnosis

Shell discoidal, with narrow venter, slanting umbilical wall and inflated

flanks, along the middle of which a row of tubercles is situated. There is almost

no broadening of the venter with development of the shell; its sides are bordered

by low, tooth-like tubercles. In the early stages of development there are three

rows of tubercles on the flanks—umbilical, upper lateral and marginal. Later, the

upper lateral tubercles disappear, but the umbilicals, in proportion to the

flattening out of the umbilical wall, shift behind its rim on to the flanks, and

occupy a central position on it. The suture line consists of a relatively deep ventral

lobe with short lateral branches, and nine lobes alternating with saddles. The first

lobe is shorter than the ventral, the fourth is a little larger than the fifth, the fifth

is larger than the sixth. The saddles, except the first, are rounded and bifid. The

first saddle is the highest, compressed and tapered toward the ventral side and

top. To this genus, beside the type species, one must also refer those described

from the Upper Santonian of Western Germany, A. schliiteri (Schliter, 1872, pl.

14 (figs 2)), A. pseudo-orbignyanum Hyatt (Schliter 1872, pl. 15 (figs 3-4)),

A. bidorsatum Miller, [and] Wolleman & Roemer (Miller & Wolleman 1906,

pl. 3 (fig. 1), pl. 4 (fig. 5), pl. 9 (fig. 1 only)).

406 ANNALS OF THE SOUTH AFRICAN MUSEUM

Comparison

Of the genus Stantonoceras Johnson, characterized by the form of the whorl

section, narrow venters, by strongly developed umbilical tubercles occupying a

central position on the flanks and by the disappearance of upper lateral tubercles

with growth of the shell.

(Free translation from original Russian.)

Baghiceras Chiplonkar & Ghare, 1976: 3

Type species: Baghiceras ambai sp. nov.

‘Diagnosis

Evolute; cross-section broadly elliptical tending to be subcircular to circular;

venter broadly rounded with clavi which may be tuberculate; strong spinose

umbilical tubercles and strong ventrolateral tubercles joined by different patterns

of ribbing; suture somewhat poorly developed placenticeratan type.

Age

Cenomanian—Turonian

Remarks

As compared to other members of the subfamily Baghiceratinae, this genus |

has less developed suture. It differs from Placenticeras (its tumid species) also by

having three rows of tubercles and the umbilical ones not migrating outwards.

From Stantonoceras it differs in having fewer but stronger ventral clavi; while

from the genus Diplacmoceras it differs in not having pinched ear-like ventral

clavi and ventrolateral tubercles. The genus Hoplitoplacenticeras differs from the

present genus in having typically trapezoidal cross-section and by outward

migration of umbilical tubercles.

By its less-developed Placenticeras type of suture this genus is much like the

more ornamented species of Knemiceras (Knemiceras), e.g. Kn. (Kn.) syriacum,

Kn. (Kn.) attenuatum and Kn. (Kn.) gabbi, and appears to have developed out of

Knemiceras (Knemiceras); and Placenticeratidae is considered to have been

derived from Engonoceratidae (Wright, 1957: 390).’

Malwiceras Chiplonkar & Ghare, 1976: 4—5

Type species. Malwiceras variabilis sp. nov.

‘Diagnosis

Rather evolute; early whorls essentially compressed tending to broadly

elliptical to subcircular to circular cross section; umbilical and ventrolateral

tubercles prominent; venter flanked by clavi; ribs weak or absenti suture with few

elements.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 407

Age

Cenomanian to Turonian

Remarks

As exemplified by the type species the cross-section varies with stages

of shell growth from compressed to circular with venter flattish narrow to broad

feebly convex. Suture of typically Placenticeras type has few elements as in

Proplacenticeras.

Fewer but stronger ventral clavi distinguish this genus from Stantonoceras.

Hoplitoplacenticeras by its trapezoidal cross-section clearly differs from this

genus. Compared to Malwiceras the genus Diplacmoceras is distinguishable by its

pinched and ear-like clavi and ventrolateral tubercles. Compared to Baghiceras,

Malwiceras has better developed suture, has no ribs and in its cross-section is

variable with shell growth from compressed to circular. The genus is named after

Malwa, the region in which the Bagh Beds occur.’

Placentoscaphites Chiplonkar & Ghare, 1977: 68-69

‘Genotype. Placentoscaphites dangerfieldi sp. nov.

Diagnosis

Shell moderately large with oval umbilicus; whorls more or less compressed

to convex and tumid; ornamentation of simple low ribs with or without weak

umbilical tubercles and ventral clavi; ventrolateral tubercles if present may tend

to be clavate. Suture placenticeratid but distorted.

Age

Cenomanian—Turonian.

Remarks

The polyphyletic origin of uncoiled ammonoids has been realized since long

by workers like Smith, Nowak, Spath, Reeside, etc., and Nowak has remarked

that ‘whether scaphitid forms do not occur in other families and genera may be

established by future workers’. The provincial development of these in indepen-

dent areas is considered by Cobban and Matsumoto as particularly apparent from

Turonian onwards.

The generic nomenclature such as Holcoscaphites Nowak, Acanthoscaphites

Nowak, Hoploscaphites Nowak and Desmoscaphites Reeside, which is already

currently used, indicates the close affinities of these heteromorphs to the genera

from which they are considered to have been derived.

Association of these heteromorphs with abundantly represented placenticer-

atids, coupled with their placenticeratid sutural pattern, indicates that these Bagh

heteromorphs have placenticeratid affinities. They are placed here, like other

heteromorphs, under the admittedly polyphyletic family Scaphitidae, and their

408 ANNALS OF THE SOUTH AFRICAN MUSEUM

apparently having been derived from placenticeratid stock has led us to name the

genus Placentoscaphites.’

Sancarlosia Chiplonkar & Ghare, 1978: 79

Type species. Placenticeras sancarlosense Hyatt, 1903 (pl. 30

(figs 1-3)).

‘Diagnosis

Form tumid; sides feebly convex, venter broad with alternating clavate

tubercles; tubercles at shoulder tending to be bullate; umbilical tubercles tending

to be bullate and migrating mid-ventrolaterally; suture Placenticeras-like with first

adventitious saddle in first lateral saddle broader than others.

Age

Senonian.

Genus is named after locality of origin, i.e. San Carlos (Mexico).’

NOTE ADDED AT PROOF STAGE

A new genus, Rapidoplacenticeras Alabushev, 1988 (type species Proplacen-

ticeras sutherlandbrowni McLearn, 1972), has been described from the Upper

Albian. It is said to differ from (Pro-) Planticeras by its lack of ornament, narrow

umbilicus and more incised suture line.

ALABUSHEV, A. I. 1988. Rapidoplacenticeras—a new genus of Cretaceous ammonoids. Paleon-

tological journal 1988 (1): 109-112, 3 figs.

SYSTEMATIC papers must conform to the International code of zoological nomenclature (particu-

dy Articles 22 and 51).

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

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

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

nd not be abbreviated; if the year is added, a comma must separate author’s name and year. The

uthor’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

erred from its original genus. The name of a subsequent user of a scientific name must be separated

rom the scientific name by a colon.

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

james 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

‘Nucl (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

he 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.

i 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

| (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 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.

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

THE AMMONITE FAMILY

PLACENTICERATIDAE HYATT, 1900;

WITH COMMENTS ON THE

SYSTEMATIC POSITION OF THE GENUS

HYPENGONOCERAS SPATH, 1924

ME 98 PART

Mi 10

| 0)

CTOBER 1989

MCZ

| . ISSN 0303-25

—2515

HARVARD

UNIV aoa

THE SOUTH AFRICAN

MUSEUM

\PE 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

Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

bered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

into consideration.

All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 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.:

‘Smith (1969) describes . . .’

‘Smith (1969: 36, fig. 16) describes. . .’

“As described (Smith 1969a, 19696; Jones 1971)’

‘As described (Haughton & Broom 1927)...’

‘As described (Haughton et al. 1927)...”

Note: no comma separating name and year

pagination indicated by colon, not p.

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, 1969b) 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 (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number 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 la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHER, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHULTzE, L. Zoologische und anthro-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 ~ Band

October 1989 Oktober

Part 10 #£Deel

LANTERNFISHES OF THE SOUTHERN

BENGUELA REGION. PART 3.

THE PSEUDOCEANIC-OCEANIC INTERFACE

P. ALEXANDER HULLEY

J. R. E. LUTJEHARMS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

1, 2(1-3,-5-8) 30-2, 4-5. 8, t=pa), SE oe

6(1; tpa.), TAA 8, 9-27), AOC 3):

11(1-2, 5, 7, t-p.i.), 14(1-2), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33, 36(2), 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 104 0

Printed in South Africa by In Suid-Afrika gedruk deur

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

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

LANTERNFISHES OF THE SOUTHERN BENGUELA REGION.

PART 3.

THE PSEUDOCEANIC-OCEANIC INTERFACE

P. ALEXANDER HULLEY

Department of Marine Biology, South African Museum, Cape Town

J. R. E. LUTJEHARMS

Earth, Marine and Atmospheric Science and Technology, CSIR, Stellenbosch

(With 8 figures and 7 tables)

[MS accepted 24 February 1989]

ABSTRACT

The distributions of lanternfishes (Myctophidae) from various cruises of the R.S. Africana

off the west coast of southern Africa were examined in order to elucidate the pseudoceanic—

oceanic interface in the eastern South Atlantic. Thirty-six night RMT-8 hauls to 120 m and

along nine transect lines across the southern African west-coast continental shelf/slope

(25 °32'-34°55'S_ 12°27'-17°35’E) were made during the Phyllosoma Survey in August 1984

(winter). An investigation of cross-shelf/slope zonation is undertaken using lanternfish abun-

dances and the Bray-Curtis similarity measure with group average sorting and multi-

dimensional scaling. Two groups of stations are recognized. Their clustering is correlated to

bottom depth and their interface is best demarcated by the 800-m isobath. The ‘in-shore’ group

is characterized by the pseudoceanic species Lampanyctodes hectoris, whereas ‘off-shore’ group

indicator species include the oceanic, mesopelagic myctophids Ceratoscopelus warmingii,

Diaphus hudsoni and Diaphus meadi. Examination of additional data suggests a seasonal, off-

shore displacement in the distribution of Lampanyctodes hectoris. During the summer upwel-

ling period the species occurs mainly in-shore of the 300m isobath and its off-shore

distributional limit approximates the 500 m bottom contour. In winter its distribution extends to

the 1 000 m isobath but it may be taken at lesser densities over depths in excess of 3 000 m.

Similar seasonal patterns are not evident for oceanic myctophids, suggesting that water-column

depth and the potential to undertake diurnal migration may be the major factor governing

the shoreward distribution of these oceanic species. In the southern Benguela region at least,

the seasonal difference in the distribution of Lampanyctodes hectoris appears to be geared to

the effect of frontal dynamics on the availability of food in summer, and to a migration into

deeper waters during the winter—spring spawning period.

CONTENTS

PAGE

REPRE ORM ee ee oe halen cs hea, wtirdne OS 58 Sm 410

ier Gittins AIM MCENOGS . 2. i055 dcx cs ce Ree ea we dee cenvcasuseswwse 411

cage PIN eae oo 2 on os Sis carat ARS He aN ES See’ end thaw Oe 417

EU UISSOE cig ESE a ae en en ga en ee ee er eee 420

ERIN PEMAMCTNS Ae lnc see nes el op A aN oe ea eee et 431

Don) EGER DES 8 Loe ek 6 gil Rp a ee ee 431

409

Ann. S. Afr. Mus. 98 (10), 1989: 409-435, 8 figs, 7 tables.

410 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

The existence of a frontal system close to the shelf break in the southern

Benguela region (Bang 1971, 1973a, 1973b; Bang & Andrews 1974), and the

association of frontal zones with increased biomass and productivity, have

recently given impetus to the study of the physical oceanography of the mixing

area seawards of and parallel to the main coastal upwelling core off the west

coast of southern Africa (Brundrit 1981; Shannon et al. 1983; Van Foreest et al.

1984; Shelton et al. 1985; Lutjeharms & Meeuwis 1987; Lutjeharms & Stockton

1987). Allied biological research on chlorophyll, copepod and fish distributions

have been reviewed by Shannon & Pillar (1986) and Crawford et al. (1987),

while Shelton (1986) has commented on the significance of the front to fish repro-

ductive strategies. The present paper deals with a preliminary investigation of

the biological effects of the interaction between bottom topography and large-

scale hydrographic features on the distribution of the predominantly oceanic

family Myctophidae in the eastern South Atlantic.

The continental slope not only limits the landward distribution of the

Oceanic mesopelagic and bathypelagic ichthyofauna (Pearcy 1964; Badcock

1981; Hopkins et al. 1981; Merrett 1986; Parin 1986), but also provides an

environment for the development of a recognizable and trophically dependent

community of benthic and benthopelagic fishes (Parin & Golovan 1976; Mar-

shall & Merrett 1977; Ratz 1984). However, pseudoceanic derivatives of certain

fish families, including the Myctophidae, may occupy the region above the upper

continental slope and inshore of the continental shelf break (Nafpaktitis &

Paxton 1968; Krefft 1970; Kawaguchi & Shimizu 1978; Nafpaktitis 1978; Hulley

1981, 1986b; Bekker 1983; Rubiés 1985; Hulley & Prosch 1987). These deriva-

tives include both obligatory pseudoceanic species and facultative pseudoceanic

species, the former often the numerical dominants of the pelagic slope popu-

lations (Merrett 1986).

Although Lampadena pontifex has been caught sporadically south to

28 °30’S and Diaphus taaningi south to about 24°S off the west coast of southern

Africa (O’Toole 1976; Hulley 1981, 1986a, 1986b; Rubiés 1985; SAM data), and

although isolated specimens of Diaphus garmani have been recorded at about

34°S in Agulhas Water, only a single, numerically dominant, pseudoceanic,

myctophid species, Lampanyctodes hectoris, is known from the southern

Benguela region (Hulley 1986b; Prosch 1986). The latter species has been fished

commercially and has accounted for up to 9,4 per cent (42 400 metric tons) of

the total annual pelagic catch by the South African purse-seine fishery (Hulley &

Prosch 1987).

Few sampling cruises have been undertaken off the west coast of southern

Africa to provide specific data for analyses of the pseudoceanic—oceanic inter-

face, but lanternfish specimens collected in August 1984 during the Phyllosoma

Survey (Cruise 023) by the Sea Fisheries Research Institute, Cape Town (SFRI)

allow for an investigation of slope—shelf distributional phenomena during their

SOUTHERN BENGUELA LANTERNFISHES 411

winter manifestation. However as a caveat, it must be pointed out that hydro-

graphic conditions during August 1984 may be atypical of winter, since the

period of sampling fell within the second phase of the 1984 Benguela El Nino

event (September 1983-January 1984) (Shannon et al. 1986). The extended

warm cycle apparently continued through to 1986 (Shannon & Agenbag 1987),

although the warming in 1984 was followed by substantial cooling between

November 1984 and October 1985 (L. V. Shannon, SFRI, pers. comm.).

Myctophid data sets from additional SFRI cruises are also analysed in order

to examine the possibility of distributional variability on a seasonal basis. The

results represent average, seasonal, large-scale patterning and do not purport to

examine the high-frequency variability that characterizes the southern Benguela

region, particularly during its summer upwelling phase (Shannon 1985).

MATERIALS AND METHODS

During the 1984-Phyllosoma Survey (SFRI Cruise 023) from 14 August to

24 August, 36 stations, situated on nine transect lines extending across the shelf

break (Table 1, Fig. 1), were occupied off the west coast of southern Africa by

R.S. Africana with an opening-closing RMT-8 net (Baker et al. 1973). For each

oblique haul, the water temperature was recorded at 5-m depth intervals by a

probe housed in the Universal Underwater Unit (U*), which also operated the

net-closing mechanism. From this record temperatures were extracted at 10-m

depth intervals for depths between 0 m and 40 m, and at 20-m depth intervals

for depths between 40m and 120m (Table 2). In addition, six CTD casts

(Fig. 1: CTDi—CTDe) were made to investigate the density structure in the off-

shore region and its variation with latitude. Temperature profiles at these

stations are given in Figure 2.

Due to malfunction of the jaw mechanism of the U? device that caused

premature (and unpredictable) net closure (Pollock 1984), data from stations

1 (1) through 2 (4) are unreliable and have therefore have not been taken into

account in the ensuing analyses; these stations yielded 153 lanternfish specimens

comprising nine genera and 15 species. No specimens were caught at station

3 (1) due to gear failure and the haul at station 9 (4) was aborted; these stations

are also not included in the analyses.

The location of the main upwelling fronts for the months August 1983 and

August 1984 were established by using thermal infra-red images from the radio-

meter on board the Meteosat II satellite. The instrument measured in the

10,5—12,5-m waveband and daily data sets were appropriately contrast-enhanced

for this ocean area. The spatial resolution in the area was less than 5 km by

5 km. No atmospheric corrections were made, so that absolute temperatures

were not available from the imagery, but temperature profiles for the stations

from the August 1984 cruise are given in Table 2. Experience has shown that

the main upwelling front is sufficiently distinct so that its identification is

412 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

RS. Africana Cruise 023: Station data. Pe = bottom depth; a No. = computer generated

number; Duration = duration of haul; FDepth = fishing depth; LMT = local mean time (hours:min-

utes:seconds); Speed = ship’s speed.

St. Comp. Position Date LMT Duration FDepth Speed BDepth

No. No. (mins) (m)_ (knots) (m)

25°32,18'S = 13®33,76'E —s_: 15.08.1984 20:18:19 60 120-20 2,60 525

25°33,00°S = 13°11,75°E_—_: 15.08.1984 = 23:15:00 60 120-20 2,40 1561

2531,91’S 12%9,9VE 16.08.1984 01:56:26 60 120-20 2,50 2500

25°29,97S 12°27,2V7E 16.08.1984 04:51:44 60 120-20 2,50 3060

26°39,75°S_ —- 13°33,02’7E_—_: 17.08.1984 06:32:45 60 120-0 2,80 950

26°40,80’S_ —s- 13°13,20"E_—: 17.08.1984 = 03:26:52 60 120-0 2,00 2000

26°44,00°S 12°54,00°7E = 17.08.1984 00:08:00 60 120-0 230553399

26°46,00’S_ 12°39,007E 16.08.1984 19:44:00 60 12020 “= Zz

28905,24S 14°29,62’7E_—_: 14.08.1984 20:05:02 60 120-20 2,80 598

28908,00’°S_ —- 14907,00°E —-_: 14.08.1984 =. 23:12:42 60 120-20 ZeQ@tZ7

28°07,00'S_ = 13°43,007E—-: 15.08.1984 =: 02:20:58 60 120-20 2,60 1433

28°09,10°S_ ——:13°19,35’°E_—s: 15.08.1984 = 05:32:47 60 120-20 2,60 1300

28°52,00’S_ — 14°21,007E_—-: 17.08.1984 =. 20:11:00 60 120-0 250 550

29°00,00’°S_ — 14°00,00°E_—-: 17.08.1984 — 23:09:00 60 120-0 2,10 1491

29°05,40’S_ — 13°38,527E_—-: 18.08.1984 02:07:46 60 120-0 2505

29°09,75’°S_ - 13°15,86’E_—_: 18.08.1984 = 05:00:53 60 120-0 2,50 2950

30°32,00’S_ — 15°17,00E_—_: 19.08.1984 ~—_- 05:30:00 60 120-20 2,50 352

30°37,80°S 14°55,10'E_—_: 19.08.1984 = 03:28:00 60 120-20" 250° "1300

30%42,74S 14°32,637E 18.08.1984 22:57:00 60 120-20 2,50 2200

30%48,37S 14°10,727E_— 18.08.1984 = 20:09:00 60 120-20 2 257

hWN— WN WN WN

—

PWN

31°44,00°S = 15°48,00°E —-. 20.08.1984 —- 05:20:00 60 120-0 2,50 742

31°49,00’'S_ —- 15°30,00°E —_. 20.08.1984 ~—- 02:30:00 60 120-0 250 1750

10 31°55,23’'S_ = 15909,46’E_—s_ 19.08.1984 —- 23:11:00 60 120-0 250 2470

11 31°59,93°S =: 1451,06E —-: 19.08.1984 ~—. 20:46:00 60 120-0 250) 22800

12 32°2,00’S = 1653,007E_—_ 21.08.1984 = 03:55:00 60 120-20 2,50 408

3259,50’'S_ —- 16°34,00’E—_ 21.08.1984 —- 01:08:00 60 12020 250 > 1500

13 33°05,14S = 1612,31’E ~—- 20.08.1984 = 22:05:00 60 120-20 2,50 2540

14 33°12,02S 15%9,4VE 20.08.1984 19:25:00 60 120-20 “2503 60

15 33°33,61’S 1732,2VE 22.08.1984 20:00:00 60 12020 2,50 404

16 33°43,89°S 17°11,007E ~—- 22.08.1984 23:04:00 60 120-20 2,50 837

17 33%1,00’S = 16°50,00’E —- 23.08.1984 —_ 02:30:00 60 120-20 2,50 2450

18 34°00,00’'S = 16°28,00°9E =—_ 23.08.1984 —- 05:52:00 60 120-20 2,50 3120

19 34°19,20'S = 1735,30°E— 24.08.1984 04:50:00 60 120-20 2,50 760*

20 34°31,50’S 17°16,50°E = 24.08.1984 02:08:00 60 120-20 2,50 2600*

21 34°43,68'S 16°5,00’7E =. 23.08.1984 22:51:00 60 120-20 2,00 3 100*

3454,95’°S _ 16°36,527E_—-_ 23.08.1984 — 20:08:00 60 120-20 2,80 3.600*

hWN PWN

den ea ee ae ee ee ee ee ee ee eee ee eee

SL RAE 7 a a a NL LN LL LN LL LLL

WN =

RWN

* = bottom depth plotted from Dingle et al. (1987).

independent of the enhancement algorithm employed (Shannon et al. 1985).

Daily images were scrutinized and the locations of the fronts on all days that

were sufficiently cloud-free were superimposed (Fig. 3).

In order to examine the possible effects of seasonality on the distribution of

lanternfishes in the southern Benguela region, additional myctophid data from

SOUTHERN BENGUELA LANTERNFISHES 413

CRUISE 023

—e RMT-8

EB so 8,

Luderitz

Port Nolloth

Fig. 1. R.S. Africana Cruise 023: August 1984. Station positions. Depth intervals given in

metres. 103.

the following SFRI cruises have also been scrutinized: Cruise 002— August

1982: 29°12’—33°43'S, 13°04'-17°40,7’E (RMT-2); Cruise 009—May 1983:

28 °38,71'—34 °56'S, 14°34,12'—18°19,38’E (RMT-8); Cruise 011— August 1983:

29 °29'—34°56'S, 14°06'-17°35'E (RMT-8); Cruise 022—July 1984: 28°55’—

35 °59'S, 14°50’—19 °58’E (BT-180); Cruise 028—January 1985: 28°49'—36°01'S,

14°46’—19°50’E (BT-180); Cruise 033—July 1985: 29°04’—36°00'S, 15°17,3’-

414 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

R.S. Africana Cruise 023: August 1984. Variation of temperature with depth.

Station Temperature (°C) at depth

No. om 10m 2m 30m 40m 60m 80m 100m 120m

102) 232 2831 182 B81 =20 36 27 | eee

12) 235 134° 234 834 £42134 +133 "13.0% 91 seen

13) 82 232 ° 132 “Wo 9 ios ees i 11,8. ¢eetis

38 13,8 13,8 13,8 13,8 13,6 13,7 133 12,3

14,3 14,2 14,3 14,3 14,3 14,4 14,4 14,0 13,2

15,0 15,0 15,0 15,0 15,0 15,0 14,8 14,3 13,9

15,1 5,1 15,1 14,8 14,5 14,1 13,8 12,4 11,0

14,8 15.7 15,7 15,8 15,8 15,8 15,7 15,8 14,2

16,0 16,0 16,0 16,0 16,0 15,8 15,7 15,7 15,0

15,6 15,6 15,6 15,6 15,6 15,6 15,6 15,6 15,0

14,9 14,9 14,9 14,7 14,6 14,2 13,7 132 11,4

14,9 14,9 14,8 14,7 14,6 14,4 13,9 13,4 12,3

15,6 15,6 15,6 15,6 15,4 14,8 14,6 14,1 13,6

15,6 15,6 15,6 15,6 15,6 15,6 15,6 15,4 13,5

15,1 15,1 15,0 15,0 15,0 14,9 14,7 14,4 13,8

15,6 15,6 15,5 15,5 15;5 155 15,5 15,4 15,2

iS:7 15,6 15,4 15,4 15,5 15,4 15,3 1533 14,9

15,3 1557, 1557 7 15,7 15,6 15,5 15,2 15;1

—

Nee Nee Nee’ eee” a _ Nee Nee Nee eee” New Nee eee Nee” a ae ee eee Nee eee” Nee eee ee ee” Ne eee ee ee a a

OOO 6 OO 00 00 OO ~~~ NNN N Aman HHH WWW WW NNN NY

1 155 15,4 15,2 15;1 15;1 15,0 14,9 14,6 14,0

2 15,4 15,3 15,2 15,1 15,1 14,9 14,8 14,4 13,7

3 15,4 2 15,2 15,2 15:2 15,2 15,1 14,8 14,1

+ 15,6 15,4 153 15,3 15,2 14,9 14,8 14,6 14,2

1 15,3 15,5 15,2 15,1 15,1 15,1 15,1 15,0 14,4

2 15,6 1555 15,3 15,3 153 15,1 15,2 15,0 14,6

3 15,4 53 15,2 15,2 15,2 15,2 15,1 14,9 14,9

4 15,7 15,6 15,5 15,5 155 15,3 15,3 15,2 15,2

1 15,5 15,5 15,4 15,3 15;2 15,4 14,7 13,7 13,3

2 15,3 15;3 15,2 15,1 15,1 [5,1 14,9 15,0 13,9

5 1555 155 1555 15,5 15,4 15,3 15,2 15,2 15,0

+ 15,4 15,4 15,4 53 15,3 15,2 15,1 15,1 15,0

1 15,1 15,1 15,1 15,1 fot jis 15,0 15,0 15,0

2 15,2 15,2 15,2 15,2 15,2 15,1 15,0 14,8 14,3

3 15,1 15,1 15,1 15,1 ist 14,9 14,8 14,5 13,6

+ iS 15,1 15,1 15,1 14,9 14,6 14,2 123 12,0

19°58,5'E (BT-180); Cruise 039—January 1986: 24°28,6’—35°59’S, 13°39,6'—

19°53,2'E (BT-180); Cruise 046—July 1986: 28°50'-35°55'S, 14°15,1’-

19°49,6’E (BT-180); Cruise 050—January 1987: 28 °27,5'—35 °44,5'S, 14°15,1’-

19°51’E (BT-180); Cruise 054—28 °22,1'-36°06,8’S, 14°36,9’-20°00,6’E

(BT-180); Cruise 060—March 1988: 32°53,9’'-35°09’S, 17°20'-19°29,7'E

(BT-180, Engels—308).

415

‘[ oInsLy Ul se SuLIoquINnN

‘suonejs GLO wor sayyoid oinjyerodwiay, “pg6, isn3ny :¢79 asin vuvIf{y ‘Sy CZ BI

00S 00L

0001

006

ie 009

2) 004 008

jee

eal 00S

ca 004

=] aS uv ao)

o0E 2 a ee)

2 a 009 2

< vw nw

n 007 H wn

s 5 = ¢

Po)

~ a s aLo a 006 fy

a a a

Gi z Z e

“ OOE 7) Bm

jaa} a)

Zz 9 G19 002 00%

jaa]

= 002 00E

a2)

e 001

002

00L

0 0 0

0'0z 0'SL 0'0L 0's 0'0 0'02 0'SL 0'0L 0's 0'0 00L 0S

Jo JYNLVYAdWAL Jo JUNLVYAdWaL Jo JUNLVYAdWIL

416 ANNALS OF THE SOUTH AFRICAN MUSEUM

LOE 20°E

AUGUST 1983

e Cruise 011

Walvis Bay

Luderitz

10°E Z0%E

AUGUST 1984

Walvis Bay e Cruise 023

Luderitz

ape Town

Fig. 3. Montage of locations of the outer limits of cold upwelled water (fronts) as evident in

thermal infra-red satellite imagery from Meteosat II of the south-east Atlantic in relation to

station position for: (A) Cruise 011—August 1983; (B) Cruise 023— August 1984.

SOUTHERN BENGUELA LANTERNFISHES 417

RESULTS

The 26 RMT-8 hauls yielded a total of 904 myctophids; the numerical dis-

tributions of these are given in Table 3 in relation to the bottom depth of the

particular station, and as pseudoceanic or oceanic species in accordance with

Hulley (1981). A similar breakdown of the lanternfish data (1 085 specimens)

obtained during the SFRI Cruise 011 (Phyllosoma Survey, August 1984) with

the identical RMT-8 net and during stratified sampling between 600 m and 0 m,

is also given in Table 3.

TABLE 3

Catch rates (specimens.hour’) for pseudoceanic and oceanic species of Myctophidae

from RMT-8 night hauls during SFRI Phyllosoma Surveys in August 1983 and August

1984. No. spp. = number of species.

Bottom PSEUDOCEANIC OCEANIC

depth

stratum No. No. Catch rate No. Catch rate

(m) hauls spp. (specs.h) spp. (specs.h1)

AUGUST 1983

<500 1 1 26,0 3 6,0

500-1 000 2 1 75,0 4 a5

1 000-2 000 5 1 2,4 29 56,4

2 000-3 000 4 0 0,0 26 109,4

>3 000 1 0 0,0 16 166,0

AUGUST 1984

<500 %) 2 83,7 0 0,0

500-1 000 4 1 20,0 11 13,8

1 000-2 000 8 | 0,4 23 Sut

2 000-3 000 8 1 0,5 20 24,5

>3 000 3 0 0,0 12 22,0

AUGUST 1983 + AUGUST 1984

<500 4 2 69,3 3 1,5

500-1 000 6 1 38,3 12 10,3

1 000-2 000 13 1 £2 if 40,8

2 000-3 000 12 1 0,3 32 SPAGS

>3 000 4 0 0,0 20 58,0

The samples from Cruise 023 resulted in a data matrix of 33 species at

26 stations. The catch rates of myctophids at five of these stations were less than

10 specimens.h"!. Preliminary cluster analysis of the entire data set resulted in

the formation of five isolated groups, whose similarity levels were low, i.e.

Outlier groups, and which tended to obscure the major groupings (fide Hulley &

Krefft 1985: 43). These five stations were therefore excluded from further analy-

sis. They include: station 3 (3) (4 specimens.h''); station 4 (3) (2 specimens.h");

station 6 (1) (3 specimens.h"); station 6 (2) (6 specimens.h"); and station 7 (2)

(3 specimens.h"). The reduced data matrix comprised 31 species at 21 stations

and was analysed according to the method described by Field et al. (1982), and

418 ANNALS OF THE SOUTH AFRICAN MUSEUM

employed by Hulley & Krefft (1985) for transect analyses of lanternfishes from

the 1979-Sargasso Sea Expedition. Standardization of the raw data matrix was

unnecessary because the duration of each haul was 60 minutes.

The dendrogram given in Figure 4 shows the station affinities based on

root-root transformed abundances of the 31 species included in the reduced data

matrix. The broken line drawn at the arbitrary similarity level of 20 per cent

delineates two major groups of stations (Group I and Group I). The ordination

of the similarity matrix using MDS (Fig. 5) gives the same groupings as the den-

21 5 7 16 16.11 10 1°20 8 17 13 7 14 & 9 “19 tees

—

= 50

GROUP | GROUP II

Fig. 4. R.S. Africana Cruise 023: August 1984. Dendrogram of station affinities for RMT-8

hauls.

SOUTHERN BENGUELA LANTERNFISHES 419

drogram. Although the formation of two subgroups among the Group I stations

is indicated at 38 per cent, these groupings are not evident in the ordination and

the level of similarity is low when compared to the dichotomy of the Group II

stations (44%). MDS-ordination (Fig. 5) supports this contention and suggests

that clustering within the Group I stations is ambiguous. Consequently the

further classification of the data sets was not undertaken. It should be noted that

levels of similarity in the dendrogram are generally low and are the result of the

large number of zeros for the less-common species of the matrix.

Results of information statistic (I—) tests for Groups I and I, which are cal-

culated from the numbers of specimens, are presented in Table 4. While 2A]i

values should only be employed as a rule of thumb for cut-off limits (Field et al.

1982), the data in Table 4 indicates two perfect indicator species for Group I,

namely Diaphus hudsoni and Diaphus meadi. In addition, high 2 AI values for

Lampanyctodes hectoris (826,880) and Ceratoscopelus warmingii (77,732)

confirm their identities as Group II- and Group I-indicator species, respectively.

As in the classification and ordination procedures, ambiguous results were

obtained for the suggested subgroups within Group I. Relatively low values for

2 AI; (maximum 15,249) were obtained in comparison to those in Table 4 and to

those from the Sargasso Sea analyses (maximum 25 026,911) presented by

Hulley & Krefft (1981, tables 4-6, 9-11).

CRUISE 023

MDS (ROOT-ROOT TRANSFORMED)

20% SIMILARITY

GROUP |

GROUP Il

Fig. 5. R.S. Africana Cruise 023: August 1984. MDS-ordination of station affinities for

RMT-8 hauls.

420 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

RS. Africana Cruise 023: August 1984. Frequencies of occurrence (F) and numbers of

individuals (N) of species, ranked according to information statistics, which distinguish

Group I (Fi, N1) from Group II (F2, Nz). 2Ali values calculated from numbers of

specimens. Species above horizontal dotted line have 2Ali>6,63; those below line have

2Ali>3,84; species with 2ATi<3,84 are not included. Maximum values for F1 and F2 are

given in parentheses.

Fi Ni F2 N2 2Ali

(16) (5)

Lampanyctodes hectoris 4 Wei B 324 826,880

Diaphus meadi 15 236 0 0 128,353

Ceratoscopelus warmingii 15 160 1 1 ThLY2

Diaphus hudsoni Oe Be 30 e

Lobianchia dofleini 6 11 0 0 5,983

Lampanyctus lepidolychnus 5) 19 1 1 5,263

Lampanyctus alatus 4 2 0 0 4,895

Lampanyctus pusillus 4 8 0 0 4,351

Notoscopelus resplendens 5 8 0 0 4,351

DISCUSSION

Since all hauls from Cruise 023 were of 60 minutes duration, were hauled in

a uniform manner (D. E. Pollock, SFRI, pers. comm.) and were all fished

obliquely at night between 120-0 m or 120-20 m at towing speeds between

2,0 knots and 2,8 knots, and since chemical and productivity measurements

were not taken during the cruise, bottom depth and temperature data are the

only variables that can be correlated to the station clustering. Temperature data

sets at selected depths superimposed on the MDS-ordination indicate little cor-

relation with the station groupings. Examination of the CTD traces (Fig. 2)

reveals that the RMT-8 oblique hauls were sampling the upper mixed layer of

the water column at all stations and that the lower limits for the mixed layer (top

of the thermocline) were similar for casts from both inshore (CTDs = 108 m;

CTD¢s = 122 m) and offshore (CTD: = 132 m; CTD2=120 m; CTD;= 108 m;

CTD, = 138 m) of the 1 000-m isobath.

A better correlation results from the superimposition of the bottom depth

data on the MDS-ordination. The initial delimitation of three bottom depth

strata demonstrates a gradation from quadrant to quadrant, with shallowest

depths in the Group II stations and with depth strata > 1 000 m confined to the

Group I stations. The close proximity of station 8 (2) (bottom depth 837 m) of

Group I to station 9 (1) (bottom depth 760 m) of Group II should be noted.

This suggests that the change in faunal composition lies at some intermediate

depth. The correlation of the groupings to the 800-m isobath (Fig. 6) yields the

best result, with Group I stations fished over bottom depths in excess of 800 m

and Group IJ stations over bottom depth less than 800 m. Geographic plots of

Group I and Group II stations are given in Figure 7. Indicator species for the

groupings reinforce these findings. Group I indicator species (Ceratoscopelus

SOUTHERN BENGUELA LANTERNFISHES 421

CRUISE 023

MDS (ROOT-ROOT TRANSFORMED)

BOTTOM DEPTH

GROUP |

GROUP II

Fig. 6. R.S. Africana Cruise 023: August 1984. Bottom depth data superimposed on

MDS-ordination for RMT-§8 hauls.

warmingii, Diaphus hudsoni, Diaphus meadi) are oceanic species, whereas the

sole indicator species for Group II (Lampanyctodes hectoris) is a pseudoceanic

species characteristic of the shelf area of the southern Benguela region (Hulley

1981, 1986a, 1986b; Hulley & Prosch 1987).

Therefore the landward distribution of oceanic mesopelagic myctophid

species and the seaward distribution of pseudoceanic myctophid species in the

central and southern Benguela regions (28°S—35°S) would appear to be gov-

erned mainly by the depth of the water column, with the major faunal change

demarcated by the 800 m bottom contour. Across the west Florida continental

slope, Hopkins et al. (1981, fig. 2) have indicated that the greatest decline in

Oceanic species occurred where bottom depths shallowed from 500 m to 275 m.

Over the Oregon slope off Newport, Pearcy (1964) has observed this decline

where bottom depths shoaled from about 1 000 m and has related this to an

inability by mesopelagic fishes to complete vertical migration cycles as the

bottom shallows, and the effects of shallow depths on vertical temperature struc-

ture and the photo-environment.

Examination of the RMT-8 myctophid data from the 1984-Phyllosoma

Survey (Table 3) reveals that catch rates for the two pseudoceanic species,

Lampanyctodes hectoris and Diaphus garmani, declined evenly from 83,7 speci-

mens.h' over the <500-m bottom depth stratum to 20,0 specimens.h! over

501-1 000 m, and dropped abruptly to 0,4—0,5 specimens.h" over the 1 000-

2 000-m and 2 000-3 000-m bottom depth strata. The species were absent from

422 ANNALS OF THE SOUTH AFRICAN MUSEUM

CRUISE “O23

@ Group I

e's

Luderitz

o not included

Port Nolloth

Fig. 7. R.S. Africana Cruise 023: August 1984. Geographic plots of RMT-8 hauls showing

distribution of Group I and Group II.

the water column where bottom depths exceeded 3 000 m. In contrast, oceanic

myctophid catch rates were more-or-less constant (22,0-31,1 specimens.h") over

bottom depths in excess of 1 000m and declined to 13,8 specimens.h! over

501—1 000 m. Oceanic species were absent where the depth of the water column

was less than 500 m. A comparative data subset (13 night hauls) from the 1983-

Phyllosoma Survey, during which myctophids were taken in discrete hauls fished

SOUTHERN BENGUELA LANTERNFISHES 423

in the upper 250 m of the water column, is presented in Table 3. Here catch

rates for the pseudoceanic species, Lampanyctodes hectoris, declined from

26,0-75,0 specimens.h"! at depths shallower than 1 000 m to 2,4 specimens.h"!

over the 1 000-2 000-m depth stratum, and the species was absent over the

2 000-3 000-m and >3 000-m depth strata. In contrast, catch rates for oceanic

myctophid species declined evenly shorewards from a value of 166,0 speci-

mens.h! (>3 000m) to 56,4 specimens.h?! (1 000-2 000 m) and decreased

abruptly over depths shallower than 1 000 m.

Pooled data sets from both cruises (Table 3) allow for a more comprehen-

sive examination of the oceanic species spectrum at shallower depths. Whereas a

few specimens of three oceanic species (Diaphus diadematus, D. hudsoni, Lam-

panyctus alatus) were taken over bottom depths shallower than 500 m,

12 myctophid species were taken over depths of 500-1 000 m. The latter include

Ceratoscopelus warmingii, Diaphus diadematus, D. hudsoni, D. meadi, D. mol-

lis, D. perspicillatus, Hygophum hygomii, Lampanyctus achirus, L. australis,

L. lepidolychnus, Notoscopelus_ resplendens, and Symbolophorus boops.

Oceanic species diversity increased from six species inshore of the 1 000-m

isobath to 13 species over bottom depths greater than 1 000 m.

Similar trends in the pseudoceanic—oceanic faunal replacement were

evident in the myctophid data from BT-180’ hauls undertaken during the July

1984—January 1986 SFRI Hake Surveys (Hulley & Prosch 1987, table 2). During

this period, oceanic species were never taken over bottom depths less than

301-400 m: shallowest bottom depth 378 m (Symbolophorus boops) during the

winter cruises; shallowest bottom depth 354m (Scopelopsis multipunctatus)

during the summer cruises. On the other hand, Lampanyctodes hectoris was

taken over bottom depths greater than 500 m in July 1984 and July 1985, and

was confined to hauls fished shallower than the 401-500 m depth range in

January 1985 and January 1986. While this suggests the possibility of a seasonal

shift in the position of the outer boundary limit for Lampanyctodes hectoris,

these data should be regarded with reservation. It must be noted that:

(1) although Roel (1986: 578) has suggested that the presence/absence of Lam-

panyctodes hectoris in BT-180' hauls is ‘representative’, this gear may not

sample the species efficiently; (2) few stations were occupied at depths greater

than 500 m during the Hake Surveys; and (3) in contrast to the grouped data

sets, absolute maximum values for this species during these cruises, and in three

subsequent cruises, were similar: Cruise 022 (July 1984)—515 m (St. A 1563);

Cruise 033 (July 1985)—502 m (St. A 2756); Cruise 039 (July 1986)—456 m

(St. A 4307); Cruise 054 (July 1987)—460 m (St. A 5899); Cruise 028 (January

1985)—488 m (St. A 2273); Cruise 039 (January 1986)—456 m (St. A 3348);

Cruise 054 (January 1987)—467 m (St. A 5306).

However, unpublished data from R.S. Africana Cruise 060 off the south-

west Cape coast in March 1988 provide additional evidence for the seasonal shift

in the distribution of Lampanyctodes hectoris. Catch statistics for this species

from two daylight and two night lines of stations, which were run in both the

424 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cape Canyon (SW Cape Columbine) and the Cape Point Canyon (SW Cape

Point) regions, are given in Table 5. For this same cruise, catch statistics for five

midwater Engels—308 hauls on acoustically identified targets are also given in

Table 5. Firstly, these data indicate that the BT-—180’ net does not effectively

sample Lampanyctodes hectoris: catch rates with this gear in the 300-m and

500-m bottom depth spectra averaged 0,23 kg.h' and 0,003 kg.h"' respectively,

whereas catch rates at similar bottom depths with the Engels—308 net were

66,40 kg.h! and 0,36 kg.h'! respectively. Targeted water-column hauls with the

Engels—308 were 15,20 kg.h' over 300m, and 0,01 kg.h! over 500m. In

addition, the frequencies of occurrence of Lampanyctodes hectoris in BT-180'

hauls from the 300 m and 500 m depth spectra was 66,7 per cent and 12,5 per

cent respectively, whereas the frequencies in the epibenthic hauls were 100,0 per

cent at identical bottom depth strata. Secondly, and from all three sampling

strategies, it can be seen that catch rates for Lampanyctodes hectoris fall mark-

edly from the 300-m to the 500-m bottom depth strata, and that the species was

absent from all stations in the 700-m and 900-m bottom depth strata. The

maximum depth of occurrence for the species during Cruise 060 was 552 m

(St. A 6986). Lampanyctodes hectoris therefore appears to be confined to depths

less than about 500 m in summer and is distributed mainly inshore of the 300-m

isobath.

Winter catch rates for Lampanyctodes hectoris in relation to bottom depth

during Cruises 011 and 023 have been discussed above. Pooled data for both

TABLE 5

Catch rates (kilograms.hour) for pseudoceanic and oceanic species of Myctophidae

from benthic stations (BT-180), epibenthic stations (Engels-308) and pelagic stations

(Engels-308) during R.S. Africana Cruise 060 (March 1988). SD = standard deviation.

; Bottom Pseudoceanic Oceanic

Station depth No. catch rate catch rate

type stratum hauls (kg.h’) (kg.h)

(m) Mean SD Mean SD

Pee 300 5 023. O27 000 seam

500 8 0,003 0,01 0,01 0,02

700 8 0,00 0,00 0,03 0,05

900 7 0,00 0,00 0,18 0,26

Epibenthic

300 5 66,40 40,92 0,004 0,01

500 7 0,36 0,74 8,73 a7)

700 | 0,00 0,00 6,68 6,89

900 8 0,00 0,00 B322 1,40

Pelagic

300 1 15,20 = 0,00 =

500 2 0,01 0,01 5,87 8,30

700 1 0,00 = 2,78 =

900 1 0,00 — 232,87 =

SOUTHERN BENGUELA LANTERNFISHES 425

cruises (Table 3) indicate a decrease in catch rates from 69,3 specimens.h"! over

bottom depths less than 501 m to 38,3 specimens.h'! over 500-1 000 m and a

marked decline to 1,2 specimens.h'! and 0,3 specimens.h' over bottom depth

ranges of 1 000-2 000 m and 2 000-3 000 m respectively. Although the species

was not taken over bottom depths greater than 3 000 m during these cruises,

three specimens were captured with an RMT-2 over 3 816 m (St. A 002023)

during the Phyllosoma Cruise in August 1982. Table 6 summarizes data for the

maximum depth of the water column for pelagic captures of Lampanyctodes hec-

toris during the autumn—winter period. These data therefore suggest that

Lampanyctodes hectoris is more widely distributed in autumn—winter than in

summer, being regularly taken to the 1 000-m isobath and less frequently to the

3 000-m isobath. An hypothesis to discuss this seasonality in distribution is

advanced below.

TABLE 6

Maximum depth of water column for pelagic capture of Lampanyctodes hectoris during the

1982 Phyllosoma Survey (Cruise 002), 1983 Phyllosoma Survey (Cruise 011), 1984

Phyllosoma Survey (Cruise 023) and 1983 Anchovy Acoustic Survey (Cruise 009) by SFRI.

BDepth = bottom depth; FDepth = fishing depth; No. specs = number of specimens.

Month Cruise Gear Station FDepth BDepth No.

No. (m (m specs

May 009 RMT-8 72-05 100-50 2 090 10

August 002 RMT-2 002023 75-0 3 816 3

August 011 RMT-8 P 001 100-25 1 030 12

August 023 RMT-8& 6 (4) 120-0 2 800 3

In the Mauritanian Upwelling Region and in summer, Badcock (1981) has

recognized both an on-slope and an off-slope myctophid fauna, and a third

fauna comprising the obligatory pseudoceanic species Diaphus taaningi and facul-

tative pseudoceanic species Diaphus dumerilii confined to the shelf-break zone.

He has stated (1981: 1489) that ‘it is naive to assume that water column depth

has much influence as a separating boundary’ and has suggested that shelf—slope

zonation is the dynamic response of each group to hydrographic and circulatory

characters that maintains core abundances as separate entities, in much the same

manner as been shown by Petersen et al. (1979) in zooplankton studies off the

Oregon coast. The latter authors have proposed that copepod distributions are

maintained by two-cell, zonal circulation patterns during active upwelling, and

that the seaward limit of nearshore populations is dictated by the dynamics of

the upwelling front and its associated longshore current. Larval fish assemblages

off Oregon parallel these findings (Richardson & Pearcy 1977; Richardson et al.

1980). Prograde shelf-break fronts that are characteristic of eastern boundary

current upwelling systems, form a boundary zone to the plankton-rich shelf

waters and Owen (1981) has suggested that periodic relaxation or breaching of

the front may be biologically important, since they allow for the release of nutri-

ents and plankton into off-shore waters. Frontal relaxation occurs with cessation

426 ANNALS OF THE SOUTH AFRICAN MUSEUM

of upwelling, which may be either seasonal or episodic depending on the wind

regime, whereas the causes of breaching probably involve meanders and eddies,

which may be produced by local instabilities of the frontal currents, by outside

eddy impingement, or by wave interactions with the front. Such perturbations

on the Benguela front, leading to the formation and cross-frontal motion of

mesoscale eddies, have been documented from satellite remote sensing (Lutje-

harms 1981). Johnson & Van der Spoel (1986) have postulated that the main

determinant in the distribution patterns of pelagic species is the food supply,

tempered by seasonality. Hulley & Krefft (1985) have examined the distribution

of the myctophid Electrona risso in relation to the 10 °C- and 15 °C-isotherms at

200 m, and to primary productivity values (50 gCm*y" isoline), whereas Yentsch

& Garside (1986) have produced a semi-empirical model of two phytoplankton

fronts that correspond to the northern and southern limits of myctophid popu-

lations in the biogeographic region of the subtropical North Atlantic Ocean.

It is hypothesized therefore that the seasonal differences in the seaward

extent of the distribution of Lampanyctodes hectoris in the southern Benguela

region is causally related to the seasonal dynamic of the frontal system and its

associated jet current, and is geared to the biological prerequisites of both the

summer feeding phase and winter spawning phase of the species (Hulley &

Prosch 1987). Whereas Andrews & Hutchings (1980) have suggested that the

influence of the Cape Peninsula upwelling plume expands in summer, seasonal

changes inferred from large-scale temporal and spatial averaging of sea-surface

temperature data suggest that the events are dominated by summer warming

and winter cooling of the sea-surface temperatures within the southern Benguela

region (Wooster 1973; Christensen 1980; Parrish et al. 1983; McLain et al. 1985;

Kamstra 1985; Shelton et al. 1985). On the basis of data collected over the

CELP Survey grid at monthly intervals, Shelton (1986) has stated that the most

noticeable feature of the region, and indeed of the whole Benguela System, is

the contraction of the area occupied by cool water in summer and the expansion

of this zone in winter, and that the influence of upwelling is modulated or

squeezed by off-shore warming from late spring to early winter. Shannon (1986:

10) has stated that the Benguela upwelling region is ‘pinched’ during summer

and early autumn between the warm Angola and Agulhas Current systems.

Averaged and smoothed sea-surface temperatures over a 50-year period reiter-

ate this seasonality (Kamstra 1985, fig. 6). The seasonal effect is also graphically

demonstrated in Gorshkov’s (1978) oceanographic atlas at depths of 0 m, 25 m

and 50 m; however, the presentations should be regarded with reservation, since

the atlas does not show the data distribution on which the figures are based. At

the sea surface at least, interannual changes in gross upwelling may be similar to

seasonal changes, and can be related to large-scale interannual variability in the

environment in the south-east Atlantic (L. V. Shannon, SFRI, pers. comm.). |

In summer, off-shore warming together with strong upwelling in-shore, the

penetration of Agulhas Bank water, and a steep shelf break close to the coast,

combine to cause the formation of an exceptionally strong thermal front and an

SOUTHERN BENGUELA LANTERNFISHES 427

associated equatorward, shelf-edge jet current. Both features approximately

track the 230-m isobath and truncate the seaward expansion of the in-shore,

upwelled water (Bang 1973a; Harris & Shannon 1979; Shelton & Hutchings

1982; Nelson & Polito 1987). The baroclinic jet, which is thought to persist

throughout the summer after its initiation in spring (Andrews & Hutchings 1980;

Brundrit 1981), appears to extend upwards from the shelf edge (250 m) towards

the surface, where some modification occurs under the direct action of wind

(Bang 1973a; Nelson & Polito 1987, fig. 3). Plots of the relative abundances of

Lampanyctodes hectoris in Engels—308 epibenthic and pelagic hauls from

Cruise 060 (Fig. 8A) suggest little correlation with temperature, although abun-

dance values greater than 10 kg.h' tend to be found in-shore of 394-m isobath

and at sea-surface temperatures less than 16,4 °C. According to the definitions

proposed by Andrews & Hutchings (1980) this temperature regime is suggestive

of ‘mixed water’.

Warm oceanic water close to the coast during the upwelling season also

reduces the extent of the productive area and there is good agreement between

the positions of the thermal front (15 °C-18 °C) and the chlorophyll (colour)

front (0,3 mg.m%) (Shannon et al. 1983, 1985; Shelton et al. 1985). Elevated

plankton standing stocks and microplankton concentrations tend to be restricted

in-shore of the front (Shelton 1986). During this season, Lampanyctodes hectoris

mirrors this distribution and occurs mainly in-shore of the 300-m isobath. Fur-

thermore, its biology is geared to feeding (mainly copepods (61,6% ), amphi-

pods (26,6%), euphausiids (11,6% )—Centurier-Harris (unpublished data) in

Prosch 1986) and to the production of a lipid energy reserve for later use in the

winter—spring spawning cycle (Hulley & Prosch 1987). Although ecological

investigations of the feeding of Lampanyctodes hectoris in the southern Benguela

region have not yet been undertaken, Young & Blaber (1986) have demon-

strated that the feeding intensity of the species off the Tasmanian coast is

greatest during the summer period. The data in Table 5 suggest that there is

some ‘leakage’ of the species to the 500-m isobath, but the causes for a distinct

off-shore distributional limit at this depth in summer are not apparent since

upwelling filaments (and presumably their plankton content) may extend to the

2 000-m isobath (Lutjeharms & Stockton 1987). Leakage of the species across the

300-m isobath may be linked to the generation of cool-water eddies at the front

(cf. Centropages brachiatus—De Decker 1984) or to the sinking of plankton-

rich surface water along the front (Clowes 1954; Bang 1973a; Andrews 1974;

Andrews & Hutchings 1980; Shannon ef al. 1983; Pillar 1986). Whereas the

chlorophyll colour front is well demarcated off the west coast of southern Africa

(Shannon et al. 1985), chlorophyll does not appear to be concentrated at the

front (Shannon & Field 1985, fig. 2—approximate chlorophyll concentrations

<0,1 mg.m°? west of front; 2-3 mg.m® in the frontal zone; and 3-9 mg.m°

inshore of the front) and Hutchings (1981) has found no particular association of

higher zooplankton biomass with the frontal zone. However, pronounced peaks

of mesozooplankton do occur immediately inshore of the front after periods of

428 ANNALS OF THE SOUTH AFRICAN MUSEUM

1000

750

(m)

DEPTH

500

BOTTOM

250 O absent @ <10 kg.h"!

@ 10-50 kg

@ >50 kg.h”!

12,0 14,2 16,5 18,7 21,0

SURFACE TEMPERATURE (°C)

@ <10 spec. hl

a 10-20 spec. hb!

-1

a >20 spec.h

(m)

DEPTH

BOTTOM

14,0 14,5 15,0 15,5 16,0

SURFACE TEMPERATURE (°C)

Fig. 8. Relative abundances of Lampanyctodes hectoris in relation to bottom depth and

sea-surface temperature: (A) epibenthic + midwater Engels—308 stations from Cruise 060;

(B) RMT-8 stations from Cruise 023. MWT = midwater Engels—308 haul.

SOUTHERN BENGUELA LANTERNFISHES 429

prolonged upwelling (Hutchings et al. 1986) and would allow shelf-edge popu-

lations of oceanic pelagic fish species (particularly myctophids) to reap the

benefit of the upwelling productivity, as has been suggested for the Mauritanian

Upwelling region (Badcock 1981). In the latter region, the zooplankton biomass

(principally copepods) is higher over the outer shelf and slope than the inner

shelf (Blackburn 1979), whereas in the southern Benguela region copepod

biomass is highest over the shelf (Pillar 1986) and is associated both with

enhanced primary productivity and pelagic fish production on the shelf

(Shannon & Field 1985). Although demersal fishes, which are indicative of less-

productive areas (Mills & Fournier 1979), are said to dominate the slope regions

(Shannon & Field 1985), high catch-rates for shelf-edge myctophid species

(mainly Diaphus hudsoni, Symbolophorus boops and Symbolophorus barnardi)

during Cruise 060 in the southern Benguela (Table 5) approximate similar

values for equivalent shelf-edge species (mainly Diaphus dumerilii and Diaphus

taaningi) in the Mauritanian Upwelling region (Gjdésaeter & Blindheim 1982,

table 77).

During winter the frequency of active upwelling is reduced in the southern

Benguela region (Lutjeharms & Meeuwis 1987). The front becomes weaker and

relaxes, with a consequent decline in sea-surface temperature values. The

seaward extent of cooler waters (less than 16 °C) increases up to 220 km off-

shore in association with a seaward expansion of the upwelling filaments (Parrish

et al. 1983; Shelton et al. 1985; Lutjeharms & Stockton 1987). This results in the

formation of a wider productive zone than in summer but one in which near-

surface chlorophyll concentrations are lower (Shannon et al. 1985). The standing

stock of plankton may increase in winter but the production may be less because

of low light levels and reduced nutrient regeneration (Shelton et al. 1985). Evi-

dence from thermal infra-red satellite imagery during August 1984 (Fig. 3B)

indicates that those stations, which were clustered as Group II in the above

analysis (stations 4 (1), 5 (1), 7 (1), 8 (1), 9 (1)), are all situated seaward of the

surface expression of the fronts. Absolute sea-surface temperatures for these

stations ranged between 14,9 °C-—15,5 °C and are similar to the 14,8 °C-—

15,7 °C range for those stations clustered as Group I. By contrast, montage of

the station positions in relation to the location of the fronts during August 1983

(Fig. 3A) reveals that all stations were inshore of the front at this time. Absol-

ute sea-surface temperatures for these stations were not available for this cruise.

These findings therefore suggest that during winter the off-shore distribution of

Lampanyctodes hectoris is unrelated to the position of the surface front.

In winter, feeding intensity may be reduced (Young & Blaber 1986) and the

lipid composition of adult Lampanyctodes hectoris decreases in association with

reproductive activity (Hulley & Prosch 1987). CELP Survey data indicated that

spawning did not occur to any great extent in-shore of the 14 °C surface iso-

therm and that egg abundances were greatest along the shelf break between the

300-m and 1 000-m isobaths (Shelton 1986, fig. 5.28). Off South West Africa—

Namibia, where the seasonality of upwelling is less marked than in the southern

430 ANNALS OF THE SOUTH AFRICAN MUSEUM

Benguela region (Shannon 1985; Lutjeharms & Meeuwis 1987), spawning also

takes place in winter—early spring (August-November) (Ahlstrom et al. 1976).

The latter authors have reported that more than 60 per cent of the larvae

occurred where sea-surface temperatures ranged between 14,0 °C and 15,5 °C,

and larval abundances were greatest seaward of the 100 fm (183 m) isobath.

Since both data sets imply the presence of large populations of spawning adults

beyond the shelf break during the winter season, it seems not unlikely that there

is a shift in the distribution of the species to deeper waters at this time. The

analyses of our data given above would confirm this supposition. Plots of the

relative abundances of Lampanyctodes hectoris at RMT-8 stations from Cruise

023 (winter) in relation to bottom depth and sea-surface temperature (Fig. 8B)

indicate that, whereas the species occurred over bottom depths to about 3 000 m,

they were most abundant at depths shallower than 760 m, where sea-surface

temperatures ranged between 14,9 °C and 15,5 °C. The temperature range

matches that proposed for the larval distribution patterns (Ahlstrom et al. 1976;

Shelton 1986).

In summary, it is postulated that the summer distribution of Lampanyctodes

hectoris mainly in-shore of the 300-m isobath is related to the availability of

suitable quantities of food in the water column. During this time a lipid energy

reserve is metabolized for use in later spawning activity. The distribution of

these food items in the southern Benguela region is itself governed by the

dynamics of the frontal system which develops during upwelling and which

restricts plankton-rich waters to the east of the shelf break. Sinking at or breach-

TABLE 7

Minimum depth of water column for pelagic and benthic captures of oceanic Myctophidae during the

Phyllosoma, Anchovy Acoustic and Hake surveys by SFRI. BDepth = bottom depth; FDepth = fishing

depth, No. specs = number of specimens.

Zone Month Cruise Gear Station FDepth BDepth No. Species

No. (m) (m Specs

Pelagic May 009 RMT-8 28-07 100-0 510 147 Symbolophorus boops

4 Symbolophorus barnardi

August 002 RMT-2 002035 100-0 540 1 Diaphus hudsoni

August 011 RMT-8 P007 250-0 290 1 Diaphus diadematus

1 Diaphus hudsoni

1 Lampanyctus alatus

August 023 RMIT-8 1(1) 120-20 525 1 Lampanyctus australis

Benthic January 028 180’°BT A2304 354 354 1 Diaphus ostenfeldi

January 039 180’°BT A345 740 740 (22 Symbolophorus boops

January 050 180°BT A5276 407 407 1 Lampadena notialis

July 022 180’°BT A1550 378 378 2 Symbolophorus boops

July 033 180°BT A2780 496 496 1 Diaphus hudsoni

1 Symbolophorus boops

July 046 180°BT A4309 298 298 2 Symbolophorus barnardi

July 054 180°BT AS5893 358 358 3 Symbolophorus boops

SOUTHERN BENGUELA LANTERNFISHES 431

ing of the front may occur and may account for the apparent ‘leakage’ of the

species to depths of 500 m. In winter the off-shore distribution expands and Lam-

panyctodes hectoris may be taken regularly over depths of 800 m, and less fre-

quently over depths between 800 m and about 3 000 m. Although relaxation of

the front takes place during this season, it seems unlikely that the distribution of

the species is geared to the wider productive zone, since the decrease in lipid

content would suggest a lower feeding intensity than in summer, and distribution

of the species does not appear to be correlated with the positions of the winter

fronts and the extent of nutrient-rich waters in the surface layers. Rather, it

appears that adult populations of Lampanyctodes hectoris move to deeper water

to spawn and catabolize lipids for the production of eggs.

Data at hand indicate that similar seasonal differences in distributional pat-

terning are not evident for oceanic myctophids and suggest that water-column

depth and, therefore, the potential to undertake diurnal migration may be the

major factor governing the shoreward distribution of such species. During the

summer months, the shallowest water-column depth (284 m) for pelagic cap-

tures of oceanic species (Table 5) is matched by an equivalent water-column

depth (290 m) during winter (Table 7), whereas their occurrence in benthic

hauls varies between 350-500 m in summer and 300-500m in winter

(Tables 5, 7).

ACKNOWLEDGEMENTS

We would like to express our sincere thanks to Dr D. E. Pollock and the

Director of the Sea Fisheries Research Institute, Cape Town, for access to the

lanternfish material and to the Master, Officers and Crew of the R.S. Africana

who were involved in the collection of the specimens. Our thanks are also due

to Professor John Field and Ms Coleen Maloney of the Institute for Marine

Research, University of Cape Town, for setting up the data for the computer

program and for ensuing discussion of the results, and to Dr Vere Shannon of

the Sea Fisheries Research Institute, Cape Town, for his comments. The satel-

lite images were prepared by the staff of the Satellite Remote Sensing Centre of

the CSIR at Hartebeeshoek. This study was made possible through grants from

the CSIR Foundation for Research Development.

REFERENCES

AHLSTROM, E. H., Moser, H. G. & O’TooLe, M. J. 1976. Development and distribution of

larvae and early juveniles of the commercial lanternfish Lampanyctodes hectoris (Giinther)

off the west coast of southern Africa, with a discussion of phylogenetic relationships of the

genus. Bulletin of the Southern California Academy of Sciences 75 (2): 138-152.

ANDREWS, W. R. H. 1974. Selected aspects of upwelling research in the southern Benguela

current. Téthys 6 (1-2): 327-340.

ANDREWS, W. R. H. & Hurcuincs, L. 1980. Upwelling in the southern Benguela Current.

Progress in Oceanography 9 (1): 1-81.

432 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bapcock, J. 1981. The significance of meristic variation in Benthosema glaciale (Pisces,

Myctophoidei) and of the species distribution off northwest Africa. Deep-Sea

Research 28 (12A): 1477-1491.

BAKER, A. DE C., CLARKE, M. R. & Harris, M. J. 1973. The N.I.O. combination net (RMT

1+ 8) and further developments of rectangular midwater trawls. Journal of the Marine

Biological Association of the United Kingdom 53 (1): 167-184.

BANG, N. D. 1971. The southern Benguela Current region in February, 1966: Part II. Bathy-

thermography and air-sea interactions. Deep-Sea Research 18 (2): 209-224.

BANG, N. D. 1973a. The southern Benguela system: finer oceanic structure and atmospheric

determinants. Ph.D. thesis: University of Cape Town.

Banc, N. D. 1973b. Characteristics of an intense ocean frontal system in the upwell regime

west of Cape Town. Tellus 25 (3): 256-265.

Banc, N. D. & ANDREws, W. R. H. 1974. Direct current measurements of a shelf-edge

frontal jet in the southern Benguela system. Journal of Marine Research 32 (3): 405-417.

BEKKER, V. E. 1983. Myctophids of the world ocean. Moskva: Akademiya nauk SSSR. (In

Russian).

BLAcKBURN, M. 1979. Zooplankton in an upwelling area off northwest Africa: composition,

distribution and ecology. Deep-Sea Research 26 (1A): 41-56.

Boyp, A. L. & AGENBAG, J. J. 1985. Seasonal trends in the longshore distribution of surface

temperatures off Southwestern Africa 18—34°S, and their relation to subsurface conditions

and currents in the area 21—24°S. Simposio international sobre las areas de afloramiento

mas importantes del Oueste Africano (Cabo Blanco y Benguela), Instituto de Investigaciones

Pesqueras, Barcelona 1: 119-148.

Brunoprit, G. B. 1981. Upwelling fronts in the southern Benguela region. Transactions of the

Royal Society of South Africa 44 (3): 309-313.

CHRISTENSEN, M. S. 1980. Sea-surface temperature charts for southern Africa south of 26°S.

South African Journal of Science 76 (12): 541-546.

Crowes, A. J. 1954. An introduction to the hydrology of South African waters. Investi-

gational Reports. Fisheries and Marine Biological Survey Division, Union of South

Africa 12: 1-42.

CRAWFORD, R. J. M., SHANNON, L. V. & PoLtock, D. E. 1987. The Benguela ecosystem.

Part IV. The major fish and invertebrate resources. Oceanography and Marine Biology.

An Annual Review 25: 353-505. Aberdeen: University Press.

DINGLE, R. V., BircH, G. F., BREMNER, J. M., DE DECKER, R. H., Du PLeEssis, A., ENGEL-

BRECHT, J. C., FINCHAM, M. J., Fitton, T., FLEMMING, B. W., GENTLE, R. I., GOODLAD,

S. W., Martin, A. K., Mitis, E. G., Morr, G. J., PARKER, R. J., RoBSON, S. H., RoGErs, J.,

SALMON, D. A., SIESSER, W. G., Simpson, E. S. W., SUMMERHAYES, C. P., WESTALL, F.,

WINTER, A. & WoopsornE, M. W. 1987. Deep-sea sedimentary environments around

southern Africa (South-east Atlantic and South-west Indian oceans). Annals of the South

African Museum 98 (1): 1-27.

FIELD, J. G., CLARKE, K. R. & Warwick, R. M. 1982. A practical strategy for analysing

multispecies distribution patterns. Marine Ecology Progress Series 8 (1): 37-52.

GIJOSAETER, J. & BLINDHEIM, J. 1982. Observations on mesopelagic fish off Northwest Africa

between 16° and 27°N. Rapport et procés-verbaux des réunions. Conseil permanent inter-

national pour l’exploration de la mer 180: 391-398.

GorsHkov, S. G. (Ed.) 1978. World ocean atlas. 2. Atlantic and Indian oceans. Oxford:

Pergamon.

Harris, T. F. W. & SHANNON, L. V. 1979. Satellite-tracked drifter in the Benguela Current

system. South African Journal of Science 75 (7): 316-317.

Hopkins, T. L., MILLIKEN, D. M., BELL, L. M., MCMIcHAEL, E. J. HEFFERNAN, J. J. & CANO,

R. V. 1981. The landward distribution of oceanic plankton and micronekton over the

west Florida continental shelf as related to their vertical distribution. Journal of Plankton

Research 3 (4): 645-658.

Huttiey, P. A. 1981. Results of the research cruises of FRV ‘Walther Herwig’ to South

America. LVIII. Family Myctophidae (Osteichthyes, Myctophiformes). Archiv fir Fis-

chereiwissenschaft 31 (1): 1-300.

Hu tey, P. A. 1986a. Family No. 86: Myctophidae. Jn: SmitH, M. M. & HeEeEmstra, P. C.

eds. Smiths’ sea fishes: 282-321. Johannesburg: Macmillan.

SOUTHERN BENGUELA LANTERNFISHES 433

Huttey, P. A. 1986b. Lanternfishes of the southern Benguela region. Part 1. Fauna! complex-

ity and distribution. Annals of the South African Museum 97 (7): 227-249.

Hu iey, P. A. & Krerrr, G. 1985. A zoogeographic analysis of the fishes of the family

Myctophidae (Osteichthyes, Myctophiformes) from the 1979-Sargasso Sea Expedition of

R.V. Anton Dohrn. Annals of the South African Museum 96 (2); 19-53.

Hu Ley, P. A. & Proscu, R. M. 1987. Mesopelagic fish derivatives in the southern Benguela

Upwelling Region. Jn: Payne, A. I. L., GULLAND, J. A. & BRINK, K. H. eds. The Ben-

guela and comparable ecosystems. South African Journal of Marine Science 5: 597-611.

HutcuHincs, L. 1981. The formation of plankton patches in the southern Benguela Current.

In: RicHarps, F. A. ed. Coastal and Estuarine Sciences 1. Coastal Upwelling: 496-506.

Washington: American Geophysical Union.

HutTcuincs, L., ARMSTRONG, D. A. & MITCHELL-INNES, B. A. 1986. The frontal zone in the

southern Benguela Current. Jn: NtHouL, J. C. J. ed. Marine Interfaces and Ecohydro-

dynamics: 67-94. Amsterdam: Elsevier.

JOHNSON, R. K. & VAN DER SPOEL, S. 1986. Summary report and recommendations. In:

PrERROT-BULTs, A. C., VAN DER SPOEL, S., ZAHURANEC, B. J. & JOHNSON, R. K. eds.

Pelagic biogeography. UNESCO Technical Papers in Marine Science 49: 201-209.

KamstrA, F. 1985. Environmental features of the southern Benguela with special reference to

the wind stress. Jn: SHANNON, L. V. ed. South African Ocean Colour and Upwelling

Experiment: 13-27. Cape Town: Sea Fisheries Research Institute.

Kawacucul, K. & SHimizu, H. 1978. Taxonomy and distribution of the lanternfishes, genus

Diaphus (Pisces, Myctophidae) in the western Pacific, eastern Indian oceans and the

Southeast Asian Seas. Bulletin of the Ocean Research Institute, University of Tokyo 10:

1-145.

KrerfT, G. 1970. Zur Systematik und Verbreitung der Gattung Lampadena Goode & Bean,

1896 (Osteichthyes, Myctophoidei, Myctophidae) im Atlantischen Ozean, mit Beschrei-

bung einer neuen Art. Berichte der Deutschen Wissenschaftlichen Kommission fiir

Meeresforschung 21 (1-4): 271-289.

LUTJEHARMS, J. R. E. 1981. Satellite studies of the South Atlantic upwelling system. In:

Gower, J. F. R. ed. Oceanography from space. Marine Science: 195-199. New York:

Plenum.

LUTJEHARMS, J. R. E. & MEEuwIS, J. M. 1987. The extent and variability of South-east Atlan-

tic upwelling. Jn: Payne, A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela

and comparable ecosystems. South African Journal of Marine Science 5: 51-62.

LUTJEHARMS, J. R. E. & Stockton, P. L. 1987. Kinematics of the upwelling front off southern

Africa. In: Payne, A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and

comparable ecosystems. South African Journal of Marine Science 5: 35-49.

MarsHALL, N. B. & Merrett, N. R. 1977. The existence of a benthopelagic fauna in the

deep-sea. In: ANGEL, M. V. ed. A voyage of discovery. George Deacon 70th Anniversary

volume. Supplement. Deep-Sea Research 24 (A): 483-497.

McLain, D. R., BRAINARD, R. E. & Norton, J. B. 1985. Anomalous warm events in eastern

boundary current systems. Report. California Cooperative Oceanic Fisheries Investiga-

tions 26: 51-64.

MerreIT, N. R. 1986. Biogeography and the oceanic rim: a poorly known zone of ichthyo-

faunal interaction. Jn: PreERRoT-BuULTs, A. C., VAN DER SPOEL, S., ZAHURANEC, B. J. &

JoHNSON, R. K. eds. Pelagic biogeography. UNESCO Technical Papers in Marine

Science 49: 201-209.

Mitts, E. L. & Fournier, R. O. 1979. Fish production and the marine ecosystems of the

Scotian Shelf, eastern Canada. Marine Biology 54 (1): 101-108.

NAFPAKTITIS, B. G. 1978. Systematics and distribution of lanternfishes of the genera

Lobianchia and Diaphus (Myctophidae) in the Indian Ocean. Science Bulletin. Natural

History Museum of Los Angeles County 30: 1-92.

NaFPAKTITIS, B. G. & PAXTON, J. R. 1968. Review of the lanternfish genus Lampadena with a

description of a new species. Contributions in Science. Los Angeles County Museum 138:

1-29.

NELson, G. & Potito, A. 1987. Information on currents in the Cape Peninsula area, South

Africa. In: PayNgE, A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and

comparable ecosystems. South African Journal of Marine Science 5: 287-304.

434 ANNALS OF THE SOUTH AFRICAN MUSEUM

O’TooLe, M. J. 1976. Incidental collections of small and juvenile fishes from egg and larval

surveys off South West Africa. Fisheries Bulletin. Division of Sea Fisheries, Republic of

South Africa 8: 23-33.

OweEN, R. W. 1981. Fronts and eddies in the sea: mechanisms, interactions and biological

effects. Jn: LonGHursT, A. R. ed. Analysis of marine ecosystems: 197-233. London:

Academic Press.

Parin, N. V. 1986. Distribution of mesobenthopelagic fishes in slope waters and around sub-

marine rises. Jn: PreRRoT-BULTS, A. C., VAN DER SPOEL, S., ZAHURANEC, B. J. &

JouHnson, R. K. eds. Pelagic biogeography. UNESCO Technical Papers in Marine

Science 49: 226-229.

Parin, N. V. & GoLovan, G. A. 1976. Pelagic deep-sea fishes of the families characteristic of

the open ocean collected over the continental slope off West Africa. Trudy Instituta

okeanologii. Akademiya nauk SSSR 104: 250-276. (In Russian).

ParRRISH, R. H., BAkuN, D. M., Hussy, D. M. & Netson, C. S. 1983. Comparative clima-

tology of selected environmental processes in relation to eastern boundary current pelagic

fish reproduction. Jn: SHARP, G. D. & CsiRKE, J. eds. Proceedings of the expert consul-

tation to examine changes in abundance and species composition of neritic fish resources.

San Jose, Costa Rica, 18-29 April 1983. FAO Fisheries Report 291: 731-777.

Pearcy, W. G. 1964. Some distributional features of mesopelagic fishes off Oregon. Journal

of Marine Science 22 (1): 83-102.

PETERSEN, W. T., MILLER, C. B. & HuTCHINSON, A. 1979. Zonation and maintenance of

copepod populations in the Oregon upwelling zone. Deep-Sea Research 26 (SA): 467-494.

PoLtockx, D. E. 1984. Report on the third phyllosoma larval survey. Internal Report. Sea

Fisheries Research Institute, Cape Town: 1-6 (mimeograph).

PittarR, S. C. 1986. Temporal and spatial variations in copepod and euphausiid biomass off

the southern and south-western coasts of South Africa in 1977/78. South African Journal

of Marine Science 4: 219-229.

ProscH, R. M. 1986. The biology, distribution and ecology of Lampanyctodes hectoris and

Maurolicus muelleri along the South African coast. M.Sc. thesis: University of Cape

Town.

RAtz, H.-J. 1984. Qualitative und quantitative Untersuchungen der Ichthozonose in der

archibenthischen Zone des Rockall-Grabens und umliegender Banke (Westbritische

Gewasser). Mitteilungen aus dem Institut fiir Seefischerei 34: 1-152.

RICHARDSON, S. L., LAROCHE, J. L. & RicHArDson, M. D. 1980. Larval fish assemblages and

associations in the north-east Pacific Ocean along the Oregon coast, winter—spring

1972-1975. Estuarine and Coastal Marine Science 11 (6): 671-699.

RICHARDSON, S. L. & PEARcY, W. G. 1977. Coastal and oceanic fish larvae in an area of

upwelling off Yaquina Bay, Oregon. Fishery Bulletin of the National Oceanic and

Atmospheric Administration 75 (1): 125-145.

RoeEL, B. A. 1987. Demersal communities off the west coast of South Africa. Jn: PAYNE,

A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and comparable ecosystems.

South African Journal of Marine Science 5: 575-584.

Rusiks, P. 1985. Zoogeography of the lanternfishes (Osteichthyes, Myctophidae) of South-

west Africa. Simposio internacional sobre las dreas de afloramiento mas importantes

del Oeste Africano (Cabo Blanco y Benguela), Instituto de Investigaciones Pesqueras,

Barcelona 1: 573-586.

SHANNON, L. V. 1985. The Benguela Ecosystem. Part I. Evolution of the Benguela, physical

features and processes. Oceanography and Marine Biology. An Annual Review 23:

105-182. Aberdeen: University Press.

*SHANNON, L. V. 1986. Synthesis of information on the tunas of the Benguela Region off

southern Africa. Internal Report of the Sea Fisheries Research Institute, Department of

Environmental Affairs, Cape Town 89: 1-18.

SHANNON, L. V. & AGENBAG, J. J. 1987. Notes on the recent warming in the Southeast Atlan-

tic, and possible implications for the fisheries of the region. Collection of Scientific Papers

of the International Commission for Southeast Atlantic Fisheries (ICSEAF) 1987: 243-248.

SHANNON, L. V., Boyp, A. J., BRUNDRIT, G. B. & TAUNTON-CLARKE, J. 1986. On the exist-

ence of an El Nifio-type phenomenon in the Benguela System. Journal of Marine

Research 44 (3): 495-520.

SOUTHERN BENGUELA LANTERNFISHES 435

SHANNON, L. V. & FieELD, J. G. 1985. Are fish stocks food-limited in the southern Benguela

pelagic ecosystem? Marine Ecology Progress Series 22 (1): 7-19.

SHANNON, L. V., MostTert, S. A., WALTERS, N. M. & ANDERSON, F. P. 1983. Chlorophyll

concentrations in the southern Benguela Current region as determined by satellite

(Nimbus-—7 coastal zone colour scanner). Journal of Plankton Research 5 (4): 565-583.

SHANNON, L. V. & PILLAR, S. 1986. The Benguela ecosystem. Part III. Plankton. Oceanogra-

phy and Marine Biology. An Annual Review 24: 65-170. Aberdeen: University Press.

SHANNON, L. V., WALTERS, N. M. & MostTeErT, S. A. 1985. Satellite observations of surface

temperature and near-surface chlorophyll in the southern Benguela region. Jn: SHANNON,

L. V. ed. South African Ocean Colour and Upwelling Experiment: 183-210. Cape Town:

Sea Fisheries Research Institute.

SHELTON, P. A. 1986. Fish spawning strategies in the variable southern Benguela Current

region. Ph.D. thesis: University of Cape Town.

SHELTON, P. A., Boyp, A. J. & ArmsTRONG, M. J. 1985. The influence of large-scale environ-

mental processes on neritic fish populations in the Benguela Current system. Report.

California Cooperative Oceanic Fishery Investigations 26: 72-92.

SHELTON, P. A. & Hurcuines, L. 1982. Transport of anchovy Engraulis capensis Gilchrist

eggs and early larvae by a frontal jet current. Journal du Conseil permanent international

pour l’exploration de la mer 40 (2): 185-198.

VaN ForeEEsT, D., SHILLINGTON, F. A. & LEGECKIS, R. 1984. Large-scale, stationary, frontal

features in the Benguela Current system. Continental Shelf Research 3 (4): 465-474.

Wooster, W.S. 1973. Upwelling in the eastern Atlantic. Abstracts of the South African Nat-

ional Oceanographic Symposium, Cape Town, 6-10 August 1973.

YENTSCH, C. S. & GarRsIDE, J. C. 1986. Patterns of phytoplankton abundance and biogeogra-

phy. Jn: PrerRot-Bu.ts, A. C., VAN DER SPOEL, S., ZAHURANEC, B. J. & JOHNSON, R. K.

eds. Pelagic biogeography. UNESCO Technical Papers in Marine Science 49: 278-284.

YounG, J. W. & BLaBEr, S. J. M. 1986. Feeding ecology of three species of midwater fishes

associated with the continental slope of eastern Tasmania, Australia. Marine

Biology 93 (1): 147-156.

* quoted with permission of the author.

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.£. . 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 heen 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.

P. ALEXANDER HULLEY

J. R. E. LUTJEHARMS

LANTERNFISHES OF THE SOUTHERN

BENGUELA REGION. PART 3.

THE PSEUDOCEANIC-—OCEANIC INTERFACE.

UME 98 PAR

etl

- NOVEMBER 1989

ISSN 030

3-2515

MCZ

LIBRARY

yan 23 1990

UNIV ieesitY

| ANNALS

oF THE SOUTH AFRICAN

MUSEUM

APE 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

Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

bered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

into consideration.

All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 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.:

‘Smith (1969) describes... .’

‘Smith (1969: 36, fig. 16) describes. . .’

‘As described (Smith 1969a, 1969b; Jones 1971)’

‘As described (Haughton & Broom 1927)...”

‘As described (Haughton et al. 1927)...’

Note: no comma separating name and year

pagination indicated by colon, not p.

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, 1969b) 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 (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number in parentheses, pagination (first and

last pages of article).

Examples (note capitalization and punctuation)

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

FiscHER, P. H: 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHer, P. H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-S1.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHULTzE, L. Zoologische und anthro-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 ~~ Band

November 1989 November

Part 11 #£Deel

FIRST RECORD OF AN EXTANT, SIGHTED,

SHALLOW-WATER SPECIES

OF THE GENUS POSEIDONAMICUS BENSON

(OSTRACODA) FROM THE CONTINENTAL

MARGIN OF SOUTH-WESTERN AFRICA

R. C. WHATLEY

R. V. DINGLE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

1, 2(1=3, 528) 3012: 4252.8. Hapa ye SlEs, See

6(1, t.-p.i.), 71-4), 8, 9(1-2, 7), 10(1-3),

11(1-2, 5, 7, t.-p.i.), 14(1-2), 15(4—5), 24(2), 27, 31(1—-3), 32(6), 33, 36Q)eaoum

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 105 9

Printed in South Africa by In Suid-Afrika gedruk deur

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

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

FIRST RECORD OF AN EXTANT, SIGHTED, SHALLOW-WATER

SPECIES OF THE GENUS POSEIDONAMICUS BENSON

(OSTRACODA) FROM THE CONTINENTAL MARGIN OF

SOUTH-WESTERN AFRICA

R. C. WHATLEY

Department of Geology, University College, Aberystwyth, UK

R. V. DINGLE

South African Museum, Cape Town

(With 11 figures and 3 tables)

[MS accepted 23 August 1988]

ABSTRACT

A new, sighted species of the ostracod genus Poseidonamicus (P. panopsus sp. nov.) is

recorded living on the continental shelf and upper slope off south-western Africa in water depths

between 120 m and 545 m. All previous records of the genus have been from water depths

greater than 1 km. The discovery indicates that sighted species survived, in shallow-water, from

the late Cretaceous/early Tertiary stocks when the genus originated in the SW Pacific area. The

habitat of the new species is defined in terms of temperature, salinity, and dissolved oxygen

values within the Antarctic Intermediate Water mass, where it forms part of the Benguela

system, and associated cells of intense upwelling.

CONTENTS

PAGE

i peer og ee kc Sh ans wi Sle aus ale «PSA ome wwe 437

i Se caw ei Manca ed ae ee wo GERMS ee Oe dad 438

Se en I OS Oe et ce fe sn oa Sb we ees ee Che whan oes 447

ee a N@h AMCTECOIOSY | 2 sae edge ees eee ee nee bees 448

MMe NIRCONSICIETAUIONS: 6-6 5 ccs cc se es nao e naan de ces ee vie views 454

aa ete TE re Fo han Se og «ais ba wld v wwe Sa a a ere Bele male Bw 456

INTRODUCTION

Cenozoic bathyal and abyssal deep-water ostracod faunas are characterized

by distinctive cosmopolitan assemblages that include a number of ornate

cytheracean taxa. Prominent amongst this group is the genus Poseidonamicus,

which was originally erected by Benson (1972) to accommodate five species

with a geological range ?Oligocene, Miocene—Recent, and which is extant

437

Ann. S. Afr. Mus. 98 (11), 1989: 437-457, 11 figs, 3 tables.

438 ANNALS OF THE SOUTH AFRICAN MUSEUM

world-wide (Table 1). All the original species considered by Benson (1972), as

well as two further Miocene species from the South Atlantic (Benson in Benson

& Peypouquet 1983), were blind. Subsequent studies by Whatley and his

colleagues (e.g. Whatley 1983, 1985; Whatley et al. 1983; Whatley et al. 1986)

have extended the geological and geographical range of the genus (now known

from Eocene to Recent). Despite the fact that Whatley et al. (1986) described one

species with distinct but non-functional ocular structures (P. ocularis Whatley et

al. 1986), all records of the genus to date have been from water depths known or

estimated to be in excess of 1 km. Poseidonamicus has, therefore, up to now been

considered typically a deep-water taxon, although the discovery of P. ocularis in

Quaternary deep-water sediments prompted Whatley et al. (1986) to predict that

a shallow-water ancestor existed at least until late Tertiary times in the SW Pacific

region.

Table 1 lists the known species of the genus Poseidonamicus, as well as their

geological and geographical ranges.

The most widely recorded species to date has been the type species P. major

Benson, which has a known geological range Miocene to Recent (southern

hemisphere), and a possible record in Oligocene of the NE Atlantic (Table 1).

This species occurs in Pleistocene—Recent sediments off south-western Africa,

where it has a known depth range 2 070-2 916 m (Dingle et al. in press).

Although the genus has been recorded from Tertiary sediments in the South

Atlantic (P. riograndensis and P. miocenica from the Miocene of Deep Sea

Drilling Project (DSDP) site 92—Benson & Peypouquet 1983), the main centre

of evolutionary activity and the area where the genus has the longest geological

record (Eocene to Recent) is the SW Pacific (e.g. Whatley 1985). No positive

identifications of the genus have so far been made in the Tertiary strata of

southern Africa (Frewin’s 1987 record is considered questionable), so that the

discovery of an extant population of a shallow-water species off southern Africa

poses some interesting evolutionary, zoogeographical, and environmental

questions.

A total of 119 valves of the new species (P. panopsus) have been recovered

in 27 sediment samples from the continental shelf and slope off south-western

Africa (Fig. 1, Table 2). All the samples were collected with a Van Veen grab by

the Marine Geoscience Unit of the University of Cape Town from the research

vessel Thomas B. Davie. (Sample numbers have a TBD prefix.)

TAXONOMY

All illustrated material is housed in the South African Museum under

catalogue numbers prefixed SAM—PQ-MF. Abbreviations: ACA = anterior

cardinal angle, AM = anterior margin, ATE = anterior terminal element, DM =

dorsal margin, LV = left valve, MS = muscle scars, PM = posterior margin,

PTE = posterior terminal element, RV = right valve, SCT = subcentral tubercle,

VM = ventral margin.

439

(6861) AepeyM 2 S2joD onuepy AN dUd90311O 6861 ‘AONeYM 2 SOD ‘Aou ‘ds snoi1uYUOplasog

(L861) S210D 29 Aopeuym onuepy N d10Z0U92) 94e] L86L ‘S210D 29 Aopey ‘ds snonuvuopiasog

(L861) S910D 2 Aopey onuepy N d10ZOUDZ) 91k] ZL6L ‘UOSUdg 10JuId ‘g ‘JO SNoIUDUOplasog

(L861) S210D 2 AONeuA onueNy N d10ZOUID 91P] ZL61 ‘UOsUdg solv ‘g ‘JD SNIIUDUOpIasog

dled MS

eR (9861) 70 12 AapeyAA 4oAng rey rey eI] dUd90I|d 9861 ‘70 Ja Aopeyy ‘ds snonuvuopiasog

= (9961) 70 Ja AopeuM dIde@q MS 2U90031[9--9u900q O86I ‘70 Ja AapeuAA snisnqos snonuvuopiasog

(9861) 70 Ja Aopeum olde@q AAS Areusajend 9861 ‘70 Ja AapeuAy Snjvjound snonuouopiasog

= (L861) S910D 2 Aopey AA onuepy N dIOZOUID 93k]

2 (9961) ‘70 Ja Aopeun aioeg MS Areusayen?--dus001|g 9861 ‘70 Ja AapeyA, Snpnuavid snonuvuopiasog

(9961) 70 Ja AopeuNM o1ise@q AAS Areusajend O86I ‘70 Ja AapeyAy SUvjNIO SnoiWDYUOpIasog

° (9961) 70 7a Aopeum dled MS AreussyenG--sus001]_ 9Q6I ‘70 Ja AapeyAy Snjvjoundouajuv snomupuopiasog

a (€g61) yonbnodhag 2 uosusg onueny § ‘76 Os dasa UdDOI] 2IPPIA E861 ‘UOSUdg voIUAIOIU SNIWUMDUOpIasOg

i (€861) onbnodAag 2 uosusg onuepy S ‘7% 21s dasa QUdDOIJY JOMOT €86[ ‘UOsUdg SisuapuDisoU SndIWMDUOpIasog

a (ZL61) uosusg ‘(QggT) Aperg uedd0O UIOYINOS juD09y (Ogst ‘Apesrg) vawiuia snoiwouopiasog

: (ZL61) uosusg suey Ss JU999y] ZL6I ‘UOSUdg 10jJuId snoIUDUOpIasog

o (ZL61) uosusg oyloeg AS ‘ueIpuy juD99y ZL61 ‘UOSUdg SnpnU snoIUDUOplasog

Z (ZL61) uosusg oloeg 7 AreurayenO ZL6I ‘UOSUdg 4OUNU SnoIWUDUOpIasog

2 (ssoid ut) 1oWI00g 2» p1OT ‘oj3uIG Jesshqe onueny AS AreurayenO

Zz ZL6[) uosusg yesshqe uvIpUuyT AAS jUD00y

< ZL6[) uosusg onuepy MS = yUd00yZ--oUDD0I[J

ZL6[) uosusg yjessXqe--jeAyyeq oloeg MS JU999Y--dUDIOIJY

ZL6[) uosusg onueny AN dUdD0TIIOZ ZL61 ‘UOsudg sofpi snoiuDUOplasog

aouadafay yjdaq M0207 asp saidad¢

“sndiubUOpiasod JO saideds poqiiosop AjsnorAosd jo uOINGIIsIp [erodur9) pue jeneds

Tl dIaVL

440 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Sample sites.

TBD Latitude Longitude Depth No. ostracod No. Percentage — Percentage

S or (m) valves _—_ P. panopsus P. panopsus mud

Walvis Shelf

3769 22,250 13,233 223 39 1 3 93

Orange Shelf

2752 29,783 16,017 170 374 1 0,1 90,4

2973 28,417 15,000 173 1 460 1 0,1 82,4

2975 28,417 14,583 180 892 1 0,1 76,4

2925 28,900 15,367 183 284 3 1 25,8

2974 28,417 14,800 186 2011 2 0,1 7

2840 30,917 15,700 205 108 9 8 8,7

2736 29,950 15,417 205 555 6 t 83

Ziki 30,400 16,433 218 112 4 4 33

2703 30,133 15,433 220 239 7 a 76,4

2485 30,917 16,000 227 114 2 2 19,6

2719 30,450 16,067 240 140 13 10 302

2361 31,313 16,858 241 70 9 13 64

2884 Z9,S17 14,717 252 159 2 1 js

2691 30,533 15,617 265 43 1 2 27,4

2690 30,517 15,833 ZyA 137 f 9 23

2460 31,233 16,625 Ze 39 1 3 90

2459 a1 253 16,383 300 29 5 2 85,7

2976 28,417 14,383 350 18 2 11 92,5

2447 31,925 16,467 350 49 1 2 22,9

3577 31,367 16,083 453 96 1 1 84,2

SW Cape

6823 34,093 18,215 120 1 098 11 1 =

3587 33,995 18,148 140 167 q ~ ZOyt

6825 34,090 18,173 160 Zee 10 5 ==

1691 34,567 18,483 220 516 5 1 35

1694 34,750 18,327 425 ol 1 3 72,4

1697 34,767 18,250 545 112 it 1 62

Phylum CRUSTACEA Pennant, 1777

Class OSTRACODA Latrielle, 1806

Order PODOCOPIDA Miiller, 1894

Suborder PopocopPina Sars, 1866

Superfamily CYTHERACEA Baird, 1850

Family Trachyleberididae Sylvester-Bradley, 1948

Subfamily Thaerocytherinae Hazel, 1967

Genus Poseidonamicus Benson, 1972

The genus was erected by Benson (1972) to accommodate a number of

deep-sea, reticulate to virtually smooth, holamphidont-hinged species. They are

characterized by certain features of their carapace morphology, such as the

absence of an ocular rib, the presence of regular, vertically aligned fossae and

muri posteromedianly, and a reticulate to punctate area medianly, the two areas

being separated by a subvertical loop in the region of the adductor scars. These

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 441

ORANGE

SHELF

Fig. 1. Locations of sediment samples containing Poseidonamicus panopsus sp. nov. on the

continental margin off south-western Africa. Isobaths at 100 m to 500 m and 1 km to 5S km.

Sample numbers and locations are listed in Table 2. Open circles are sites with living specimens.

Inset shows location of study area.

442 ANNALS OF THE SOUTH AFRICAN MUSEUM

features are present even in smooth species, such as P. nudus, if viewed in

transmitted light. Benson (1972) based the genus on five species (see Table 1;

type species P. major Benson, 1972), and the diagnosis has been amended by

Whatley et al. (1986) on the basis of their study of six additional species from the

SW Pacific (Table 1).

Poseidonamicus panopsus sp. nov.

Figs 2-3, 4A—-E, 5C

Bradleya? sp. Boomer, 1985: 42-43, pl. 3 (figs 35-36).

Derivation of name

Greek panops, all seeing. With reference to the well-developed ocular

structures of this obviously sighted species, and also because Gods and their

friends tend to be all seeing.

Material

Holotype SAM—PQ-MF0502, female LV, TBD 2719, 240 m.

Paratypes SAM—PQ-MF0503, female LV, TBD 2719, 240 m.

SAM-—PQ-MEF0504, ?male RV, TBD 2690, 271 m.

SAM-—PQ-MF0505, ?male LV, TBD 2459, 300 m.

SAM-—PQ-MF0506, female carapace, TBD 2840, 205 m.

SAM-PQ-MEF0507, juvenile LV, TBD 2840, 205 m.

SAM-—POQ-MEF0508, juvenile RV, TBD 6823, 120 m.

Dimensions (mm)

length height width

MF0502 Oot 0,51 —

MF0503 0,89 0,52 —

MF0504 0792 0,49 —

MF0505 0,89 0,48 —

MF0506 0,85 0,47 0,48

MF0507 0,64 0.37 —

MF0508 0,70 0,39 —

Diagnosis

A medium-sized, sighted species of Poseidonamicus with large hemispherical

eye tubercle and distinct internal ocular sinus. Ornament reticulate with large

open fossae, the solae of which are secondarily reticulate. SCT prominent;

medianly inflated. Dorsal margin with prominent anterior and posterior hinge

ears in LV.

Description

External features. Medium-sized (see relative size scale for genus, Whatley et

al. 1986: 388), thick-shelled and robust. Subrectangular in lateral view; strongly

inflated medianly and bluntly acuminate terminally in dorsal view. AM

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 443

| Fig. 2. Poseidonamicus panopsus sp. nov. SAM-—PQ-MF0502, holotype, LV, female,

TBD 2719, 240 m. A. External view. B. Internal view. C. Oblique internal view. D. Central

muscle scars. BE. ATE. F. PTE.

Scale bars: A—C = 100 microns; D-F = 10 microns.

444 ANNALS OF THE SOUTH AFRICAN MUSEUM

3813 1KV X126 166vm WOie

3987 1@KV X12@ 100vm WDl2

9817 18KV 4126 10@vm W012

915 1@KU ¥120 100vm WD11

Fig. 3. Poseidonamicus panopsus sp. nov. A. SAM-—PQ-MF0503, LV, female, TBD 2719,

240 m. B—-D. SAM—PQ-MF0504, RV, male, TBD 2690, 271 m. B. External view. C. ATE.

D. Internal view. E. SAM—PQ-MF0505, LV, male, TBD 2459, 300 m.

F. SAM-—PQ-MF0506, carapace, dorsal view, TBD 2840, 205 m.

Scale bars: A-B, D-F = 100 microns; C = 10 microns.

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 445

5818 1@KY X170 100¥m WD12 3911 1okv- R178 100m

een: 7 6 CE rome

¥;

9828 1@KY X17@ 160rm WO12

Fig. 4. A-E. Poseidonamicus panopsus sp. nov. A-D. SAM-—PQ-MF0507, LV, juvenile,

TBD 2480, 205 m. A. External view. B. Internal view. C. ATE. D. Oblique internal view.

E. SAM-PQ-MF0508, RV, juvenile, TBD 6823, 120m. F. Poseidonamicus major Benson,

1972, SAM-PQ-MF0495, LV, TBD 6851, 2 916 m, continental slope off south-western Africa.

Scale bars: A-B, D—F = 100 microns; C = 10 microns.

446 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Comparison of central MS patterns. A. Poseidonamicus major Benson, 1972,

SAM-—PQ-MF0497, RV, TBD 6851, 2 916m, continental slope off south-western Africa

(Dingle et al. in press). B. Poseidonamicus major Benson, 1972, USNM 174354, RV,

TIOE 363B, 2 995 m, Mozambique Channel (Benson 1972). C. Poseidonamicus panopsus sp.

nov. SAM—PQ-MF0504, RV, TBD 2690, 271 m.

Scale bars = 100 microns.

asymmetrically rounded in RV; with more pronounced antero-dorsal slope in LV;

bearing numerous small marginal denticles. PM with blunt apex below mid-height

and with short, convex denticulate postero-ventral slope and longer straight or

concave postero-dorsal slope. DM straight with prominent anterior and posterior

hinge ears in LV, particularly in the female. VM straight to very gently convex.

Greatest height at the ACA, greatest length just below mid-height; greatest width

just behind mid-length. Eye tubercle large, hemispherical and prominent, more

peripheral in the RV than the LV due to the presence of the hinge ear in the

latter. Ornament reticulate with large open fossae of fairly regular size, which are

more circular anteriorly than elsewhere. Muri somewhat preferentially aligned

vertically postero-dorsally, and horizontal to subradiate anteriorly. Solae

secondarily reticulate in a radiate pattern. SCT slightly elevated in the region of

the adductor scars, bearing small deep fossae and being embraced posteriorly and

ventrally by the mural loop characteristic of the genus. Marginal rims prominent

anteriorly and posteriorly; smooth except for a thin ridge along their crests.

Ventro-lateral ridge prominent and sub-ponticulate. A dorsal ridge, divided into

two parts, extends subparallel to the dorsal margin.

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 447

Internal features. Calcareous inner lamella rather narrow, but strongly

developed, selvage prominent. Avestibulate. Internal ocular sinus distinct, open

in juveniles, somewhat restricted at its base in adults. Hinge holamphidont. MS

comprising four undivided adductors, two frontals and two mandibular scars.

Remarks

This species differs from all other known species of the genus by possessing

a prominent eye tubercle, a distinct internal ocular sinus, an inflated carapace,

and in details of its ornamentation. The only other species of Poseidonamicus with

ocular features is P. ocularis Whatley et al., 1986. The latter species, from the

Quaternary of three DSDP sites in the southern part of the area between

Australia and New Zealand, was so named because it has a small (much smaller

than that of P. panopsus sp. nov.) but prominent eye tubercle and an internal

ocular sinus. The two species, despite the fact that they both possess strong hinge

ears in the LV, can readily be distinguished in that they possess quite different

ornament, and because P. ocularis lacks a SCT.

Although the external morphology of the new species differs significantly

from P. major Benson and there is no possibility of confusion, we include a

comparison of the central MS of the two species, because P. major is the only

other representative of the genus recorded from southern Africa (Mozambique

Channel, Benson 1972; Cape Basin, Dingle et al. in press). The MS of the two

species are very similar (Fig. 5), with the anterior scars consisting of an ovate

dorsal scar and a partially subdivided ventral scar, whereas the adductors are all

elongate/ovate, with a ‘dog’s bone’-shaped second scar. The third scar of the

adductor quartet in P. major is relatively longer than in P. panopsus.

Range

Pleistocene—Recent.

DISCUSSION

The discovery of a sighted species of Poseidonamicus living on the continen-

tal shelf off south-western Africa has a number of important implications for

understanding the evolution of the genus.

DISTRIBUTION AND ECOLOGY

Poseidonamicus panopsus sp. nov. has been found on the continental margin

of south-western Africa between latitudes 22°S and 35°S, where it has a depth

range of 120 m (mid-continental shelf) to 545 m (upper continental slope) (Fig. 1,

Table 2). With the exception of sample TBD 3769 (NW of Walvis Bay), however,

all the occurrences lie south of 28°S, and cluster into two distinct groups: on the

Orange Shelf (28°S—32°S); and off the SW Cape (34°S—35°S). The isolated site

off Walvis Bay consists of one valve, and this is separated by 6 degrees of latitude

(650 km) from the Orange Shelf population.

448 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Distribution of Poseidonamicus panopsus sp. nov. (values are means for n sites).

Area (n) Percentage Percentage Depth Range

P. panopsus mud (m)

A. Walvis Shelf (1) 3 9 223 225

B. Orange Shelf (20) 4 56 248 170-453

C. SW Cape (6) 3 54* 268 120-545

D. Overall (27) 3 a” 251 120-545

Living specimens occur at two sites on the Orange Shelf. At these sites the following parameters apply:

Percentage Percentage Depth

P. panopsus mud m

8 9 205

13 64 241

* = excludes samples 6823 and 6825 for which no mud values are available.

mud

+ P. panopsus

Percentage

100 200 300 400 500

Water depth (m)

Fig. 6. Mud content of sediment samples and relative abundance of Poseidonamicus panopsus

sp. nov. (percentage of ostracod fauna x 10) plotted against water depth. Mud and percentage

ostracod values are smoothed 5-point means. Samples TBD 6823 and TBD 6825 have not been

included (no analysis of mud contents are available). Note the inverse relationship between the

mud content and percentage P. panopsus, with peak values of the latter on the outer continental

shelf (c. 250 m water depth).

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 449

Table 3 lists some of the parameters of the areas of occurrence. In the

Orange Shelf and SW Cape populations, the species has a similar depth range

(170-453 m, and 120-545 m, respectively), and the mean mud contents of the

sediments in which the species is recorded (53% and 54%) and the mean

abundance of the species (4 % and 3 %) are also very close. These two areas are

separated by approximately 200 km of shelf which is barren of P. panopsus. Plots

of the abundances of the species (as percentage of the ostracod population)

against the mud contents of the sea-floor sediments (Fig. 6) indicate that the

species peaks (up to 60 % total ostracod population) at about 250 m water depth,

where the mud content of the sediment is lowest (mean of 30 %). On the adjacent

inner shelf and outer shelf/upper slope, where the mean mud contents are >60 %,

the abundance of P. panopsus rapidly declines. This indicates that the species is

generally not tolerant of high mean mud values (>70 %). However, the fact that

the region between the Orange Shelf and SW Cape populations, which is barren

of the species, has a mean mud content of 30 per cent (compared to 50 % for all

sediments in the two populated areas) suggests that a minimum mean mud

threshold (30 %) is one of the factors necessary to sustain the species.

+ %P. panopsus

Thousands

150 250 350 450 550

Water depth (m)

Fig. 7. Abundance of ostracod population (number of valves/100 g sediment) and relative

abundance of Poseidonamicus panopsus sp. nov. (percentage of ostracod fauna x 100) plotted

against water depth. Ostracod population and P. panopsus values are smoothed 3-point means.

Samples TBD 6823 and TBD 6825 are not included (no ostracod population abundance values

available). Note the high ostracod population values on the mid-shelf, which decline rapidly into

deeper water, with a concomitant increase in relative abundance of P. panopsus.

450 ANNALS OF THE SOUTH AFRICAN MUSEUM

A further limiting factor within the two main P. panopsus population centres

is the abundance of the whole ostracod fauna (Fig. 7). Mean values (measured as

number of valves/100 g of original dry sediment) peak on the inner shelf (>500

valves/100 g in <200 m water depths), and fall rapidly towards the outer shelf and

slope. Concomitant with this decline in the overall abundance of the ostracod

fauna, there is a marked increase in the percentage of this fauna made up by

P. panopsus. This trend suggests that the species has limited toleration of

competition from large numbers of individuals belonging to other species,

although at present the nature of this factor remains unknown.

Living specimens were collected at two locations (TBD 2840 and 2361), both

in the Orange Shelf area, but plots of the population structure show that in both

this area, and within the SW Cape occurrences, the mean percentages of adult

valves varies between 20 and 70 per cent, with the higher values concentrated in

water depths between 200 m and 250 m in the Orange Shelf area (Figs 8, 9).

These assemblages of mixed adults and juveniles suggest that both the main areas

of occurrence reflect viable extant populations. The isolated site off Walvis Bay is

a single dead adult valve, and we are not in a position to determine its true status.

population seems unlikely, but we have good sample coverage over the whole of

the adjacent continental shelf, and it is equally unlikely that a population centre

larger than 300 km’ would have been undetected.

100

80 Orange Shelf

Percentage

smoothed data

raw data

Latitude °S

Fig. 8. Percentage of adult valves of Poseidonamicus panopsus (raw and smoothed 5-point

means) plotted against latitude. Sample TBD 3769 off Walvis Bay is omitted. Mean values:

Orange Shelf = 49 per cent adult, SW Cape = 29 per cent adult.

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 451

Percentage

100 200 300 400 500

Water depth (m)

Fig. 9. Percentage of adult valves of Poseidonamicus panopsus (smoothed 5-point means)

plotted against water depth. Note that peak values (>60 %) lie around 200 m water depth, which

is slightly shallower than the peak values of abundance of the species (Fig. 6—250 m).

The three deepest sites at which the species was recovered (TBD 1694, 425 m;

TBD 3577, 453 m; and TBD 1697, 545 m) all consist of single juvenile valves,

which possibly indicate allochthonous occurrences due to downslope postmortem

transportation. If this is the case then the true lower depth limit of the species is

350 m.

The oceanographic climate off south-western Africa is complex, with

large-scale upwelling over the continental shelf area (Shannon 1985). Figure 10

shows sea-floor variations in temperature, salinity, and dissolved oxygen across

the continental margin at the southern end of the Orange Shelf (32°S). The upper

depth limit of P. panopsus on the Orange Shelf is 170 m, and this occurs at

approximately the same depth as two important changes in physico-chemical

parameters: the base of the inner shelf thermocline (c. 13°C-<9,5°C), and the

western limit of the continental shelf low dissolved oxygen zone. The slightly

shallower upper depth limit (120m) off the SW Cape (34°S) may reflect

somewhat lower sea-floor temperatures at shallower depths: the 9°C isobath

intrudes to depths of 100 m for much of the year in this area (see Shannon 1985,

fig. 21).

Low dissolved oxygen values (<2 ml/1) may also be important limiting factors

in the latitudinal distribution of the species. A major source of oxygen deficient

water lies to the north of the Orange Shelf, where Chapman & Shannon (1985)

452 ANNALS OF THE SOUTH AFRICAN MUSEUM

[ae SW Cape Ee eet re

aba

«a——.. Orange Shelf —____._»

Top of SMZ |

0 200 400 600

Water depth (m)

+ __ Salinity

oO temperature l

Fig. 10. Variations in water temperature, salinity, and dissolved oxygen on the sea floor along

a transect across the continental margin at approximately 32°S. Data from Fuglister (1960),

Stander (1964), Shannon (1966, 1985), Shannon & Van Rijswijk (1969). Depth ranges of

P. panopsus on the Orange Shelf and off the SW Cape are indicated by arrows. Note that the

western limit of the ‘oxygen deficient zone’ (<2 ml/l, as defined by Chapman & Shannon 1985)

lies at approximately 180 m water depth (Stander 1964). This is somewhat deeper than predicted

by the straight line connecting the two data points used to create the dissolved oxygen curve.

Parameters: temperature =°C, salinity = [(°/99 —34) x 10], dissolved oxygen = ml/I.

Abbreviations: oxygen = dissolved oxygen, Ox. min. = oxygen minimum, SMZ = salinity

minimum zone of AAIW.

identified a double low on the continental shelf, whereas a smaller, but persistent

source is centred on St Helena Bay. We suspect that the latter may contribute to

the P. panopsus-barren region between 32°S and 34°S. Similarly, the oxygen-low

area immediately off the Orange River mouth (Chapman & Shannon 1985, fig. 6)

may be a factor restricting the species to water deeper than 170 m on the Orange

shelf (compared to 120 m off the south-western Cape).

At the lower depth limit (545 m or 350 m, depending on whether our three

deepest specimens are allochthonous), gradients in the variation of sea-floor

temperature and dissolved oxygen are relatively low, but Shannon (1985) has

located the top of the salinity minimum zone (SMZ) within the Antarctic

Intermediate Water mass (AAIW) at approximately 600 m along most of the

continental margin of south-western Africa. Data in Fuglister (1960) show this

phenomenon to be closer to 500 m at 32°S (Fig. 10), and Dingle et al. (in press)

have recognized the bottom of the SMZ (at about 950 m) to be a major faunal

barrier, separating neritic and bathyal ostracod faunas. We suspect that the lower

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 453

depth limit of P. panopsus is regulated by the level of the top of the SMZ within

the AAIW mass.

In summary, we believe that the following environmental parameters influ-

ence the distribution of Poseidonamicus panopsus sp. nov.:

1. The species prefers a mud content in the sediment of between 30 and 50 per

cent, with the lower value limiting its distribution.

2. It is most abundant where it lives within ostracod populations with low

abundances (<400 valves/100 g).

3. Limiting water temperatures are approximately 9,0°C-—5,8°C.

4. Limiting salinities are 34,70-34,40°V/o9.

5. Limiting dissolved oxygen values are 2,7—4,2 ml/I.

6. The species tolerates the nutrient-rich conditions of the upwelling cells

associated with the Benguela system, but where these cells result in intense

biological productivity and oxygen deficiencies in the water column (to the north

of the Orange Shelf area, and between the Orange Shelf and the SW Cape area),

dissolved oxygen values on the sea floor are too low for its survival.

Percentage

fee! a

Water depth (km)

Fig. 11. Relative abundances of Poseidonamicus panopsus sp. nov. and Poseidonamicus major

Benson (percentage of ostracod population) plotted against water depth. The populations of the

two species are separated by 1 500 m of water on the continental slope which contains the

following physico-chemical barriers: salinity minimum zone of the Antarctic Intermediate Water

mass; the shear zone of the Antarctic Intermediate Water/North Atlantic Deep Water masses

boundary; and the top of the core of the North Atlantic Deep Water. Note also that

Poseidonamicus major is a more important component of the abyssal ostracod fauna than is

P. panopsus of the neritic ostracod fauna (means 22 % and 3,3 %, respectively).

Abbreviations: AAIW = Antarctic Intermediate Water; CZ = core zone; ML = mixed layer;

NADW = North Atlantic Deep Water; SMZ = salinity minimum zone; SZ = shear zone.

454 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dingle et al. (in press) discuss some of the factors that may be responsible for

confining various deep-water (>950 m) ostracod assemblages to their respective

depth ranges. They concluded that the upper depth limit of P. major off

south-western Africa is probably controlled by the position of the top of the core

zone of the North Atlantic Deep Water mass (NADW), above which higher

temperatures (>3,0°C) and lower salinities (<34,86°/o9) occur. Figure 11 shows

the depths and abundances (as percentage of ostracod fauna) of samples

containing Poseidonamicus major and P. panopsus off south-western Africa and

emphasizes that, although the two species are both extant, they are separated by

1 500 m of water column that contains two major barriers to faunal mixing: the

SMZ of the AAIW;; and the contact zone between the AAIW and the NADW. As

Dingle et al. (in press) point out, both these are powerful barriers that control the

Neritic/Upper Bathyal, and Upper Bathyal/Lower Bathyal faunal boundaries,

respectively. In addition, the top of the NADW core regulates the Lower

Bathyal/Abyssal faunal boundary. Because the water masses that give rise to the

controlling physico-chemical gradients have been in existence since at least

mid-Tertiary time, the separation of the two Poseidonamicus species off south-

western Africa is likely to have been a long-term phenomenon. i

GENERAL CONSIDERATIONS

Benson (1972: 21), apart from speculating that the ancestry of Poseidon-

amicus ‘reaches far back into the Cretaceous’, could not, at the time, suggest a

possible ancestor for the genus. Subsequently, Whatley et al. (1983) suggested

that it evolved in the area of the SW Pacific Ocean, and more particularly in the

region between Australia and New Zealand, where they encountered it in DSDP

samples dating back to the Lower Eocene. The two oldest species that they found

in the Lower Eocene, P. rudis and P. robustus (Whatley et al. 1986), were shown

to exhibit a considerable resemblance to certain Upper Cretaceous taxa from

western Australia. These authors (1983: 497, fig. 2) suggested that Hermanites

sagitta Bate, 1972, from the Campanian of the Carnarvon Basin, or some similar

species, could be the ancestor of Poseidonamicus. They argued that H. sagitta

could have given rise to the hypothetical ancestral Poseidonamicus species, which

would probably have been sighted, but which gave rise, once the genus had been

involuntarily inducted into the deep sea, to all the blind species in the deep-water

environments of the world’s oceans. Since the two Australian Lower Eocene

species are blind (a characteristic of all podocopid ostracods living in waters

deeper than about 500 m) then this induction must have taken place at some stage

prior to the Eocene.

The same authors suggested, however (1983: 479, fig. 2; 1986: 391), that

P. ocularis evolved from a fully sighted, shallow, shelf-dwelling ancestor in the

Quaternary. Poseidonamicus ocularis was only recovered from bathyal environ-

ments in the southern part of the area between Australia and New Zealand

(DSDP sites 207, 277 and 281) and only from Quaternary strata. They argued

(1983: 479) that the discovery of P. ocularis ‘implies that there has persisted in

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 455

shallow waters of this area until recent times a population with eyes, derived

direct from a postulated sighted ancestor’. They also comment on the ‘exciting

prospect that a living sighted species of the genus may be encountered in due

course’.

The discovery of the sighted shelf species P. panopsus sp. nov. has justified

these expectations, but the present authors admit to surprise that it should have

been encountered off the southern African Atlantic coast rather than in

Australasian waters. Given the Quaternary age of P. ocularis, however, it

remains quite conceivable that a shallow-water shelf species of the genus remains

to be discovered living in the antipodes.

In logic, P. panopsus must be considered a lineal descendant of a late

Cretaceous or early Palaeogene hypothetical sighted ancestral Poseidonamicus

species that subsequently give rise to all the blind deep-water and sighted

shallow-water species of the genus. Direct contiguity of continental shelf

environments between what is now southern Africa and Australasia probably

ceased at about 100 m.y. Bp (Albian), when southern Africa finally separated

from the Falkland Plateau extension of South America (see Dingle et al. 1983).

Circuitous shallow-water connections probably continued until Turonian times via

the Walvis Ridge/Rio Grande Rise archipelago across the South Atlantic, and the

Drake Passage shallows between South America and Antarctica but, since the

latter date, the two areas have been separated by barriers of abyssal depths.

Consequently, the ancestry of Poseidonamicus panopsus is uncertain, given the

fact that the genus probably arose in Australasia from shallow-water Upper

Cretaceous stock. Dingle (1981) discussed the similarity between southern

African and western Australian Campanian shallow-water ostracods, and

recorded a relatively high similarity (27 %) at the generic level. However, there

were no species in common, and no evidence suggests any contact between the

ostracod populations since at least the mid-Cretaceous. Hermanites kennedyi

Dingle (Campanian) from SE Africa is similar to H. sagitta Bate, a possible

progenitor for the SW Pacific Poseidonamicus populations (Whatley et al. 1983),

but even the possibility of homeomorphy from a similar ancestor is not attractive,

because there is no record of Poseidonamicus in the Tertiary of southern Africa.

A lineal connection with the Australasian stocks seems the most likely solution,

but it may be that P. panopsus or its ancestors do not have a long history on the

continental margin of southern Africa, and migrated into the area relatively

recently (?late Tertiary) from other areas (such as eastern Africa). Whatever the

case, it is remarkable that sighted species have survived so long apart (to the

Recent in southern Africa, and to at least the Pleistocene in Australasia) without

either becoming extinct or being subject to major evolutionary change. Although

this can only be confirmed if and when sighted shelf species of Poseidonamicus are

recovered from Tertiary shallow-water deposits in the Southern Hemisphere, it

does suggest that, when compared to the considerable degree of evolutionary

change experienced by deep-water species (Whatley 1985), the genus has been

more stable in shallow environments.

456 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

RVD gratefully acknowledges research facilities provided by Drs M. A.

Cluver and Q. B. Hendey at the South African Museum, and Dr A. R. Lord at

University College, London. The samples were collected whilst RVD was in the

Marine Geoscience Unit at the University of Cape Town, and the University, the

South African National Committee for Oceanographic Research, and the Foun-

dation for Research Development are thanked for funding sea-time. The authors

are grateful to the Editorial Board of UCT and the South African Museum for

subventing publication costs.

REFERENCES

Bate, R. H. 1972. Upper Cretaceous ostracods from the Carnarvon Basin, Western Australia.

Special Papers in Palaeontology 10: 1-85.

Benson, R. H. 1972. The Bradleya problem, with descriptions of two new psychrospheric

genera, Agrenocythere and Poseidonamicus (Ostracoda: Crustacea). Smithsonian Contri-

butions to Paleobiology A12: 1-138.

BENSON, R. H. & PEypouquetT, J. P. 1983. The upper and mid-bathyal Cenozoic ostracode

faunas of the Rio Grande Rise found on Leg 72, Deep Sea Drilling Project. Jn: BARKER, P. -

F. et al. Initial Reports of the Deep Sea Drilling Project 72: 805-818. Washington: U. S.

Government Printing Office.

Boomer, I. 1985. A reconnaissance survey of Recent and Holocene Ostracoda from the

continental shelf of South West Africa. Unpublished M.Sc. thesis, University College,

London. 98 pp.

Brapy, G. S. 1880. Report on the Ostracoda dredged by ‘HMS Challenger’ during the years

1873-1876. Report of the Scientific Results of the Voyage of HMS Challenger

(Zoology, 1) 3: 1-184.

Cotes, G. & WuaTLey, R. C. 1989. New Palaeocene to Miocene genera and species of

Ostracoda from DSDP sites in the North Atlantic. Revista Espanola de

Micropaleontologia 21 (1): 81-124.

CHAPMAN, P. & SHANNON, L. V. 1985. The Benguela ecosystem. Part II. Chemistry and related

processes. Oceanography and Marine Biology Annual Review 23: 183-251.

DincLe, R. V. 1981. The Campanian and Maastrichtian Ostracoda of south-east Africa.

Annals of the South African Museum 85 (1): 1-181.

DINGLE, R. V., Lorp, A. R. & Boomer, I. In press. Deep-water Quaternary Ostracoda from

the continental margin off south-western Africa (SE Atlantic Ocean). Annals of the South

African Museum.

DINGLE, R. V., SIESSER, W. G. & Newton, A. R. 1983. Mesozoic and Tertiary geology of

Southern Africa. Rotterdam: Balkema.

FREWIN, J. 1987. Palaeogene ostracods from the South African continental shelf. Unpublished

M.Sc. thesis, University of Cape Town. 171 pp.

FUGLISTER, F.C. 1960. Atlantic Ocean atlas of temperature and salinity profiles and data from the

International Geophysical Year of 1957-1958. Woods Hole: Woods Hole Oceanographic

Institution.

SHANNON, L. V. 1966. Hydrology of the south and west coasts of South Africa. Investigational

Report Division of Sea Fisheries, South Africa 58: 1-22.

SHANNON, L. V. 1985. The Benguela ecosystem. Part I. Evolution of the Benguela, physical

features and processes. Oceanography and Marine Biology Annual Review 23: 105-182.

SHANNON, L. V. & vAN Ruswick, M. 1969. Physical oceanography of the Walvis Ridge region.

Investigational Report Division of Sea Fisheries, South Africa 70: 1-19.

STANDER, G. H. 1964. The Benguela Current off South West Africa. Investigational Report

Marine Research Laboratory, South West Africa 12: 1-43.

Wua Ley, R. C. 1983. Some aspects of the palaeobiology of Tertiary deep-sea Ostracoda from

the S.W. Pacific. Journal of Micropalaeontology 2: 83-104.

A NEW SIGHTED SPECIES OF POSEIDONAMICUS 457

WuHuatTLeEyY, R. C. 1985. The evolution of the ostracod genera Bradleya and Poseidonamicus in

the deep sea Tertiary and Quaternary of the South-West Pacific. Special Papers in

Palaeontology 33: 103-116.

WuatLey, R. C. & Cotes, G. 1987. The late Miocene to Quaternary Ostracoda of Leg 94,

Deep Sea Drilling Project. Revista Espanola de Micropaleontologia 19 (1): 33-97.

WHATLEY, R. C., Downina, S. E., KESLER, K. & HARLow, C. J. 1986. The ostracod genus

Poseidonamicus from the Cainozoic of D.S.D.P. Sites in the S.W. Pacific. Revista

Espanola de Micropaleontologia 18 (3): 387-400.

WHATLEY, R. C., HARLow, C. J., DowNnING, S. E. & KESLER, K. 1983. Some observations on

the origin, evolution, dispersion and ecology of the genera Poseidonamicus Benson and

Bradleya Hornibrook. In: Mappocks, R. F. ed. Applications of Ostracoda, Proceedings of

the Eighth International Symposium on Ostracoda, Houston, 1982: 51-77.

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, ctc.., 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: Nickles, 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.

+ SPECIAL HOUSE RULES

Capital initial letters

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

? ‘ ,

_*... the Figure depicting C. namacolus .. .’: ‘. ..in C. namacolus (Fig. i ae

(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.

R. C. WHATLEY

R. V. DINGLE

FIRST RECORD OF AN EXTANT, SIGHTED,

SHALLOW-WATER SPECIES

OF THE GENUS POSEIDONAMICUS BENSON

(OSTRACODA) FROM THE CONTINENTAL

MARGIN OF SOUTH-WESTERN AFRICA

MCZ

LIBRARY

AUS 06 1999

HARVARD

UNIVERSITY

O 5 oS >

a) ANNALS OF THE SOUTH AFRICAN MusEuM 98 (1)

University of Cape Town South African Museum

f WALVIS BAY

DEEP-SEA SEDIMENTARY ENVIRONMENTS AROUND SOUTHERN AFRICA : in : se

(SE ATLANTIC & SW INDIAN OCEANS) ae

R. V. DINGLE, G. F. BincH, J. M. BREMNER, R. H. DE DECKER, A. DU PLEssISs, J. C. ENGELBRECHT,

M. J. FINCHAM, T. FITTON, B. W. FLEMMING, R. I. GENTLE, S. W. GOODLAD, A. K. MARTIN,

E. G. MILLs, G. J. Morr, R. J. PARKER, S. H. ROBSON, J. ROGERS, D. A. SALMON, W. G. SIESSER,

E. S. W. SIMPSON, C. P. SUMMERHAYES, F. WESTALL, A. WINTER, & M. W. WOODBORNE.

LUDERITZ [ SHELF SOURCE AREAS STEADY — STATE ACCRETION & EROSION

terrigenous muds sediment drift

] diatomaceous muds drift crest LAYER B sediments

West Coast carbonate sands & muds basin fill

| East Coast dunes | ridge and plateau billows & other LAYER A sediments

relict shelly sands sediment waves Mfolozi R

authigenic sediments (>5% content) terrigenous construction: submarine fan

Orange! | f ? Naud KS

Pre-Mesozoic basement constructional terrace / Ase

LLOCHTHONOUS ENVIRONMENTS & FEEDERS SCOUlmsCOle m2 ONe

turbidites, channel-fill, LAYER C sediments “+ _+ ] ferro-manganese deposits

chute deposits

canyons & feeder valleys outcrop

rise lobes =] surface ridge ; deep-water basement

LAYER B

slumps & slides sediments buried ridge

probable allochthonous masses ITIMMIII seamount, uncertain composition

glide planes & scars undefined areas

are fissured & unstable zones

Olifants

MER = Marginal Fracture Ridge ‘bathymetry in km

CANYONS ——77 approximate limit of west coast aerosols

'.* VEMA

Pa 1 Ntabende 3 Egosa 5 Mzimvubu 7 Black Beacon 9 Hamburg 11 Fish 13 Padrone

2 Mzimkulu 4 Mbotyi 6 Morgan 8 Cove Rock 10 Stalwart 12 Boesmans 14 St Francis.

Mercator Projection. Scale 1:3 200000

Fish R.

C. Padrone

CAPE TOWN

Breede R

3).

— aa

little or no sediment cover

SCHMITT-OTT

10°

MCZ_.

LIBRARY

aug 06 1992

HARVARD

UNIV ERSITY

ft-hpe)

WALVIS BAY

University of Cape Town South African Museum

BATHYMETRY AROUND SOUTHERN AFRICA e

(SE ATLANTIC & SW INDIAN OCEANS) ea) 2

4 R. V. DINGLE, G. F. BIRCH, J. M. BREMNER, R. H. DE DECKER, A. DU PLESSIS, J. C. ENGELBRECHT,

\ M. J. FINCHAM, T. FITTON, B. W. FLEMMING, R. I. GENTLE, S. W. GOODLAD, A. K. MARTIN,

\ E. G. MILLs, G. J. Morr, R. J. PARKER, S. H. Rosson, J. ROGERS, D. A. SALMON, W. G. SIESSER,

E. S. W. SIMPSON, C. P. SUMMERHAYES, F. WESTALL, A. WINTER, & M. W. WOODBORNE.

LUDERITZ ta

Mercator Projection. Scale 1 3 200 000

Isobaths: regional 1 km

Zips »\\\

DURBAN Cpe p ty (\\N A {\)

Vem}. (PS n\} ;

LWT,

ASIN

Mzimkulu R.

Y i, 3

PORT SHEPSTONE WY Uy

Figures 7 |

draftedby INDY WOODHRORD

St a ha

MEER ah gh ae fi ME OE AE 2

Be a ern ct oa ae ote NE a Eon pe ae ate aiede i ’

a ce AR TE SO ate, achat Jens GPE Oe ca ths

‘ -H “eee en pn gtndinsneop tynk recat | ni f

= : Ne At Naan Ain nhasat eae ' . . 3 men ’ tun

tah eet 5 6h ttl ethan ER TREND Fold ot SD BAD hiatal ot , ‘ or ;

See Ce ae . tee 9 e te oar f ' euadce : : we an

iil

<r ty OTT eS

. wa < . ?

Tha SRO + ete ‘ ‘ ‘ :

see Sa nansan® % wy a, ‘ . . .

t AF adadaten do % : 7 hy 4

29.8 MrADPO MS.» L i Pig \ ae . ;

oy A ER i “sy :

ere ’ et vo Ns ‘ . . ‘ :

. : A es : : 8 at «a

a>. » . on ‘ ’ . .

eee ee 's bd MAD ' : . 4

2 Pap warm ¢ wy 4 . . 7 . < .

ere tts aa 6 bree . P ’ ’ . ‘ : : :

Sopa yeast? add “ ‘ ws ‘ ’ : sere

Os. GAAP Fes fs” é Mate . : . . ‘ i

SPN ede OD prehant® Ss foes ye ¢ te. use a

ett eet et tL fOr Or tay Soa

RL Qdumessr Vase + shaeeae va SUR WN “yh itive rs eee

aS “ea yeiad : . °% ve . eas’ $8 oan ; . ‘ a

Tm eegn.t . . sae . 3 ‘ :

Beas bowde Os ree TY) qe ery . . 3

vA ebasg deed : : ‘ sesh gost ‘ ,

eer hy . % ee . . oe ‘ ’

soba tes eu 8 ast . 1 ‘

so peavesat . a sia NY v5 7

: : Bearers ‘ sasbinny x oe . . ‘ ey hg- Ma

PES ar wy Cr coer | St ey S| A er ices . , ‘ ‘ .

LPNS SSL \*_@ x . oth . rm : .

¢ . pare 08 Lg etOtsn* 84 ‘ : ‘ a 48 . ‘

Coe rer Cy wa a yy . c% Rm Sew e .

Wi a. a we ' on ea . y a aa . . .

rere eee arr ‘ ‘ xe . . . &

. . . ee Gta . ace , .

et, ie . . . ‘ ‘

OL ‘ees . : wy hss re toe ‘

sales. Rt e-'5 Are eae ak ae fis . .

Cass . SGice sy ear outta ty one 5

ake. ee eke ste oe ‘ .

seth) ese « sesh JET ote our e ae Fis Sey 9? . . n

wae . . ‘ an ‘ Cr A N i

ee ‘ stem ay Sete a'< . .

see * 4 a . . :

eset . "ee. . ‘ rs 7 s . :

ae ‘ . ara Sate tees a? ‘ .

eee . “0a « ‘ ee ’ . x . ;

aot Ae LET THY vpn € : r . Ww : ‘

: guy Choe de ge \ ‘ =?

be’ se bebe yet hs . Nie ga SS P ’ . Paar A ‘

TS A + % 2 : <5 . oer 7 ‘

. eo ‘ ‘

. roe . hs . eee hd . ,

. 4 ate 4 . , ote : 7

a Setar Wey vee . 2 . . .

caren 3 ‘ ste : wate

a ere Oe . . .

. oe 3 re ta = 85 . . .

A+ * Feeb eye pense § are tet ae ss dry ly a. gt Pe Se

eee ore

co oa

SVE ae

ne eer et ‘ :

pe nee eee Fare < - . : 2 < J are 7 iL

fiw etary sy me %

Oe ee ee ’

es eee

ives * : ‘ : . i, Were

cep e . a . :

ee tt ;

Oe me ee se ae

a a ti eateeT

Be tag Be,

Prarie ow, oleratl

ata £28 ye oH ERO

ee tot dee

Feet ea ear

eee Eee

ONES HRN rg fs ; one

Sk hee ee Oe : ’ ! r¢) ‘ ) woot . : A . ene .

Ns eed : 3 se R