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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
VOLUME 74 BAND
THE TRUSTEES OF THE DIE TRUSTEES VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD
1977-1978
# SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY
THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE
524
LIST OF CONTENTS
Avery, D. M.
Past and present distribution of some rodent and insectivore species in the southern
Cape Province, South Africa: new information. (Published December 1977.)
EAsTWwoop, E. B.
Notes on the scorpion fauna of the Cape. Part 3. Some observations on the distri-
bution and biology of scorpions on Table Mountain. (Published January 1978.)
EASTWOop, E. B.
Notes on the scorpion fauna of the Cape. Part 4. The burrowing activities of some
scorpionids and buthids (Arachnida, Scorpionida). (Published January 1978.)
GLAESSNER, M. F.
Re-examination of Archaeichnium, a fossil from the Nama Group. (Published April
1978.) .. ie me
GRIFFITHS, C.
The South African Museum’s a tea Naude cruises. Part 6. Giese area
November 1977.) é ;
GRINDLEY, J. R.
A new species of Tortanus (Crustacea, Copepoda) from South Africa. (Published
January 1978.) : a se We :
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. A Jauberticeras from
the Mzinene Formation (Albian). (Published November 1977.)
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. A Flickia from the
Cenomanian of northern Zululand. (Published January 1978.)
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family
Lytoceratidae Neumayr, 1875. (Published February 1978.)
_KENSLEY, B.
The South African Museum’s Meiring Naude cruises. Part 5. Crustacea, Decapoda,
Reptantia and Natantia. (Published December 1977.) .. :
KENSLEY, B.
The South African Museum’s Meiring Naude cruises. Part 7. Marine Isopoda.
(Published January 1978.) ..
KLINGER, H. C. see KENNEDY, W. J.
McKENZIE, K. G.
Illustrated generic key to South African continental Ostracoda. (Published Novem-
ber 1977.) ;
MILLARD, N. A. H.
The Se distribution of southern African hydroids. (Published January
1978.).. : a Ke me By aN es S% id
Page
201
Vipes)
249
335
105
US)
ZAM
PEST)
13
2S
45
159
NEW GENERIC NAMES PROPOSED IN THIS VOLUME
Page
Izinkala Griffiths, 1977 ae ie ra A: he ra ar od An 115
Kapcypridopsis McKenzie, 1977 Bes Ae a: a ae ae 4 ae 46
Sarscypridopsis McKenzie, 1977 ae ae at fe Bat Ls ake He 48
Spinarcturus Kensley, 1978 .. as Me Bie a as ae Spe Se)
Valettiella Griffiths, 1977 5 Sc =i on oe Be si si’ a 2G)
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. ; ey Ct eA et Ae “an
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VOLUME 74 PART 1 NOVEMBER 1977 | ISSN 0303-2515
MUS. COMP. ZOOL.
LIBRARY
‘SAN 3 1 1978
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Konan, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
November 1977 November
Part 1 Deel
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
A JAUBERTICERAS FROM THE MZINENE
FORMATION (ALBIAN)
By
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
Cape Town Kaapstad
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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
A JAUBERTICERAS FROM THE MZINENE FORMATION (ALBIAN)
By
WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford
&
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
(With 6 figures)
[MS. accepted 28 April 1977]
ABSTRACT
The ammonite subfamily Gabbioceratinae Breistroffer, 1953, is represented in the
Cretaceous of southern Africa by a unique specimen of Jauberticeras collignoni Murphy from
the Albian Mzinene Formation exposed along the Mzinene River, Zululand. The specimen
shows details of suture, lirae and constrictions not previously known in the species, until now
based on the unique Madagascan holotype alone, allowing a fuller diagnosis, description and
comparison with other species. The holotype and the types of the closely related species
J. jaubertianum (d’Orbigny) and J. villoutreysi Murphy are reillustrated for comparative
purposes.
CONTENTS
PAGE
Introduction : : ; i i l
Systematic palaeontology . : ; 2
Acknowledgements . ‘ , 4 1]
References : , ‘ , ; 11
INTRODUCTION
In two previous papers (Kennedy & Klinger 1977, and in press) the authors
have documented the representatives of the great majority of the ammonite
superfamily Tetragonitaceae Hyatt, 1900 known from the Cretaceous sediments
of Zululand and Natal. In the course of examining the collections of the late
Dr E. C. N. van Hoepen, recently transferred to the South African Museum
from the National Museum Bloemfontein, one of the authors (H. C. K.) recog-
nized a further representative of this superfamily, in the form of a Jauberticeras
Jacob, 1907, from the Upper Albian Mzinene Formation of Zululand.
Jauberticeras is one of the two genera referred to the subfamily Gabbio-
ceratinae Breistroffer, 1953 (the other is the genus Gabbioceras Hyatt, 1900);
the specimen represents not only the first record of the genus from southern
Africa, but also the first record of the subfamily from the region.
l
Ann. S. Afr. Mus. 74 (1), 1977: 1-12, 6 figs.
D) ANNALS OF THE SOUTH AFRICAN MUSEUM
SYSTEMATIC PALAEONTOLOGY
Class CEPHALOPODA Cuvier, 1797
Order AMMONOIDEA Zittel, 1884
Suborder LYTOCERATINA Hyatt, 1889
Superfamily TETRAGONITACEAE Hyatt, 1900
Family Tetragonitidae Hyatt, 1900
Subfamily Gabbioceratinae Breistroffer, 1953
The subfamily Gabbioceratinae has been the subject of two exhaustive
accounts in recent years (Wiedmann 1962; Murphy 1967), and there is a diver-
gence of opinion over which species should be referred to Gabbioceras versus
Jauberticeras, as well as over the systematic position of a number of species
regarded by Wiedmann as Gabbioceras and by Murphy as Eofetragonites. More
serious is the disputation of the position of Parajaubertella Matsumoto, 1943,
regarded by Wiedmann as a synonym of Gabbioceras and by Murphy as an
independent homoeomorphous offshoot of the Eogaudryceras stock. There is,
however, agreement that the subfamily has its origin in the group of Eogaudry-
ceras (Eogaudryceras) numidum (Coquand). From this the line of descent runs
to Gabbioceras, with G. lamberti (Breistroffer) as the earliest known member of
that genus giving rise to Jauberticeras via J. jaubertianum during early late, or
late early Aptian time. Jauberticeras is confined to the late Aptian and Albian;
Gabbioceras extends to the Cenomanian.
Genus Jauberticeras Jacob, 1907
Type species
Ammonites jaubertianus d Orbigny (1850: 200, figs 9-10) by original designa-
tion (see Howarth 1974 for clarification of the date and place of publication of
Jauberticeras and of Ammonites jaubertianus as type species).
Diagnosis
Relatively small cadicones with a funnel-shaped umbilicus, no flank and a
marked angulation demarcating umbilicus from the broad, flattened venter.
Shell surface lirate with constrictions (sometimes only visible on the mould).
Suture with three umbilical lobes.
Discussion
The authors take this opportunity to illustrate (Fig. |!A—G) a plaster cast
of the lectotype of J. jaubertianum, the original of which is from the Aptian of
Hyeges, Basses Alpes, France. When compared with Gabbioceras (a typical
example of which is shown in Fig. 2A—G), the two genera are best distinguished
on whorl section (following Murphy 1967: 604): in Gabbioceras the venter is
strongly curved (Fig. 2A, C, D, G): in Jauberticeras it is flat (Fig. 1A—B, D-F).
CRETACEOUS FAUNAS FROM SOUTH AFRICA 3
The angulation is commonly sharper in Jauberticeras when compared with early
Gabbioceras (Murphy 1967, fig. 1).
Occurrence
Late Aptian to mid Albian of southern France, Spain, the Caucasus, north
Africa, Madagascar and Zululand.
F G °
Fig. 1. Jauberticeras jaubertianum (d’Orbigny). The lectotype from the Aptian of Hyéges,
Basses Alpes, France (Peticlerc Collection, Sorbonne, Paris). A-C x 2; D-G ~x 1.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Gabbioceras lamberti (Breistroffer). BMNH C10780, from the Aptian of Castellane,
Basses Alpes, France. A-C x 2; D-G x 1].
CRETACEOUS FAUNAS FROM SOUTH AFRICA 5
Jauberticeras collignoni Murphy, 1967
Figs 3-5
Jauberticeras latecarinatum Collignon non Anthula, 1963: 28, pl. 251 (fig. 1080).
Jauberticeras collignoni Murphy, 1967: 606, pl. 64 (fig. 16).
Holotype
M. Collignon Collection no. 435, the original of Collignon (1963: 28, pl. 251
(fig. 1080)) by monotypy. Reproduced here as Fig. 3A-H. From the Albian
sensu lato of Betioky, Madagascar.
Material .
Specimen D293 in the E. C. N. van Hoepen Collection.
Emended diagnosis
A Jauberticeras with a blunt lateral angle, moderately expanding whorls,
weak lirae and irregularly spaced prorsiradiate convex constrictions and a
prominent second saddle on the internal suture.
Description
Specimen D293 is a beautifully preserved mould of a phragmocone 21,5 mm
in diameter, infilled with sparry calcite, and retaining a short portion of the body
chamber filled with black quartzose glauconitic sandstone. Traces of the original
shell material with delicate external ornament are preserved on the umbilical
wall, but on the outer surface the shell is partially or wholly absent. Dimensions
of the specimen, and others with which it is compared, are shown below.
(D = diameter; Wb = whorl breadth; Wh = whorl height; U = umbilical
diameter from seam to seam; K = umbilical diameter from keel to keel.
Figures in parentheses are dimensions expressed as percentages of the total
diameter.)
D Wb Wh Wb:Wh U K
J. collignoni 22,4 (100) 16,3 (74) 6,9 (31) 2,36 ~—-9,8 (44)_—‘:18,7 (84)
D293 8200), 13.3,a)) 5.5 CO. 24N 74441) 14.978)
MC435 OFS) (OO) lS (68) 523 (27) ae 4 8,1 (42) 14,9 (77)
J. yilloutreysi |
BMNH C68145 27,8 (100) 18,5 (67) 8,8(32) 2,13 12,0(43) 22,5 (81)
J. jaubertianum
lectotype SOR COO) Fe IOSD Os G2) e485 058833) = (=)
The coiling is very evolute, so that the angulation of the successive whorls
is exposed in the umbilicus, giving it terraced, rather than evenly sloping sides.
The umbilical wall is gently convex, whilst the angulation between it and the
venter is abruptly rounded and blunt. The venter is broadly and evenly rounded,
the distance between angulations comprising 84 per cent of the maximum
diameter. The whorls expand quite slowly, so that there is a distinct tapering
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the whorls towards the apex in both dorsal and ventral views. The mould
bears low, faint, irregular folds and interspaces, which are markedly prorsiradiate
on the flanks and sweep across the venter with a broad and shallow convex
curvature. Two of these folds are strengthened into broad collars, one at the
beginning of the body chamber, the other at a diameter of just over 20 mm.
These are succeeded by distinct if shallow constrictions.
On the venter, patches of partially exfoliated shell appear smooth, save for
the low folds already mentioned; a few tiny areas preserve fine, dense lirae,
invisible to the naked eye. In contrast, areas of umbilical wall where shell is
well preserved show distinct if fine prorsiradiate lirae, concave close to the
umbilical seam, but weakly flexed so as to be gently convex across the angulation.
So far as can be discerned the whole of the shell was finely lirate, lirae paralleling
the folds and constrictions visible on the mould.
The external suture is well exposed (Fig. 4) and 1s deeply and intricately
subdivided, as is characteristic of other species of the genus. The median saddle
of the external lobe is prolonged into a series of narrow, tapering processes
which strongly recall the siphonal band of certain tetragonitids (Kennedy &
Klinger 1977).
Fig. 3. Jauberticeras collignoni Murphy. Holotype, M. Collignon Collection no. 435, from the
Albian sensu lato of Betioky, Madagascar. A-D x 2; E-H ~x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 7
Fig. 4. Sutures of Jauberticeras collignoni Murphy. Based on specimen D293 (Van Hoepen
Collection) from the Mzinene Formation of the Mzinene River, Zululand, and of late middle
or early late Albian age. A « 12: B < 6 to show details of siphonal structures.
3) ANNALS OF THE SOUTH AFRICAN MUSEUM
E F G H
Fig. 5. Jauberticeras collignoni Murphy. D293 (Van Hoepen Collection) from the Mzinene
Formation of the Mzinene River, Zululand, and of late middle or early late Albian age.
A-Dix 22. E=H << Ii,
CRETACEOUS FAUNAS FROM SOUTH AFRICA 9
Discussion
Murphy based this species on the specimen figured by Collignon (1963,
pl. 251 (fig. 1080)) which he refigured in dorsal view only (Murphy 1967, pl. 64
(fig. 16), text-fig. 1 (9)). The original description is short, and in it Murphy
characterized the species by its low expansion rate, smooth shell, and strong
second saddle in the internal suture, lack of lirae and constrictions and similar
whorl section to that of the type species. As can be seen from the holotype, here
reillustrated (Fig. 3), the specimen does not quite fit Murphy’s description in
possessing constrictions. There are at least three distinct constrictions on the
outer whorl, markedly prorsiradiate on the flank, and feebly convex across the
venter, each being preceded by a slight swelling. The whorl section, with a
breadth to height ratio of 2,54 at the maximum preserved diameter and 2,08
360° before this, comprises only 68 per cent of the total diameter whereas in
J. jaubertianum, to which it was said to be similar, the figure is 71 to 94 per cent.
The whorl angulation is also significantly blunter. The latter, however, may be
partially due to preservation or even of intra-specific value only, being dependent
on ontogeny (see Murphy 1967, pl. 64 (fig. 23)).
The second internal saddle is, however, prominently developed, whilst the
shell surface is smooth, even under a lens. Close inspection shows that this may
be no more than an artefact of preservation; the originally aragonitic shell has
been replaced by iron-stained sparry calcite, and the whole of the surface of the
umbilicus of the specimen is corroded and pitted. The venter is similarly pitted,
and in places the angulation is covered and accentuated by traces of a succeeding
whorl, now lost. This preservation and the remaining adherent matrix of fine
sandstone suggest the specimen to be from a precisely similar setting to many
of the Albian and Cenomanian ammonites from Zululand which the authors
have studied, especially those found weathering out along the terraces of the
Mzinene and on the Skoenberg, where identically preserved specimens of diverse
ammonites have lost delicate growth lines and lirae by post-mortem processes.
Lack of liration is not, therefore, a diagnostic feature of the species in the
authors’ view. With these qualifications in mind, the Zululand specimen, although
larger than the Madagascan type, can be referred to Jauberticeras collignoni on
the basis of similarities of whorl section, bluntness of angulation, relative propor-
tions (which are similar but not identical), expansion rate, form and direction of
constrictions, and associated collar-like swellings. Because the present specimen
does not show the septal face (retaining as it does part of the body chamber)
the nature of the internal suture is not determinable.
The principal difference between J. collignoni and the type species, J. jauber-
tianum (of which J. latecarinatum (Anthula) (1899: 101, pl. 7 (fig. 2a—c)) is
probably a synonym according to Murphy (1967: 605-6)), lies in the more
prominent development of the second saddle in the internal suture.
Jauberticeras villoutreysi Murphy (1967: 606, pl. 64 (figs 25—29)) (the holo-
type BMNH C68148 is reproduced here as Fig. 6A—G) has much coarser lirae
on the umbilical wall, which give a distinctive appearance to the whorl angulation
10 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. Jauberticeras villoutreysi Murphy. Holotype, BMNH C68148, from the Albian of Clar,
Alpes-Maritimes, France (Astier Collection). A-C x 2; D-G x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA II
(Fig. 6A). In addition, the expansion rate is such that the last half whorl is
almost parallel-sided in profile (Fig. 6B, F), the venter flatter, and constrictions
straight across the venter with a broad, band-like collar associated. In addition
the saddles in the internal suture are of different sizes (Fig. 6C).
Jauberticeras lanternoi Wiedmann (1962: 32, pl. 2 (fig. 2)) has quite different
proportions and whorl section, and may possibly be a juvenile Gabbioceras
(M. A. Murphy pers. comm. 7.3.1977).
J. subbeticum Wiedmann (1962: 31, pl. 2 (fig. 3a—c)} has much coarser
umbilical liration and a different form of whorl section and constrictions, and
again is of dubious generic affinity. According to Murphy (pers. comm. 7.3.1977)
it is probably a Scaphites. |
Occurrence
The holotype of Jauberticeras collignoni is from the Albian sensu lato of
Collignon’s locality 435, ‘Pied Est du Vohimaranitra (Betioky), Madagascar’.
The Zululand specimen is from the Albian part of the Mzinene Formation along
the Mzinene River near Hluhluwe. Precise locality data are lacking, but preserva-
tion suggests it to be from locality 51 of Kennedy & Klinger (1974: 288, fig. 4)
where concretions are exposed in the stream bed and bank extending around
the broad meander of the river east-north-east of the Sisal Factory at 27°53'43’S,
32°19'22”E. The specimen is thus of late middle or early late Albian age (Albian
IV-V).
ACKNOWLEDGEMENTS
Our best thanks are due to General M. Collignon of Moirans (Jsere), who
kindly allowed us to study the holotype of Jauberticeras collignoni, and has
helped us throughout our work on the Zululand Cretaceous. We are also
grateful to Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural
History) for allowing us to study the specimens in their care, and Dr J. Sornay
of the Muséum Nationale d’Histoire Naturelle, Paris, for help in our study of
Specimens in the D’Orbigny collection.
REFERENCES
ANTHULA, D. J. 1899. Ueber die Kreide Fossilien des Kaukasus. Beitr. Paldont. Geol. Ost. Ung.
12: 54-159.
COLLIGNON, M. 1963. Atlas des Fossiles caractéristiques de Madagascar (Ammonites), Fascicule
X (Albien). Tananarive: Service Géologique.
Howarth, M. K. 1974. The Lower Cretaceous ammonite genera proposed by C. Jacob in
1907. Palaeontology 17: 727-728.
KENNEDY, W. J. & KLINGER, H. C. 1974. Cretaceous faunas from Zululand and Natal, South
Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.
KENNEDY, W. J. & KLINGER, H. C. 1977. Cretaceous faunas from Zululand and Natal, South
Africa. The ammonite family Tetragonitidae Hyatt, 1900. Ann. S. Afr. Mus. 73: 149-197.
KENNEDY, W. J. & KLINGER, H. C. In press. Cretaceous faunas from Zululand and Natal,
South Africa. The ammonite family Gaudryceratidae Spath, 1927. Bull. Br. Mus. nat. Hist.
(Geol.).
1 ANNALS OF THE SOUTH AFRICAN MUSEUM
Murpny, M. A. 1967. The Ammonoid subfamily Gabbioceratinae Breistroffer. J. Paleont. 41:
595-607.
OrBIGNY, A. D’. 1850. Note sur quelques nouvelles espéces remarquables des étages Neocomien
et Aptien de France. J. Conchyliol. 1: 196-201.
WIEDMANN, J. 1962. Die Gabbioceratinae Breistroffer. Newes Jb. Geol. Paldont. Abh. 115: \-43.
6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to n the text
e.g. “.. . the Figure depicting C. namacolus...’; ‘. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
A JAUBERTICERAS FROM THE MZINENE
FORMATION (ALBIAN)
\ as WKS 2 DECEMBER 1977 ISSN 0303-2515
; MLSs,
NOV 1 9 ygQQn
Ft A. ine 47 fovin, at me)
ANNALS
OF THE SOUTH AFRICAN
UM
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INSTRUCTIONS TO AUTHORS
1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:
(a) Centred masthead to consist of
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(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
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(f) Summary, if paper is lengthy
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3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of
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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...’
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‘As described (Haughton & Broom 1927)...’
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Note: no comma separating name and year
Dagination indicated by colon, not p.
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et al. in text for more than two joint authors, but names of all authors given in list of references.
(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Carus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4 ): 51.
THIELE, J. 1910. Mollusca: B. Boleeceehees? Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band |
December 1977 Desember
Part 2 Deel
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 5
CRUSTACEA, DECAPODA, REPTANTIA
AND NATANTIA
By
BRIAN KENSLEY
Cape Town Kaapstad
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are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town
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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 5
CRUSTACEA, DECAPODA, REPTANTIA AND NATANTIA
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 16 figures)
LMS. accepted 10 May 1977]
ABSTRACT
Sixty-one species of Reptantian and Natantian decapod crustaceans from deep water off
Natal are listed. Of the thirteen new records of Natantia dealt with from the area, three are
described as new species. These are Benthesicymus expansus, Hymenopenaeus kannemeyeri and
Pasiphaea meiringnaudei.
CONTENTS
PAGE
Introduction : ‘ i hk
Species List : : : ene AS
Systematic Discussion . : 5 4N)
Acknowledgements ck : SiS
References . : : : « |) 43
INTRODUCTION
The present paper forms part of a series based on material collected by the
South African Museum during two cruises off the Natal coast, on the R/V Meiring
Naude (see Fig. 1). For the scope of, and the background to, this programme,
as well as for all station data, the reader is referred to Louw (1977).
All the species of Reptantia and Natantia collected are listed systematically,
but only new species, new records for South Africa, and a few additional species
requiring further comment are dealt with in the systematic text.
The following abbreviations are used throughout this paper: SAM—South
African Museum catalogue number; SM—Meiring Naude station numbers;
CB—carapace breadth; CL—carapace length; RL—rostral length; TL—total
length; ovig.— ovigerous; juv.—juvenile.
SPECIES LIST
Where specimens are not identified to species level, these were either immature or damaged.
‘ * New record for South African waters.
SM Station Ovig.
no. 33 29 29 Juv.
PALINURA
Family Eryonidae
Stereomastis sculpta (Smith) . A i : 58 — ~- 1 —
(Barnard 1950: 572) 83 1 — — —
(Eryoneicus caecus) ; 5 F ee LOT — — _ 1
13)
Ann. S. Afr. Mus. 74 (2), 1977: 13-44, 16 figs.
14
ANNALS OF THE SOUTH AFRICAN MUSEUM
SOG Ta
PAP INGA
ho72
50'
A DURBAN
Fig. 1. Map showing localities of collecting stations.
50°
28°
29
SI9453 Sd os
%
a) 5
oY)
@ ve
j 33° 10 20°
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
SM Station
no. 3d eee
Family Palinuridae
Palinurus delagoae Barnard . : : : 22 I =
(Berry & Plante 1973: 374)
Projasus parkeri (Stebbing) . , ' : 38 — ox
(George & Grindley 1964: 87)
ASTACURA
Family Astacidae
Nephropsis atlantica Norman. : 38 3 D
(Barnard 1950: 530) 66 3 —
NATANTIA PENAEIDEA
Family Penaeidae
Subfamily Sicyoninae
Sicyonia longicauda Rathbun . ‘ : 86 — , 1
(Barnard 1950: 635)
Subfamily Aristeinae
Aristaeomorpha foliacea (Risso). , ; fi 1 y)
(Barnard 1950: 625) 22 2 4
28 1 -
38 3 3
44 — 2
66 — 1
83 1 _
92 — 2
99 1 _
100 3 3
* Aristeus semidentatus (Bate) . : ; ee Os 1 _
(see systematic section, p. 20)
*Aristeus virilis (Bate) . ; : : : 58 2 —
(see systematic section, p. 21) 91 1 —
107 ~- 1
*Benthesicymus expansus sp.nov. . : ee hO7 1 —
(see systematic section, p. 22)
*Benthesicymus investigatoris Alcock & Anderson 38 — |
(see systematic section, p. 26) 58 — 1
66 -- 1
107 _ y)
Gennadas bouvieri Kemp : : ; ; 5 = 1
(Kensley 19716: 273) 56 1 —
63 — 4
88 — i
105 — 1
111 -- 1
Gennadas capensis Calman. : ; : 99 — 1
(Kensley 19715: 277) 105 1 1
Gennadas clavicarpus De Man : : 5) 1 1
(Kensley 19715: 278) 63 — 1
88 — 1
96 1 1
105 — 1
111 3 —
Gennadas gilchristi Calman . : 5 : 25 — 3
(Kensley 19715: 280) 33 2 4
49 -- 2)
55 3 4
—
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
SM Station Ovig.
no.
= +O
|
|
Gennadas parvus Bate . : ; : F 25
(Kensley 19715: 287) ) 33
Gennadas scutatus Bouvier . : x 4 56
(Kensley 19715: 288) 71
Gennadas tinayrei Bouvier. : ; : 56
(Kensley 19715: 290) 63
|
|
Plesiopenaeus edwardsianus (Johnson) . ; 7
(Crosnier & Forest 1973: 292) 28
lswleeal
|
|
|
|
|
Plesiopenaeus ni‘idus Barnard ; ‘ , 7
(Barnard 1950: 622) 22
— me OY
BRENANIUNERE XA
|
|
ww
Subfamily Solenocerinae
Haliporoides triarthrus (Stebbing) : : 2)
(Barnard 1950: 619) 92
* Hymenopenaeus kannemeyeri sp.nov. . ; 38
(see systematic section, p. 27)
Hymenopenaeus sp. : dU 1 _ — =
== an
|
|
|
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
SM Station
no.
Subfamily Penaeinae
*Funchalia villosa (Bouvier) . ,
(see systematic section, p. 29)
Parapenaeopsis acclivirostris Alcock
(Barnard 1950: 604)
Penaeopsis rectacuta (Bate)
(Hall 1962: 18)
Penaeus indicus Edwards
(Barnard 1950: 588)
Family Sergestidae
Sergestes (Sergestes) armatus Kroyer
(Kensley 1971la: 232)
Sergestes (Sergestes) atlanticus Edwards .
(Kensley 1971la: 234)
Sergestes (Sergestes) curvatus Crosnier & Forest
(Crosnier & Forest 1973: 315)
Sergestes (Sergestes) orientalis Hansen
(Kensley 1971a: 238)
Sergestes (Sergestes) sargassi Ortmann
(Kensley 1971la: 241)
Sergestes (Sergestes) pectinatus Sund
(Kensley 1971a: 240)
lwme leo!
3d
Psi |
w
—_—
| Sy eal
op load ol
Ovig.
e2
juv.
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
SM Station ovig.
no.
+O
+O
+O
+O
a
c
<
Sergestes (Sergia) laminatus Burkenroad . : 56
(Kensley 1971la: 251) 88
\O
~
Ox
| RerRUmOBRWN OG
NwUNueas |
|
Sergestes (Sergia) potens Burkenroad . : U
(Kensley 1971la: 253)
Sergestes (Sergia) prehensilis Bate . : 5
(Kensley 1971a: 253)
Ne
00
—
e ||
(oe)
(oe)
| Ne WN) DB KH OW — RK Pe
Sergestes (Sergia) regalis Gordon . : F DS
(Kensley 1971a: 256) 55
ore)
SS
loos fool
Sergestes (Sergia) scintillans Burkenroad . : WS)
(Kensley 1971a: 257) 88
Sergestes (Sergia) splendens Sund . : : 88
(Kensley 1971a: 260) .. ‘ : : 96
* Sergestes (Sergia) talismani Barnard . ; 55)
(see systematic section, p. 31) 63
—
—
N
b—
|
|
—
N
|
NATANTIA CARIDEA
Family Oplophoridae
* Acanthephyra armata Edwards i ; 38 4
(see systematic section, p. 31) 58 3
Acanthephyra eximia Smith . : : 2 3 —
(Crosnier & Forest 1973: 34) 38 —
—=re UN = COR WD
|
|
* Acanthephyra indica Balss_ . 4 ; é 56
(see systematic section, p. 31) 88
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
Acanthephyra quadrispinosa Kemp .
(Barnard 1950: 668)
Oplophorus gracilirostris Edwards .
(Chace 1936: 30)
Oplophorus spinicauda Edwards
(Kensley 1969: 169)
Oplophorus spinosus (Brullé) .
(Crosnier & Forest 1973: 25)
Systellaspis debilis (Edwards)
(Crosnier & Forest 1973: 87)
Family Nematocarcinidae
Nematocarcinus sp.
Family Pasiphaeidae
* Eupasiphae gilesii Wood-Mason
(see systematic section, p. 32)
*Pasiphaea sivado (Risso) ,
(see systematic section, p. 32)
*Pasiphaea meiringnaudei sp. nov. .
(see systematic section, p. 34)
Family Hippolytidae
Hippolysmata tugelae Stebbing
(Barnard 1950: 712)
SM Station
no.
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hes hoes]
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CON = = =
—
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N
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
SM Station Ovig.
no.
Oy
+0
+0
+0
+O
et
|
<
Family Pandalidae
Heterocarpus dorsalis Bate. . ; 3 7
(see systematic section, p. 38) 28
Heterocarpus laevigatus Bate . ; : : 38
(see systematic section, p. 38) 66
co
WwW
Heterocarpus tricarinatus Alcock & Anderson . 22
(see systematic section, p. 38)
Parapandalus richardi (Coutiére) . : s 56
Plesionika martia (Edwards) . : , ' 7
(Crosnier & Forest 1973: 212) 22
ereel) cytes 4
Samm csp) es
S
|
|
Family Glyphocrangonidae
Glyphocrangon dentatus Barnard . ' ; 22
(see systematic section, p. 40) 28
ieee
| | @&
| ld
*Glyphocrangon regalis Bate . : 3 : 7
(see systematic section, p. 40) 28
| sree |
|
|
|
oneeenl
|
Ne)
Ne)
|
Family Crangonidae
Pontophilus sculptus (Bell) . ; . ° 86 _
(Barnard 1950: 810)
—
|
|
SYSTEMATIC DISCUSSION
(new species, new records, and selected other species)
PENAEIDEA
Family Penaeidae
Aristeus semidentatus (Bate)
Figs 2A, 3A
Hemipenaeus semidentatus Bate, 1888: 305, pl. 49 (fig. 1).
Aristaeus semidentatus: Wood-Mason & Alcock, 1891: 280. Alcock, 1901: 31. Alcock &
McArdle, 1901: pl. 49 (fig. 3).
Aristeus semidentatus: De Man, 1911: 29.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 21
Previous records
Laccadive Islands; Cape Comorin; Bay of Bengal; off Banda; Kei Islands;
Kermadec Islands.
Material
SAM-A15139 SM 107 12 gg CL + RL 26,8 mm x TL 77,0 mm to
CL + RL 31,5 mm x 92,0 mm
Remarks
The genus Aristeus is characterized by the lack of an hepatic spine, an
indistinct cervical groove, the third pereiopods possessing an epipodite but no
podobranch, the fourth pereiopods lacking an epipodite, the exopodite of the
second maxilliped longer and stronger than the exopodite of the third maxilliped.
There is some variation in the rostral dentition of the present material,
several specimens showing a small dorsal and ventral tooth close to the apex
(Fig. 2A). (Burucovsky & Romensky 1972, also describe rostral variability in
Aristeus varidens.)
Aristeus virilis (Bate)
Figs 2B, 3B
Hemipenaeus virilis Bate, 1888: 303, pl. 44 (fig. 4).
Hemipenaeus tomentosus Bate, 1888: 307, pl. 49 (figs 2-3).
Aristaeus virilis: Wood-Mason & Alcock, 1891: 279. Alcock, 1901: 30:
Aristeus virilis:; De Man, 1911: 27.
Fig. 2. A. Aristeus semidentatus. B. Aristeus virilis.
DY ANNALS OF THE SOUTH AFRICAN MUSEUM
Previous records
Andaman Sea; China Sea; Philippines; Bali Sea; New Hebrides.
Material
SAM-A15159 SM 58 2 3g CL 41,2 mm x TL 110,5 mm; 37,0 mm x
+ 95 mm (telson damaged)
SAM-A1I5160 SM 91 1 ¢43,0 mm x 109,0 mm
SAM-A1I5161 SM 107 1 ° (rostrum damaged)
Remarks
The present material shows the characteristic thickening of the distal part
of the scaphocerite, but the males lack the pubescence of the integument. A twist
in the base of the inner antennular flagellum is also present. The two species of
Aristeus dealt with here may be distinguished by the differences in the petasmata
(Fig. 3) and rostra (Fig. 2), and the pubescent carapace of some specimens of
A. virilis.
Benthesicymus expansus sp. nov.
Figs 4—5
Description
Male
Carapace (Fig. 4A) integument soft, membranous. Rostrum an entire crest,
apex missing in holotype, ventral border bearing slight ridge carrying setae. No
A
Fig. 3. A. Aristeus semidentatus petasma. B. Aristeus virilis petasma.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 23
antennal spine; orbitoantennal groove extended posteroventrally to branchial
region; rounded ridge situated above orbitoantennal groove; branchiostegal
spine small, supported by short carina, no hepatic spine; cervical groove strong,
almost reaching orbitoantennal groove; suprabranchial ridge fairly strong,
rounded. Thoracic sternite between fourth pereiopods bearing short narrowly
rounded, forwardly directed lobe. Posterior half of third pleon segment, and
pleon segments four to six with mediodorsal carina, none bearing posterior spine.
Telson carrying four pairs of lateral spines on distal third, apex acute (Fig. 4B).
Antennal scaphocerite (Fig. 5F) blade two and one-third times longer than
wide, outer margin reinforced by tapering ridge, ending in small spine some
distance from apex of scale; basal segment carrying small dorsal hook-like
process.
Antennular peduncle segments setose.
Ocular tubercle present on inner margin of proximal half of eye-stalk.
Mandibular palp two-segmented, basal segment one and a half times longer
and considerably broader than distal segment; mandible consisting of strong,
straight cutting edge and three small teeth on inner face (Fig. 5A).
First maxilliped (Fig. 5B) exopod narrowing abruptly to segmented distal
portion. Second maxilliped (Fig. 5E) with merus very expanded, distally rounded,
inner margin setose.
Third maxilliped longer than first perelopod (Fig. 4A), merus and ischium
expanded, fringed with setae on ventral margin; propodus and carpus relatively
slender, of equal length, dactylus broadest at midpoint, tapering distally. First
pereiopod chela relatively slender, finger and thumb slightly longer than palm;
carpus slender; merus and ischium expanded.
Second pereiopod (Fig. 4A) longer than first, very slender, finger and thumb
of chela equal in length to palm; carpus and merus of equal length.
Third to fifth pereiopods missing in holotype.
Petasma slender (Fig. 4C), divided distally into median narrow lobe con-
nected to truncate, rounded portion, and outer smoothly convex lobe bearing
spinules.
Inner scale of appendix masculina narrow, elongate, apically rounded,
bearing short setae; outer scale broad, longer than inner.
Material
Holotype SAM-A15140 SM 107 1 ¢ CL 50,5 mm TL 150 mm
Remarks
The lack of mediodorsal pleonal spines, the ocular tubercle being proximal
in position, the lack of an hepatic spine, and the presence of a mediodorsal keel
on pleonal segments 3-6, place this specimen close to B. iridescens Bate (see
Burkenroad 1936: 45; Roberts & Pequegnat 1970: 42). The latter authors note
the difficulty of sorting out the ‘Brasiliensis’ group of species of Benthesicymus,
due mainly to a lack of information on intraspecific variation. In spite of this
24 ANNALS OF THE SOUTH AFRICAN MUSEUM
Z Yi YY YY A
Vy
Fig. 4. Benthesicymus expansus sp. nov. A. Holotype, carapace in lateral view. B. Telson.
C. Anterior and posterior views of petasma.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 25
limitation, the present new species is described, albeit from a single specimen.
The species has two features which easily separate it from B. iridescens. These
are the non-dentate rostrum, and the expanded meri and ischia of the third
maxilliped and first pereiopods. (The specific name is derived from this latter
feature.) In addition, differences from B. iridescens are seen in the petasma, the
appendix masculina, the telson, the lack of an antennal spine, and in the carapace
sculpture.
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Fig. 5. Benthesicymus expansus sp. nov. A. Mandible. B. First maxilla. C. Second maxilla.
D. First maxilliped. E. Second maxilliped. F. Antennal scaphocerite.
26 ANNALS OF THE SOUTH AFRICAN MUSEUM
Benthesicymus investigatoris Alcock & Anderson
Fig. 6
Benthesicymus investigatoris Alcock & Anderson, 1899: 282. Alcock, 1899: pl. 41 (fig. 2);
1901: 44. Burkenroad, 1936: 25.
Previous records
Saya de Malha (mid Indian Ocean); Andaman Sea; Gulf of Manar;
Halmahera Sea; Ceram Sea; Hawaii.
Material
5 292 CL + RL 24,0 mm x TL 76 to 31,0 mm x 89,5 mm
Fig. 6. Benthesicymus investigatoris. A. Carapace in lateral view. B. Thelycum.
THE SOUTH AFRICAN MUSEUM’S ME/RING NAUDE CRUISES pe
Hymenopenaeus kannemeyeri sp. nov.
Fig. 7
Description
Carapace (Fig. 7A) firm, glabrous. Rostrum about one- Ania length of
carapace directed obliquely upward at angle of about 15°, with eight dorsal
teeth (three postorbital) in uninterrupted series, apical tooth very small, ventral
margin bearing fringe of setae. Carapace with strong antennal and postantennal
spine, each supported by stout carina; hepatic spine lacking keel; branchiostegal
spine marginal, supported by keel extending posteriorly to hind margin of cara-
pace; cervical sulcus well developed, with posterior margin of groove at midpoint
marked by a short keel, joining the hepatic groove ventrally. Hepatic groove
marked posteroventrally by short keel. Branchiocardiac sulcus well developed
with supra-branchial portion having a straight, rounded ridge ventrally. First
pleon segment dorsally rounded, segment two with posterior half keeled, third
to sixth segments fully keeled, segments four to six with short posterior spine.
Telson (Fig. 7D) with two fixed spines subterminally, with strong dorsolateral keel
extending from spines to proximal margin; outer uropodal ramus just reaching
telsonic apex.
Eye reaching slightly beyond midpoint of rostrum, tubercle at midpoint of
medial eye-stalk margin (Fig. 7E).
Prosartema of antennule short and rounded, carrying long setae; basal
peduncular segment with strong distal spine; stylocerite a strong spine; terminal
peduncular segment with lateral setose ridge and thick setose patch dorsally;
flagellum circular in cross-section.
Scaphocerite extending well beyond antennular peduncle.
Mandibular palp conspicuous, setose; distal segment slender, tapering,
shorter than proximal segment, latter broadly oval.
Third maxilliped extending slightly beyond scaphocerite.
First pereiopod with strong distoventral spine on ischium and basis.
Rudimentary exopods on first to fifth pereiopods and third maxilliped;
epipods on second and third maxillipeds and first to fourth pereiopods.
Inner margin of fifth perelopod basis in male with anteriorly directed
triangular lobe above genital aperture.
Thelycum of female (Fig. 7C) relatively simple, sternites of fourth and fifth
pereiopods with rounded, convex central areas.
Petasma divided distally into two lobes; inner lobe longer than outer,
truncate, with distal hollowed portion produced laterally into triangular point;
outer lobe also truncate, with posterior strengthening ridge. Appendix masculina
of second pleopod (Fig. 7F) about half length of appendix interna; both bearing
marginal spinules; leaf-like expansion present on outer side of base of endopod.
Material
Holotype SAM-A15141 SM 38 3 CL + RL 43,5 mm TL 119,5 mm
Allotype SAM-A15141 SM 38 9 CL + RL 39,8 mm TL 110,8 mm.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
The present specimens with the rostral and post-rostral teeth not separated,
with a strong branchiostegal spine, and lacking a pterygostomian spine, fall into
Burkenroad’s (1936) Group II.
This group includes H. robustus, H. modestus, H. lucassi and H. muelleri.
H. kannemeyeri differs from all these in the structure of the petasma and
thelycum. The latter most closely resembles that of H. fattahi Ramadan, 1938
(as figured by Crosnier & Forest 1973, fig. 85g—h) but the petasma of H. fattahi
differs markedly from the present species.
Fig. 7. Hymenopenaeus kannemeyeri sp. nov. A. Carapace in lateral view. B. Anterior and
posterior view of petasma. C. Thelycum. D. Telson. E. Eye. F. Pleopod 2, 3, appendix
interna and appendix masculina.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 29
The species is named for S. X. Kannemeyer, of the Department of Marine
Biology, South African Museum, in appreciation for his invaluable help in the
Meiring Naude cruises.
Genus Funchalia Johnson
As Funchalia villosa and F. woodwardi have been confused in the past, the
petasma and thelycum of each is figured, and all the South African Museum’s
holdings of both species are listed.
Funchalia (Funchalia) villosa (Bouvier)
Fig. 8C-D
Funchalia (Funchalia) villosa: Burkenroad, 1936: 129. Crosnier & Forest, 1973: 296.
Previous records
Eastern and western North Atlantic; Caribbean; southern central South
Atlantic.
Material and South African records
SAM-A15368 SM 18 12 CL 22,8 mm
SAM-A15369 SM 33 12 CL 21,0 mm
SAM-A15370 SM 88 13192CL 13,0 mm, 14,2 mm
SAM-A15371 SM 111 13 CL 18,4 mm
SAM-A13195_ off Natal, 30°30’S 31°45’E 200 m 1 3
SAM-A13197_ off Natal, 26°30’S 42°40’E 500 m 1 d
SAM-A13198 off Agulhas Bank, 37°10’S 21°50’E 500 m1 4d
SAM-A13236 off Natal, 31°44’S 44°35’E 500 m 3 gd
Funchalia (Funchalia) woodwardi Johnson
Fig. 8A-B
Funchalia (Funchalia) woodwardi: Barnard, 1950: 609, fig. 112. Gordon & Ingle, 1956: 478
Paulinose, 1974: 433.
Previous records
Eastern North Atlantic; Mediterranean; south-eastern South Atlantic.
South African records
SAM-A8481_ off Table Bay, from stomach of stock fish 9 ¢3
SAM-A10914 off Cape Peninsula, from tunny stomach several dg & 29
SAM-A10995_ off Cape Peninsula, from tunny stomach |
SAM-A11025 off Dassen Island, from tunny stomach 25 g¢ 19 9°
SAM-A11027 off Cape Peninsula, several g¢ & 99
SAM-A12014 off Dassen Island, 500 m 2 ¢¢ 2 99
SAM-A12482 Vema Seamount, from tunny stomach 60 immature 3g & 2°
SAM-A13196 off Cape Peninsula, 250 m 3 99
SAM-A13237 off Cape Peninsula, 350 m7 gg 7 22
SAM-A13238 off Cape Peninsula, 500 m 2 g¢ 3 99
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
B
Fig. 8. A. Funchalia woodwardi petasma. B. F. woodwardi thelycum. C. Funchalia villosa
petasma. D. F. villosa thelycum.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Sil
Family Sergestidae
Sergestes (Sergia) talismani Barnard
Sergestes talismani Barnard, 1947: 384.
Sergestes (Sergia) talismani:; Crosnier & Forest, 1973: 325, figs 11la—c, 112a—b.
Previous records
Cape Verde Islands; Portuguese Guinea; Gabon; Congo.
Material
See species list.
Remarks
This would seem to be the first record of the species from the Indian Ocean.
CARIDEA
Family Oplophoridae
Acanthephyra armata Edwards
Fig. 9B
Acanthephyra armata: Bate, 1888: 744, pl. 125 (fig. 2). De Man, 1920: 61, pl. 6 (fig. 13).
Chace, 1936: 27.
Previous records
West Indies; Bali Sea; Straits of Macassar; off Kei Islands; off Banda
Island.
Material
SAM-A15154 SM 38 434499
SAM-A15150 SM 58 3 3¢3 99
gd CL 16,9 mm-29,0 mm RL 27,0 mm—37,5 mm TL 92,0 mm—137 mm
oe) 24,5 mm-—33,0 mm 34.7 mm-—35,2 mm 121 mm—-146 mm
Acanthephyra indica Balss
Fig. 9A
Acanthephyra indica Balss, 1925: 264, fig. 34. Chace, 1936: 26.
Acanthephyra sp. De Man, 1920: 68, pl. 6 (fig. 16).
Previous records
Between Sumatra and Ceylon; Banda Sea; off Ambon.
Material
See species list.
3 gg 123 ovig. 22 gG¢CL10,0 mm-10,.2mm 2 CL 10,9 mm-13,1 mm
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 9. A. Acanthephyra indica. B. Acanthephyra armata.
Family Pasiphaeidae
Eupasiphae gilesii (Wood-Mason & Alcock)
Fig. 10B
Parapasiphae (Eupasiphae) gilesii Wood-Mason & Alcock, 1893: 166. Alcock, 1892: pl. 3 (fig. 8);
1901: 66.
Eupasiphae gilesii: Holthuis, 1955: 36. Tirmizi, 1969: 214, figs 1-4. Foxton, 1970: 958.
Eupasiphae gilesi: Fisher & Goldie, 1961: 78. Crosnier & Forest, 1973: 150, fig. 44.
Previous records
Bermuda; Cape Verde Islands; Canary Islands; Madeira; Arabian Sea;
Gulf of Oman; Andaman Sea.
Material
SAM-A15151 SM 105 1g CL + RL 32,4 mm TL 97 mm
Pasiphaea sivado (Risso)
Fig. 10A
Pasiphae sivado: Wood-Mason & Alcock, 1893: 161. Alcock, 1892: pl. 3 (fig. 6); 1901: 59.
Sivertsen & Holthuis, 1956: 29. Crosnier & Forest, 1973: 133.
33
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
Fig. 10. A. Pasiphae sivado. B. Eupasiphae gilesi,
34 ANNALS OF THE SOUTH AFRICAN MUSEUM
Previous records
Eastern North Atlantic; Mediterranean; Red Sea; Bay of Bengal; Andaman
Sea.
Material
SAM-15153 SM 63 2 99
SAM-A15152 SM 85 12°
2° CL 10,0 mm-14,5 mm
Pasiphaea meiringnaudei sp. nov.
Figs 11-14
Description
Carapace (Fig. 11) mid-dorsally carinate for almost its entire length.
Rostrum a postfrontal triangular spine directed obliquely upward, only rarely
reaching beyond anterior carapace margin. Anterior margin of rostrum straight,
almost vertical, apparently more variable in females than in males (Fig. 13).
Orbital angle a rounded lobe; branchiostegal spine directed outwards; supra-
branchial ridge rounded, not reaching posterior carapace margin.
Second to sixth abdominal segments dorsally keeled, not ending in a
posterior spine; sixth segment with curved lateral ridge. Telson shorter than
Fig. 11. Pasiphae meiringnaudei sp. nov. Holotype in lateral view,
THE SOUTH AFRICAN MUSEUM’S MEIJRING NAUDE CRUISES 35
uropodal endopod, dorsally grooved, distally forked, with seven spines on inner
margin of sinus (Fig. 121).
Antennular peduncle extending for two-thirds of scaphocerite (Fig. 11);
stylocerite (Fig. 12G) broadening distally, twisted, apically acute, dorsolateral
flagellum with about eighteen enlarged basal segments in male and female.
| Antennal base bearing ventrodistal spine.
Fig. 12. Pasiphae meiringnaudei sp. nov. A, Anterior carapace and rostrum. B. Mandible.
C. First maxilla. D. First maxilliped. E. Second maxilliped. F. Second maxilla. G. Antennular
base. H. Antennal base. I. Telsonic apex. J. Pleopod 2, ¢, appendix interna and appendix
masculina. K. Pleopod 1, 3. L. Pleopod 1, 3, endopod.
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
Third maxilliped just reaching to end of scaphocerite.
First perelopod overreaching scaphocerite by half length of palm plus
fingers (Fig. 11); palm longer than fingers; merus with one to four ventral spines;
basis with ventral distal triangular spine.
Second pereiopod overreaching scaphocerite by half length of palm plus
fingers; palm equal in length to fingers; carpus with broad ventral tooth; merus
with fifteen to twenty-one ventral spines; ischium with one spine, basis with four
to seven spines (Fig. 14B).
Third pereiopod overreaching anterior carapace margin by about half
length of propodus plus dactylus; very slender; dactylus tapering; propodus one-
third length of dactylus; carpus very elongate, almost four times length of merus.
Fourth pereiopod reaching to midpoint of carpus of third pereiopod.
Fifth pereiopod longer than fourth pereiopod, reaching to end of carpus of
third pereiopod.
Endopod of first pleopod ¢ short, with triangular process on inner margin
bearing very small denticles (Fig. 12L).
Second pleopod 3 appendix masculina twice length of appendix interna,
latter curved, with seven marginal setae (Fig. 12J).
Uropodal endopod two-thirds length of exopod, latter with small terminal
spine on outer margin.
Material |
Holotype SAM-A15316 SM 107 ovig. 9 CL 57,0 mm TL 166 mm
SM 38 134 CL 39,9 mm TL 126 mm
SM 58 4 493 CL 35,5 mm-36,3 mm TL 109 mm—-110 mm
SM 107 5 ovig. 992 50,0 mm—58,0 mm 153 mm—-168 mm
SM 91 192 42,0 mm 125 mm
SM 28 12 34,5 mm 108 mm
SM 40 2 99 31,8 mm—42,4 mm 95-130 mm
Fig. 13. Pasiphae meiringnaudei sp. nov. rostral variation.
Top row, ovigerous 22. Bottom row, gd. Bottom right,
rostrum modified by ellobiopsid parasites.
EE
a
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 3)
Remarks
J. C. Yaldwyn (National Museum of New Zealand) has kindly supplied
most of the following information on the closely related species of Pasiphaea,
from an as yet unpublished review of the genus by him and F. A. Chace Jr
(Smithsonian Institution). The combination of three characters, viz. the distally
forked telson, carinate carapace and abdomen, and basis of the second pereiopod
ventrally spined, places the present species in a group with four previously
described species. These are P. multidentata Esmark, P.: tarda Kroyer, P. rath-
bunae Stebbing, and P. barnardi Yaldwyn. The rostral shape of P. meiringnaudei
differs from these four, but comes closest to P. barnardi. The finger-to-palm
ratio of the first and second pereiopods, however, differs from Yaldwyn’s species
where the fingers of the first pereiopod are subequal to the palm and the fingers
of the second pereiopod are distinctly longer than the palm. P. meiringnaudei
overlaps with P. tarda (as P. princeps in Kemp 1910, pl. 4 (figs 1—-7)) in the spine
counts of the first and second pereiopods, but differs markedly in the rostral
shape and in the relative lengths of the thoracic appendages.
The species is named for the R/V Meiring Naude.
Fig. 14. Pasiphae meiringnaudei sp. nov. A. First pereiopod, with chela further enlarged.
B. Second pereiopod, with chela further enlarged.
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Pandalidae
Heterocarpus dorsalis Bate
Fig. ISA
Heterocarpus dorsalis: De Man, 1920: 156, 171, pl. 15 (fig. 43). Barnard, 1950: 684, fig. 127a
Previous records
Off Durban; off East Africa; East Indies; Japan.
Material
See species list.
7 33 CL 20 mm—41 mm RL 35 mm-48 mm TL 98 mm-179 mm
IQD 27 mm—42 mm 35 mm-44 mm~—~—s«126 mm-—171 mm
Remarks
The ovigerous female of total length 171 mm is at least 20 mm longer than
the largest ovigerous female recorded by De Man (1920).
Heterocarpus laevigatus Bate
Fig. 15B
Heterocarpus laevigatus Bate, 1888: 636, pl. 112 (fig. 3). Stebbing, 1914: 40. De Man, 1920:
154, 159, pl. 13 (fig. 37). Barnard, 1950: 684, fig. 127b. Crosnier & Forest, 1973: 195, fig.
6lc. ;
Previous records
Cape Verde Islands; Madeira; off East London; Arabian Sea; East Indies;
Hawaii.
Material
See species list.
233 CL 65 mm-74 mm RL 28 mm-38 mm TL 121 mm—160 mm
39° 64 mm—-105 mm 36 mm—5!1 mm 121 mm—200 mm
Remarks
The East London record was of an immature specimen measuring approxi-
mately 63 mm TL.
Heterocarpus tricarinatus Alcock & Anderson
Fig. 15C
Heterocarpus tricarinatus: De Man, 1920: 155, 161, pl. 13 (fig. 38), pl. 14 (fig. 38). Calman,
1939: 204. Barnard, 1950: 682, fig. 127c—d.
Previous records
Off East London; Arabian Sea; East Indies.
Material
SAM-A15264 SM 22 | 3 CL 35mm RL 30 mm TL 135 mm
ovig. 9 37 mm 32 mm 146 mm
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 39
a Wal
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Fig. 15. A. Heterocarpus dorsalis. B. Heterocarpus laevigatus. C. Heterocarpus tricarinatus.
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
Using both Barnard’s (1950) and De Man’s (1920) keys to the genus
Heterocarpus, H. tricarinatus is arrived at. The genus is obviously variable with
regard to the rostral dentition, proportions of the pereiopodal dactyli, and the
lengths of the carapace keels. This led Calman (1939) to doubt the validity of
this species, suggesting that it might be synonymous with H. gibbosus Bate. The
variability of this latter species is well illustrated by De Man (1920). The present
material could well be placed in Bate’s species, but until this species complex
is resolved, the lower carapace keel reaching back about two-thirds of the cara-
pace length necessitates placing these two specimens in H. tricarinatus.
Family Glyphocrangonidae
Glyphocrangon dentatus Barnard
Glyphocrangon gilesii var. dentata Barnard, 1926: 128.
Glyphocrangon mabihissae Calman, 1939: 217, fig. 8.
Glyphocrangon dentarus Barnard, 1947 (typ. err.): 387.
Glyphocrangon dentatus Barnard, 1950: 722, fig. 134h.
Previous records
Off Mozambique, 540 m; off Zanzibar, 640-658 m.
Material
8 dd 1 2 8 ovig. 29, see species list.
So CL 16,5 mm-19,8 mm RL 17,8 mm-22 mm TL 73 mm-85 mm
ovig. 2° 19,1 mm—23 mm 20,8 mm-27,5 mm 83 mm—-102 mm
Remarks
Only slight differences are apparent between Calman’s specimens from
Zanzibar and the present material. These differences are to some extent probably
due to individual variation, as well as to variations between populations. The
spine on the midpoint of the posterior antennal carina is present in some of the
present specimens, absent in others, while the row of small tubercles just lateral
to the submedian carina seen in the Zanzibar specimens is represented by one
or two almost indistinct tubercles. Although the rostral length always exceeds
the carapace length, the ratio is variable.
Glyphocrangon regalis Bate
Fig. 16
Glyphocrangon regalis Bate, 1888: 517, pl. 93 (figs 3-4). De Man, 1920: 220.
Description
Rostrum curved ventrally, with apex upturned, slightly less than two-thirds
carapace length, with pair of lateral rostral teeth above eyes, second pair of
spines in postorbital position. Antennal spine strong, directed forward, separated
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 4]
by short. curved carapace margin from longer branchiostegal spine directed
anteroventrally. Anterior submedian carinae consisting of seven elongate
tubercles. Posterior submedian carinae consisting of four elongate tubercles.
Row of small, rounded tubercles median to both anterior and posterior sub-
median carinae. Small median spine at base of rostrum flanked by two postero-
lateral spines. Anterior antennal carina forming wing-like flattened structure
ending in outward-pointed strong spine. Posterior antennal carina situated some
distance above anterior antennal carina, ending anteriorly in outwardly directed
spine, not as strong as that of anterior carina; at about midpoint, a slight indica-
tion of a notch, never strong enough to be called a spine or tubercle. Anterior
intermediate carina not well defined, consisting of three or four large irregularly
placed tubercles. Posterior intermediate carina consisting of four or five elongate
tubercles. Area between anterior submedian and anterior intermediate carinae
with two rows of small tubercles; three rows of tubercles between posterior
submedian and posterior intermediate carinae. Area between anterior antennal
carina and lateral groove carrying a scattering of small rounded tubercles. Areas
between posterior antennal and posterior lateral carinae also carrying irregular
Fig. 16. Glyphocrangon regalis.
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
scattering of rounded tubercles. Anterior and posterior lateral carina entire, not
tuberculate. Sublateral carina only distinct in posterior region, consisting of a
foveolate ridge. Region between sublateral carina and margin bearing a foveolate
band.
Abdomen (excluding telson) equal in length of carapace plus rostrum.
Median carina of abdominal segments strong, only on first segment forming a
forwardly directed spine. Pleura of first segment considerably shorter than second,
with forwardly directed spine. Pleura of second segment ventrally bispinose,
spines of equal strength. Pleura of third segment ventrally bispinose, anterior
spine longer than posterior. Pleura of fourth segment ventrally bispinose, spines
subequal. Pleura of fifth segment bispinose, spines directed posteroventrally.
Sixth abdominal segment ending posteroventrally in strong spine. Telson with
median spine at base, elongate triangular, with strong dorsolateral keel, apex
acute, flexed slightly dorsally.
Eyes well developed, with large black corneas, small spinose tubercle on
inner distal margin of eye-stalk.
Scaphocerite ovate, fringed with setae, lacking any spine on inner margin.
Second pereiopods unequal, left shorter and slightly squatter than right,
carpus of right (30-31 segments) considerably longer than left (19 segments),
chela markedly smaller.
Third pereiopod dactylus lanceolate, oval in cross section, one-third length
of propodus.
Fourth pereiopod dactylus slightly more than one-third length of propodus,
dorsally grooved.
Fifth pereiopod dactylus similar to fourth.
Previous records
Banda Island; off Makassar; Kermadec Island; Fiji; Paternoster Island.
Material
5 33 7 98 10 ovig. 29, see species list.
$3o CL 20,5 mm—-29,9 mm RL 16,5 mm-19,9 mm TL 82,5 mm—108 mm
ovig. 2° 29,3 mm—34,9 mm 17,8 mm—21,0 mm 108 mm—-128 mm
Remarks
The species recorded by Wood-Mason (1891) as G. investigatoris and
G. investigatoris var. andamanensis, was stated by De Man (1920) to be possibly
synonymous with G. regalis. Wood-Mason’s species, however, shows some
differences, e.g. in the degree of ‘flare’ of the spine of the postantennal carina, the
possession of midline tubercles on the rostrum which together suggest that this
is not the same species. Examination of the ‘Challenger’ types of G. regalis reveal
almost no differences from the present material.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 43
ACKNOWLEDGEMENTS
In addition to the people and institutions thanked in the introductory paper
of this series, I should like to thank Mr S. X. Kannemeyer of the South African
Museum for the photographs, Dr R. B. Manning of the Smithsonian Institution
for the loan of Pasiphaea material, Dr J. Yaldwyn of the National Museum of
New Zealand and Dr F. A. Chace Jr of the Smithsonian Institution for informa-
tion on Pasiphaea, and Prof. J. H. Day of the University of Cape Town and
Dr J. Yaldwyn for reading the manuscript and for their many useful criticisms.
REFERENCES
Atcock, A. 1892. Crustacea Part I. ///ustrations of the Zoology of the Investigator Plates 1-S.
Calcutta: Superintendent of Government Printing.
Atcock, A. 1899. Crustacea Part VII. ///lustrations of the Zoology of the Investigator Plates
36-45. Calcutta: Superintendent of Government Printing.
Awcock, A. 1901. A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura
and Anomala, in the Indian Museum. Calcutta: Trustees of the Indian Museum.
ALcock, A. & ANDERSON, A. R. S. 1899. Natural history notes from H.M. Royal Indian
Marine Survey Ship ‘Investigator’, Commander T. H. Heming, R.N., commanding.
Series III, no. 2. An account of the deep-sea Crustacea dredged during the surveying
season 1897-98. Ann. Mag. nat. Hist. (7) 3: 278-282.
AuLcock, A. & McARDLE, A. F. 1901. Crustacea Part IX. //lustrations of the Zoology of the
Investigator Plates 49-55. Calcutta: Superintendent of Government Printing.
BALss, H. 1925. Macrura der Deutschen Tiefsee-Expedition. 2. Natantia, Teil A. Wiss. Ergebn.
dt. Tiefsee-Exped. ‘Valdivia’ 20: 221-315.
BARNARD, K. H. 1926. Report on a collection of Crustacea from Portuguese East Africa.
Trans. R. Soc. S. Afr. 13: 119-130.
BARNARD, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with
notes on synonymy and new records. Ann. Mag. nat. Hist. (11) 13: 361-392.
BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and
shrimps). Ann. S. Afr. Mus. 38: 1-837.
Berry, P. F. & PLANTE, R. 1973. Revision of the spiny lobster genus Palinurus in the south-
west Indian Ocean. Trans. R. Soc. S. Afr. 40: 373-380.
BATE, C. S. 1888. Report on the Crustacea Macrura dredged by HMS Challenger during the
years 1873-1876. Rep. Voy. Challenger, 1873-1876 24: 1-942.
BURKENROAD, M. D. 1936. The Aristaeinae, Solenocerinae and pelagic Penaeinae of the
Bingham Oceanographic Collection. Bull. Bingham oceanogr. Coll. 5 (2): 1-151.
BURUCOVSKY, R. N. & ROMENSKY, L. L. 1972. On rostrum changeability of Aristeus varidens
Holthuis (Decapoda, Penaeidae). At/. Nauchno-Issled. Inst. Rybn. Khoz. Okeanogr. 42:
156-161.
CALMAN, W. T. 1939. Crustacea: Caridea. Scient. Reps. John Murray Exped. 6: 183-223.
CHACE, F. A. 1936. Revision of the bathypelagic prawns of the family Acanthephyridae, with
notes on a new family, Gomphonotidae. J. Wash. Acad. Sci. 26: 24-31.
CROSNIER, A. & Forest, J. 1973. Faune Tropicale. XIX. Les Crevettes profondes de I’ Atlantique
Oriental Tropical. Paris: O.R.S.T.O.M.
FisHER, L. R. & Goupig, E. H. 1961. New records of two deep-sea decapods. Crustaceana 2:
78-79.
Foxton, P. 1970. The vertical distribution of pelagic decapods (Crustacea: Natantia) collected
on the SOND cruise 1965. I. The Caridea. J. mar. biol. Assoc. U.K. 50: 939-960.
GEORGE, R. W. & GRINDLEY, J. R. 1964. Projasus—a new generic name for Parker’s crayfish,
Jasus parkeri Stebbing (Palinuridae: ‘Silentes’). J. Proc. R. Soc. West. Aust. 47: 87-90.
Gorpon, I. & INGLE, R. W. 1956. On a pelagic penaeid prawn, Funchalia woodwardi Johnson,
new to the British fauna. J. mar. biol. Assoc. U.K. 35: 475-481.
HALL, D. N. F. 1962. Observations on the taxonomy and biology of some Indo-West Pacific
Penaeidae. Col. Off. Fish. Publs 17: 1-230.
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ho.tuuts, L. B. 1955. The recent genera of the Caridean and Stenopodidean shrimps (Class
Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination.
Zool. Verhandl. 26: 1-157.
Kemp, S. 1910. The Decapoda Natantia of the coasts of Ireland. Scient. Invest. Fish. Brch Ire.
1908 (1): 1-190.
KENSLEY, B. F. 1969. Decapod Crustacea from the south-west Indian Ocean. Ann. S. Afr. Mus.
52: 149-181.
KENSLEY, B. F. 1971a. The Family Sergestidae in the waters around southern Africa (Crustacea,
Decapoda, Natantia). Ann. S. Afr. Mus. 57: 215-264.
KENSLEY, B. F. 1971b. The genus Gennadas in the waters around southern Africa. Ann. S. Afr.
Mus. 57: 271-294.
Louw, E. M. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data
1975, 1976. Ann. S. Afr. Mus. 72: 147-159.
Man, J. G. DE. 1911. The Decapoda of the Siboga Expedition. Part I. Family Penaeidae.
Siboga Exped. monogr. 39A: 1-131.
Man, J. G. DE. 1920. The Decapoda of the Siboga Expedition. Part IV. Families Pasiphaeidae,
Stylodactylidae, Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalido-
podidae, Gnathophyllidae, Processidae, Glyphocrangonidae, and Crangonidae. Siboga
Exped. monogr. 39A3: 1-318.
PAULINOSE, V. T. 1974. Decapod Crustacea from the International Indian Ocean Expedition:
The species of Funchalia (Penaeidae) and their post-larvae. J. nat. Hist. 8: 433-443.
RAMADAN, M. M. 1938. Crustacea, Penaeidae. Scient. Reps John Murray Exped. 5: 35-76.
RosBertTs, T. W. & PEQUEGNAT, W. E. 1970. Deep-water decapod shrimps of the Family
Penaeidae. Jn: Pequegnat, W. E. & Chace, F. A. Texas A & M University Oceanographical
Studies 1. Contributions on the Biology of the Gulf of Mexico. Houston, Texas: Gulf
Publishing Co.
SIVERTSEN, E. & HOLTHUuIS, L. B. 1956. Crustacea Decapoda (the Penaeidae and Stenopodidae
excepted). Rep. scient. Results Michael Sars N. Atlant. Deep Sea Exped. 5 (12): 1-54.
STEBBING, T. R. R. 1914. South African Crustacea. Ann. S. Afr. Mus. 15: 1-55.
TirMiIZzI, N. 1969. Eupasiphae gilesii (Wood-Mason, 1892) from the northern Arabian Sea
(Decapoda, Caridea). Crustaceana 16: 213-218.
Woop-Mason, J. & ALCOcK, A. 1891. On the results of deep-sea dredging during the season
1890-91. Ann. Mag. nat. Hist. (6) 8: 268-286.
Woop-Mason, J. & ALcock, A. 1893. On the results of deep-sea dredging during the season
1890-91. Ann. Mag. nat. Hist. (6) 11: 161-171.
~~
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, 1.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
‘not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. °... the Figure depicting C. namacolus ...’; ‘. .. in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
BRIAN KENSLEY
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 5
CRUSTACEA, DECAPODA, REPTANTIA
AND NATANTIA
’ ale, =
VOLUME 74 PART 3 NOVEMBER 1977 ISSN 0303-2515
MUS. COMP. Z
LIBRARY
JAN 3 1 1978
HARVARD
UNIVERSITY.
OF THE SOUTH AFRICAN
~ MUSEUM
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
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(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction
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(f) Summary, if paper is lengthy
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(a) Author’s name and year of publication given in text, e.g.:
‘Smith (1969) describes .. .’
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‘As described (Haughton & Broom Fees
‘As described (Haughton et al. 1927) .
Note: no comma separating name and: year
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Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. zen. 74: 627-63 4.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 1960b. Spawning behaviour, eee! masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4):
THIELE, J. 1910. Mollusca: B. Polypiecohorss Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 ~ Band
November 1977 November
Part 3 Deel
ILLUSTRATED GENERIC KEY TO
SOUTH AFRICAN CONTINENTAL OSTRACODA
By
K. G. McKENZIE
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
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Price of this part/Prys van hierdie deel
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ISBN 0 908407 24 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
ILLUSTRATED GENERIC KEY TO
SOUTH AFRICAN CONTINENTAL OSTRACODA
By
K. G. MCKENZIE
Riverina College of Advanced Education,
Wagga Wagga, Australia
(With 24 figures)
[MS. accepted 7 June 1977)
ABSTRACT
An illustrated generic key to South African continental ostracodes is provided together
with the diagnoses of two new cypridopsinid genera, Sarscypridopsis and Kapcypridopsis.
CONTENTS
PAGE
Introduction . : ; » 45
Systematics . ; , ft 46
Key : : : : . 49
Acknowledgements . ; : 53
References. A We HS
Appendix : 2 ; Bhs)
INTRODUCTION
In a previous paper (McKenzie 1971a) the known continental and mainly
freshwater Ostracoda of South Africa were listed together with a complete
bibliography of such ostracode studies in the Republic.
This paper complements that previous one by providing an illustrated key
to the known ostracode genera of South Africa. The key is meant to be used with
reference to illustrations in earlier papers by Sars (1896, 1924a, 19246) and Rome
(1965) as well as to the illustrations in this text.
Ostracoda are microscopic crustaceans with a bivalve shell which usually
completely covers the body when it is closed. This shell is commonly called the
carapace. The ostracode body has 5-7 pairs of limbs—7 pairs in freshwater
taxa—and, usually, a furca at the posterior, the rami of which attach to paired
chitin supports. From the anterior, the limbs are termed as follows: antennule;
antenna; mandible; maxillule; and the first through to the third thoracic limbs
(PI to PIII). The furca normally carries two claws and two bristles on each
ramus but can be strongly modified, e.g. into a whip-like seta in Cypridopsinae.
Sometimes, as in Darwinula, it is absent.
The furca in the genus Gesa is described in the key as having two claws
only. This follows the original description (Hartmann 1957). However, this
45
Ann. S. Afr. Mus, 74 (3), 1977: 45-103, 24 figs.
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
original description was of a juvenile form not of an adult. It is probable that
the furca in an adult Gesa will be found to carry the normal complement of
claws and bristles on each ramus.
Some characters which are illustrated in the Figures are not normally
recorded for ostracodes. These include the «, 6 and y bristles on the mandible
palp which have been so named by Danielopol (personal communication 1970)
and which appear to have some utility in taxonomy (e.g. compare Figures 6
and 14). Similarly, rake-like organs are very rarely illustrated but may also
prove to be useful taxonomic characters when known for more genera
(McKenzie 19715).
There is also a special terminology for characters of the ostracode carapace.
These include: adductor muscle scar patterns (Fig. | et seq.); marginal or radial
pore canals; normal pore canals which can be either simple and open or sieve-
like (Fig. 1); and various types of hingement. The hingement terminology used
here is that used in the American Treatise on Ostracoda (Moore ed. 1961) and
is illustrated in Figure 1.
Marginal pore canals are used to distinguish between genera in the tribe
Stenocypridini. In particular, the genus Chrissia Hartmann, 1957 is distinguished
by its very short marginal pore canals, whereas the species which were ascribed
to Parastenocypris (McKenzie 1971a) have well-developed branching marginal
pore canals (Figs 15-17).
In this paper the new taxa comprise two new cypridopsinid genera which
were foreshadowed earlier (McKenzie 1971a), namely: the genus Sarscypridopsis ;
and a new genus from Table Mountain, Kapcypridopsis. These new genera are
described in the systematics section below.
SYSTEMATICS
Family Cyprididae
Subfamily Cypridopsinae
Tribe Cypridopsini
Genus Kapcypridopsis gen. nov.
Figs 6-7
Type species
Kapcypridopsis barnardi sp. nov.
Derivation of name
Kap = Cape (Afrikaans), barnardi for the late Dr K. H. Barnard, a
tireless worker on South African crustaceans and molluscs and former director
of the South African Museum.
Diagnosis
In lateral view, carapace small; subreniform; smooth; greenish or yellowish;
valves subequal; dorsal margin convex; ventral margin inflexed medially:
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 47
anterior margin slightly more broadly rounded than the posterior; valve height
slightly more than half valve length. In dorsal view, elliptical. Internally, inner
lamellae broad anteriorly and posteriorly; inner margin regularly curved;
vestibule large; line of concrescence following the outer margin; marginal pore
canals short, fairly numerous; normal pore canals open and simple; muscle scar
pattern including 4 adductor and 2 mandibular scars (sometimes individual
adductor scars are divided and the adductor group appears to consist of 5 or
6 scars); hingement of simple ridge and groove type.
Antennule 7-segmented; length ratios of the 5 proximal segments are
20:13:10: 10:15 (or 15:10: 8:8: 11). Antenna 4-segmented, length ratios
of the 3 proximal segments are 7:4: 1; ‘natatory’ setae 6 in number, short,
reaching about the middle of the second endopod segment. Mandible coxa
normal; respiratory epipodial plate with 8 Strahlen; palp 4-segmented; «, f
and y bristles as illustrated. Maxillule respiratory epipod with 4 downwards-
directed setae plus 17 Strahlen; lobes and palp elongate; palp 2-segmented,
3rd lobe with two smooth Zahnborsten. PI epipod with 4 Strahlen; palp
strongly dimorphic as is usual in cypridaceans with 3 terminal bristles in females
but modified into a clasping organ in males. PII normal, with a powerful
terminal claw. P HI with a short terminal segment which bears a fairly short
reflexed seta. Furca strongly modified, whip-like. Rake-like organs each with
about 10 teeth. Eye with 2 ommatophores. In males, other dimorphic characters
include a Zenkers organ with 13 whorls; and hemipenis as illustrated (Fig. 7: 7).
Dimensions
Length, about 0,6 mm; height, about 0,35 mm.
Material
Small population of six individuals including mature males and ovigerous
females. Some forms were greenish, others yellowish, when collected.
Syntypes: South African Museum catalogue number SAM-A15451.
Locality
Top of Table Mountain, in small pool about 400m from the cable car
terminus; Cape Town, Cape Province, the Republic of South Africa.
Collector and data collected
K. G. McKenzie; September 1970.
Discussion
The regularly subreniform shape of this taxon is sufficient to distinguish it
from Plesiocypridopsis Rome, Cypridopsis Brady, and Sarscypridopsis gen. nov.
since Plesiocypridopsis and Sarscypridopsis are subtriangular in shape while
Cypridopsis is subovate and this new genus is subreniform. Also, unlike these
genera, Kapcypridopsis has broad anterior and posterior inner lamellae.
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
Genus: Sarscypridopsis gen. nov.
Fig. 8
Type species
Sarscypridopsis gregaria (Sars) 1896.
Derivation of name
For G. O. Sars, pioneer worker on South African Ostracoda.
Diagnosis
In lateral view, carapace small-medium; subtriangular; surface pitted and
frequently hirsute; often greenish to brown; valves subequal; dorsal margin
strongly convex; ventral margin inflexed medially; anterior margin rounded,
tending to narrow anteroventrally; posterior margin more broadly rounded
than the anterior margin; valve height usually well over half valve length. In
dorsal view elongate subovate moderately broad (breadth about half the length);
greatest breadth behind the middle. Internally, inner lamellae moderately broad
anteriorly and narrow posteriorly; inner margin regularly curved, vestibule
large, line of concrescence following the outer margin; marginal pore canals
short, numerous; normal pore canals simple, open; central muscle scar pattern
of 4 adductors and 2 mandibulars; hingement of simple ridge and groove type.
Antennule 7-segmented; antenna 4-segmented; natatory setae extending
well beyond the tips of the terminal claws; mandible normal, «, 8 and y bristles
as illustrated (Fig. 8: 2); maxillule palp 2-segmented, elongate-cylindrical;
3rd lobe with 2 smooth Zahnborsten; PI epipod with 2 Strahlen; endopod
modified in males into a clasping organ which is dissimilar between the right
and left limbs (Fig. 8: 10, 13); PII normal with powerful terminal claw;
PIII with a short reflexed seta; furca whip-like. In males, other dimorphic
characters include a Zenkers Organ with 10-15 whorls; and hemipenis as
illustrated.
Discussion
Of the three species in the original designation of Cypridopsis by Brady and
Norman in 1889, one, Cypridopsis vidua (O. F. Miller, 1776), is the type species;
C. villosa (Jurine, 1806) was made the type species of Cypridopsella Kaufmann,
1900, which is now synonymised with Potamocypris Brady, 1868, and C. aculeata
Costa, 1847, was included by Sars (1924a: 160) with the South African species
of Cypridopsis in his understanding of the genus at that time and indeed was
made its type. However, C. vidua has priority as the type species of
Cypridopsis s.s. which means that a different generic name is required for the
African and possibly one European species (note that the type material of
Costa’s species has been lost and his original figures and descriptions were poor).
The tribe Cypridopsini is differentiated from other cypridopsine ostracodes
by the fact that it possesses an elongate maxillule palp and lobes, the palp being
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 49
further characterized by an elongate second segment. This feature is in contra-
distinction to the broad palp, especially at the second segment, of Potamocypris
and its related cypridopsine genera. Considered in this sense, Cypridopsini
include the genera Cypridopsis, Plesiocypridopsis, Neocypridopsis, Kapcypridopsis,
the present new genus Sarscypridopsis, and two Korean subspecies (McKenzie
1972) which appear to be generically distinct from these others although similar
in some features to Kapcypridopsis.
This genus differs from Cypridopsis in that the shape is subtriangular and
moderately broad rather than subovate and broad; the anterior inner lamellae
are narrower, the ventral margin of the right valve is not denticulate; and there
are only up to two Strahlen on the epipod of the PI. It differs from Plesio-
cypridopsis in lacking the elongation of male PI palps and the characteristic
upturned anterior hemipenes of that genus. Kapcypridopsis is subreniform in
shape with broader anterior and posterior inner lamellae and short antennal
‘natatory’ setae, differing in all these characters from Sarscypridopsis. The
Korean taxa differ somewhat in shape especially posteriorly and have broad
anterior and posterior inner lamellae and short antennal ‘natatory’ setae. The
genus Neocypridopsis differs from Sarscypridopsis in general shape and also in
the setation of the terminal segment of the PIII, the shapes of the clasping
palps of the male P I and in possessing an upturned not a downturned anterior
hemipenis.
Of the cypridopsinid species listed by Sars (1924a) aculeata Costa, 1847,
spinifera Sars, 1924a, and echinata Sars, 1924a, form a subgroup characterized
by an armature of spines as well as hairs on the shell surface. These species are at
present classed with Sarscypridopsis but may merit transfer to a new category
when aculeata Costa, 1847, is better understood. Apart from this latter species,
which has been recorded from Iceland, Europe, central Asia, the Americas and
Africa, Sarscypridopsis has a southern distribution.
KEY
1. All three pairs of thoracic legs more or less pediform and non-
reflexed a ve sae a ue ae ae = Been ie
First pair of thoracic legs non- or subpediform, strongly dimorphic;
second pair pediform; third pair strongly reflexed i Cy priGidaen. /-
2. Shell small; smooth; elongate-ovate in lateral view; adductor-muscle
scar pattern feather-shaped; first pair of thoracic legs (PI) sub-
pediform; males not reported since 1870
Darwinulidae, Darwinula (Fig. 1)
Shell small to medium; surface usually ornamented; subrectangular,
subreniform, subtrapezoidal or subrhomboidal in lateral view;
adductor-muscle scar pattern of four discrete scars in a subvertical
row; all three pairs of thoracic limbs pediform; typically bisexual
Cytheridae 3.
50
Il,
ANNALS OF THE SOUTH AFRICAN MUSEUM
Shells small or medium; ornamented; subrectangular to subreniform;
radial pore canals few, straight; normal pore canals simple; hinges
modified lophodont ot i. Limnocytherinae 4.
Shells small or medium; prnamentede sulargetenoule, subtrapezoidal
or subrhomboidal; adel pore canals few or numerous, usually
straight; normal pore canals of sieve type; hinge types merodont,
entomodont or gonglyodont He Cytherideinae, Loxoconchinae 5.
Shell small; subrectangular to subreniform; non-dimorphic Limnocythere
Shell medium; subrectangular; strongly dimorphic (females very
broad, but males slender, posteriorly) .. .. Gomphocythere (Fig. 2)
Shell small to medium; subrectangular or subtrapezoidal; radial pore
canals few or numerous; hinge type merodont or entomodont
Cytherideinae 6.
Shell small; subrhomboidal; radial pore canals few, straight; hinge
type gonglyodont ..__.... a ie Loxoconcha (Fig. 1)
Shell small-medium; subtrapezoidal; adic pore canals few; hinge
type merodont ee ah .. Perissocytheridea
Shell medium; dubrestaneulan: bacliel pore canals numerous; hinge
type ene medont ne a ie a Cypriden (Fig. 1)
Shell medium; subrectangular; “ gipiaes pitted, dorsomedial sulci
present; palps of first pair of thoracic legs two-segmented
Ilyocypridinae, I/yocypris (Fig. 3)
Shell differing from this; palps of first pair of thoracic legs non-
segmented .. a 8.
Third thoracic limb beating at ee one 5 lone fefiexed feaninall ae:
Zenkers Organ (males only) with 5-6 whorls
Paracypridinae, Candoninae and Cyclocypridinae 9.
Third thoracic limb with short reflexed terminal seta; Zenkers Organ
(males only) with more than 6 whorls .. ae a,
Shell medium, subreniform or elongate sutenernaulle in fareral view;
elliptical in dorsal view, antennal natatory setae poorly developed or
absent e ae Paracypridinae, Candoninae 10.
Shell small- aaceliunns “gtibewathats in lateral view; antennal natatory
setae well developed (extending to the terminal claws or beyond)
Cyclocypridinae 11.
Shell medium, elongate-subtriangular in lateral view; furca with a
posterior bristle, furcal claws without spurs Paracypridinae, Paracypris
Shell medium, subreniform in lateral view; furca lacking a posterior
bristle, furcal claws with spurs... es Candoninae, Candonopsis
Shell small-medium; right valve denticulate ventrally especially on
anteroventral margin; compressed in dorsal view; furca normal
Cyclocypridinae, Physocypria (Figs 3-4)
Shell small; non-denticulate ventrally; broadened in dorsal view, furca
normal io a cn - .. Cyclocypridinae, Cyclocypris
12?
20.
Ale
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 51
Shells small to very large; furca with at least two terminal claws .. 19.
Shells small to medium; furca whip-like. . _ Cypridopsinae 13.
Shell with distinctive pustulose surface ornamentation and charac-
teristic lunette-bearing inner lamellae; maxillule 3rd lobe with at least
four Zahnborsten .. ie Oncocypris (Fig. 5)
Shells different; maxillule ard fone with ine Zahnborsten .. Ae 14:
Shell subtriangular, produced at each end; left valve overlapping right
valve anteriorly and posteriorly, but overlapped by the right valve
dorsally; maxillule palp distal segment spatulate Cyprilla (Fig. 5)
Shell of a different type; maxillule palp elongate oo Ue me ilps):
. Shell small-medium; subtriangular to subovate in lateral view; ovate
in dorsal view; left valve overlapping right valve anteriorly; anterior
inner lamellae broad, posterior inner lamellae narrow ue: ae NG:
Shells of a different type .. we we Ld
. Shell typically whitish to arcenich: Rewally Ge 408 or - nies irregular
dark strigae on each valve in life; right valve denticulate anteriorly;
inner lamellae as illustrated Ns wl Cypridopsis (Fig. 4)
Shell typically whitish; ornamented with nite arranged concentrically
or else tuberculate; without strigae; inner lamellae as illustrated
Zonocypris (Fig. 9)
. Shell small-medium to medium; subtriangular in lateral view;
relatively compressed in dorsal view; right valve overlapping left valve
anteriorly; anterior and posterior inner lamellae rather narrow;
antennal natatory setae well developed .. Wf Me AS:
Shell small to small-medium; valves subequal; aiibrenifornn in etal
view; without strigae on the valves; anterior and posterior inner
lamellae both relatively broad; antennal natatory setae short
Kapcypridopsis (Figs 6-7)
. In males, palp of first thoracic legs elongate, hemipenis anterior lobe
curved upwards .. set a Plesiocypridopsis (Fig. 9)
In males, palp of first ores. ines of normal length, hemipenis
anterior lobe pointing downwards Me Sarscypridopsis (Fig. 8)
Shell small-medium to medium; smooth (Cyprettini, Bradycyprini) or
pitted (Cyprettini) or striate (Bradycyprini); subglobose in lateral and
dorsal view; radial septa well developed at least in right valve; furca
‘normal’ but slender (Bradycyprini) or very slender (Cyprettini) |
Cyprettinae 20.
Shell medium to very large, typically smooth; compressed to globose
in dorsal view; without radial septa (exception Stenocypris, cf. 29);
furca ‘normal’, not conspicuously slender i ie ie Hi SohDD
Radial septa developed in both valves .. Cyprettini—Cypretta (Fig. 10)
Radial septa developed in right valve only ae Bradycyprini 21.
Shell striate .. ie ae ee ss Paracypretta (Figs 10-12)
Shell smooth iM ie as, a .. Bradycypris (Figs 11-12)
3
DD,
D3:
24.
2c
26.
Die
IES.
aD),
30)
Sill
Se
33)
ANNALS OF THE SOUTH AFRICAN MUSEUM
Chitin supports with loculi where they branch proximally, furca
flexuous he cz Cypricercini 23.
Chitin supports “fle oeull here hey branch jraees furca
typically straight or evenly curved, rarely flexuous... se 29
Shell oval, with distinct posterior process in some species; chitin
supports with anterior branch well developed; spermatic vesicles
($ only) coiled anteriorly .. xy ae S Cypricercus (Fig. 12)
Shell oval-oblong, without posterior processes; chitin supports
without an anterior branch ie < oh wm, 2Ae
Furca normally developed (2 claws and 2 BnBtes) ti Tanycypris
Furca with two claws only © ee) .. Gesa (cf. Introduction)
Shell without a posterior inner lamella (the inner margin coincides
posteriorly with the posterior shell margin cf. Rome 1965) .. wo 203
Shells with a posterior inner lamella es i ake 208
Distal segment of the. PII with a short donsal isi (Rome 1965)
Isocypridini, [socypris (Fig. 13)
Distal segment of the PII with a dorsal bristle which is almost as
long as the terminal claw (Rome 1965)
Amphibolocypridini, Amphibolocypris
Chitin supports widened distally into triangular chitinous processes
Herpetocypris, Stenocypridini 28.
Chitin supports without distal triangular chitinous processes ie” Sale
Furcal rami similar, furca with a posterior bristle
Herpetocypris (Figs 13-14)
Furcal rami dissimilar; furca without a posterior bristle
Stenocypridini 29.
Shells with radial septa... es on Ze eh Stenocypris
Shells without radial septa RS I ae ee 7 Lie
Marginal pore canals very short .. oe an .. Chrissia (Fig. 17)
Marginal pore canals well developed and branched
Parastenocypris (Figs 14-17)
Either right or left valve of shell denticulate ventrally except
Homocypris which, however, is of medium size only (length 1,1—1,4 mm)
and has anormal furca.. as .. Cyprinotini 32.
Neither right nor left valve a shel dendioulic ventrally (except
Apatelecypris cf. 42); carapace large, at least 1,75 mm, except
Mesocypris (0,91 mm) which has an unique furca, cf. 34 te .. 34.
Shell elongate-subreniform in lateral view; anterior inner lamellae
distinctly broader than posterior inner lamellae
Homocypris (Figs 17-18)
Shell elongate-ovate in lateral view; both anterior and posterior inner
lamellae relatively, and subequally narrow ae a We OSBE
Right valve denticulate antero- and/or poten ea uciln
Heterocypris (Fig. 18)
34.
3)
30:
Bi:
38.
jy).
40.
41.
42.
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA Do)
Left valve denticulate antero- and/or posteroventrally Hemicypris
Furca normally developed (2 claws, 2 bristles) and not modified as
below se » We
Furca with the oeted ar iyaeile ransomed ate a sialbibs Clas fie
spine; furcal rami weakly dissimilar sf E, Mesocypris
Carapace large, with prominent secondary eieedl zone in right valve
Afrocypris (Figs 18-20)
Carapace large, without prominent secondary fused zone in right valve 36.
Carapace large; maxillule palp elongate a ce er Oil
Carapace large; maxillule palp relatively broad (i.e. approximate
square or spatulate) = , oi
Carapace large, without ventral ‘ale: oemulifanete seamen: o P Ul
divided oe ae .. Eucypridini, Eucypris (Fig. 19)
Carapace large, quant or quot ventral ala; penultimate ‘segment’ of
PII not divided .. se oe 5s bts ee CyoriGdinigess
Carapace large, with ventral ala .. oA .. Pseudocypris (Fig. 20)
Carapace large, without ventral ala iu 3 Cypris
Carapace large, tuberculate in one species; anfennal natatory setae
well developed, reaching about as far as the distal part of the terminal
segment iy us ve Sclerocypris (Fig. 21)
Carapace large, eiiooth 6 distinetly hirsute; antennal natatory setae
strongly reduced .. a . 40.
Carapace large, in general shape dexeineline Pidioewprise PI male
($ only) strikingly dissimilar on left and right limbs; hemipenes
(¢ only) unlike those of any other ostracode; shape of female genital
lobe also distinctive cS, iar Liocypris (Fig. 22)
Carapace large; general shape nnlike that of Homocypris; P 1 palps
(g only) not strikingly dissimilar on left and right limbs, hemipenes
(¢ only) different to Liocypris .. ae ue Megalocypridini 41.
Carapace large; with a small but distinct posteroventral caudal spine
on each valve ss ne Hypselecypris
Carapace large; without ouch a spine on (Sihen selke ts oe . 42.
Carapace large; in dorsal view indented at both ends; not denuetiate
posteroventrally .. er a Megalocypris (Figs 23-24)
Carapace large; in dorsal view narrowing anteriorly, but rounded, not
indented, posteriorly; denticulate posteroventrally Apatelecypris (Fig. 24)
ACKNOWLEDGEMENTS
I am grateful to the National Institute for Water Research, Pretoria and
to the British Museum (Natural History) London for the opportunity to work
in South Africa and to Dr R. G. Noble who organized my visit. My original
drawings were traced for publication by Mr D. Goode of the Transvaal Museum.
My thanks are due to Professor P. H. A. Sneath and Dr M. J. Sackin of the
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
University of Leicester, who prepared the computer key of the appendix, using
the PANKEY program.
I am particularly grateful to Dr T. H. Barry, Director of the South African
Museum, for the extended use of its laboratory and library facilities during my
working visit to Cape Town, also to Dr B. F. Kensley at the South African
Museum for his considerable assistance in reorganizing the illustrations.
Mrs L. Vennell, Riverina College of Advanced Education, typed the
manuscript.
REFERENCES
HARTMANN, G. 1957. Ostracoden aus dem Namaland und Transvaal. Veroff. naturw.
Osnabruck 28: 50-60.
McKeEnzIE, K. G. 1971a. Species list of South African freshwater Ostracoda with an appendix
listing museum collections and some further determinations. Ann. S. Afr. Mus. 57:
157-213.
McKenzie, K. G. 1971b. Ostracoda from Lake Peunde, near Mt. Wilhelm, New Guinea.
Zool. Anz. 186: 391-403.
McKeEnzigE, K. G. 1972. Results of the speleological survey in South Korea 1966. XXII.
Subterranean Ostracoda from South Korea. Bull. Natl. Sci. Mus., Tokyo 15: 155-166.
Moore, R. C. ed. 1961. Treatise on Invertebrate Paleontology. Part Q. Arthropoda 3 Crustacea
Ostracoda. Lawrence: Geological Society of America and University of Kansas Press.
Rome, D. R. 1965. Crustacea: Ostracoda. Jn Hanstrom, B., Brinck, P. & Rudebeck, G., eds.
South African animal life 11: 9-58. Stockholm: Swedish Natural Science Research Council.
SARS, G. O. 1896. On some South African Entomostraca raised from dried mud. Skr.
Vidensk Selsk. Christiania 1895 (8): 1-56.
Sars, G. O. 1924a. The freshwater Entomostraca of the Cape Province (Union of South
Africa). Part Il: Ostracoda. Ann. S. Afr. Mus. 20: 105-193.
Sars, G. O. 1924b. Contributions to a knowledge of the fauna of South West Africa.
Ann. S. Afr. Mus. 20: 195-211.
APPENDIX
South African Freshwater Ostracoda Generic Key
(prepared by the Program PANKEY).
il. Shell: surface not smooth De
2: Furca reduced to whip-like seta 8:
3, Dorsal view of shell broad 4.
4, Maxillule 3rd lobe with 4 or more Zahnborsten, maxilla epipod with
less than 3 Strahlen, shell denticulation in one or both valves absent,
shell: radial septa or lunettes in inner lamellae present in both valves,
shell surface pustulose not tuberculate-costate. Oncocypris
4. Maxillule 3rd lobe with 2 Zahnborsten, maxilla epipod with more than
3 up to 6 Strahlen, shell denticulation in one or both valves present,
shell: radial septa or Junettes in inner lamellae, shell surface tuberculate-
costate not pustulose Zonocypris
3 Dorsal view of shell ‘normal’ 5)
Da Parthenogenesis: males virtually unknown, shell denticulation in one
or both valves present. Cypridopsis
De Parthenogenesis: both sexes known 6.
6. Maxilla male epipod not elongated. : Cyprilla
6. Maxilla male epipod elongated Plesiocypridopsis
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 53
De, Furca ‘normal’ ie
Us 7th limb reflexed, terminal seta long oye
8. Maxilla endopod 2-segmented, chitin supports: distally with
simple point, Zenker’s Organ with more than 15 whorls, shell:
dorsal sulci present Ilyocypris
8. Maxilla endopod unisegmented ;
9. Shell denticulation in one or both valves present Physocypria
oF Shell denticulation in one or both valves absent Cyclocypris
Is 7th limb reflexed, terminal seta short
10. Furcal rami very slender, Zenker’s Organ with 7-14 whorls,
shell: radial septa or lunettes in inner lamellae‘ present in both
valves, shell length 0,45—0,90 mm Cypretta
10. Zenker’s Organ with more than 15 whorls Ht
Wi. Furcal rami slender, shell: radial septa or lunettes in inner
lamellae present in one valve, shell surface striate, shell length
0,90-1,75 mm Paracypretta
iN Furcal rami ‘normal’ IZ,
12. Dorsal view of shell broad Cypris
I. Dorsal view of shell ‘normal’ Sclerocypris
Shell: surface smooth (includes weak pitting) 13.
I. 7th limb penultimate segment undivided 14,
14. Parthenogenesis: both sexes known, maxilla epipod with less than
3 Strahlen Candonopsis
14. Parthenogenesis: males virtually unknown IS
Sy Mandible palp comb present, maxilla epipiod with more than
7 Strahlen, maxilla endopod 3-segmented, 7th limb not reflexed,
furca absent, shell: inner lamellae absent. Darwinula
15. Mandible palp comb absent, maxilla epipod with more than 3 up to
6 Strahlen, maxilla endopod unisegmented (but ‘ghost’ segmentation
indicated), 7th limb reflexed, furca ‘normal’, shell: inner lamellae
present. Paracypris
13. 7th limb penultimate segment divided ! 16.
16. Natatory setae of antennae weakly developed Ws
We Maxilla epipod with less than 3 Strahlen, furca: posterior margins
dissimilar, shell densely hirsute Mesocypris
7: Maxilla epipod with more than 3 up to 6 Strahlen 18.
18. Furca reduced to whip-like seta, Zenker’s Organ with 7-14 whorls,
Shell length 0,45—0,90 mm Kapcypridopsis
18. Furca ‘normal’ 19.
19. Chitin supports: distally with simple point Liocypris
19. Chitin supports: distally elaborated 20.
20. Shell denticulation in one or both valves absent, shell length
3,60-7,30 mm Megalocypris
20. Shell denticulation in one or both valves present, shell length
1,75-3,60 mm Apatelecypris
16. Natatory setae of antennae well developed Die
le Chitin supports: distally elaborated DI
UD, Furca: posterior bristle absent M23
D3) Shell length 1,75-3,60 mm 24.
24. Shell: radial septa or lunettes in inner lamellae present in
both valves Stenocypris
24. Shell: radial septa or lunettes in inner lamellae absent
‘Parastenocypris’
W3). Shell length 0,90-1,75 mm WD
Di: 6th limb: distal bristle almost as long as the distal claw,
furca: posterior margins similar, chitin supports:
triangular process absent, dorsal view of shell very
compressed, shell: inner lamellae absent posteriorly
Amphibolocypris
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dy 6th limb: distal bristle short, furca: posterior margins
dissimilar, chitin supports: triangular process present,
dorsal view of shell normal, shell: inner lamellae present Chrissia
DOP Furca: posterior bristle present 26.
26. Chitin supports: triangular process present, shell length
1,75—-3,60 mm Herpetocypris
26. Chitin supports: triangular process absent 24
Dap Dorsal view of shell very compressed, shell: inner
lamellae absent posteriorly, shell length 0,90-1,75 mm __Isocypris
ile Dorsal view of shell ‘normal’ 28.
28. Shell posterior processes absent Afrocypris
28. Shell posterior processes present Hypselecypris
Ml. Chitin supports: distally with simple point ;
BS) Dorsal view of shell broad 30.
30. Furcal rami slender, shell: radial septa or lunettes in
inner lamellae present in one valve, shell length
0,90-1,75 mm Bradycypris
30. Furcal rami ‘normal’ 31.
31. Dorsal view of shell: alae absent Eucypris
31. Dorsal view of shell: alae present Pseudocypris
IES). Dorsal view of shell ‘normal’ Syd
oe Chitin supports: loculi present where they branch
proximally 338
33. Chitin supports: loculi present and prominent
anterior branch, shell posterior processes present
Cypricercus
333), Chitin supports: loculi present but without
prominent anterior branch, shell posterior
processes absent Tanycypris
3, Chitin supports: loculi absent where they branch
proximally 34.
34. Shell denticulation in one or both valves absent,
shell inner lamellae: anterior broad posterior
narrow Homocypris
34. Shell denticulation in one or both valves present 335).
35.0 Shell denticulation in right valve Heterocypris
35}, Shell denticulation in left valve Hemicypris
REFERENCE
PANKHURST, R. J., 1970. A computer program for generating diagnostic keys. Computer J. 12:
145-151.
Fig. 1
1. Darwinula sp.; ovigerous 2; adductor muscle scars, left valve. x 175.
2. Darwinula sp.; ovigerous °; outline right valve. x 125.
3. Generalized rimmed normal pore canal. x 1400.
4. Generalized rimless normal pore canal. x 2500.
5. Generalized sieve type normal pore canal. x 1000.
6. Generalized lophodont hingement right valve (after Moore ed. 1961).
7. Generalized merodont hingement right valve (after Moore ed. 1961).
8. Generalized entomodont hingement right valve (after Moore ed. 1961).
9. Generalized gonglyodont hingement right valve (after Moore ed. 1961).
10. Cyprideis sp.; hingement right valve. x 100.
11. Cyprideis sp.; adductor muscle scars and fulcral scar (dashes). x 225.
12. Loxoconcha sp.; adductor muscle scars and fulcral scar (dashes). x 300.
13. Loxoconcha sp.; anterior hingement right valve. x 250.
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA >)//
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 59
Fig. 2
Gomphocythere expansa Sars, 1924; SAM-A11303; ovigerous 9; PI and PII. x 400.
Posterior of body with caudal lobes. x 400.
P Ill. x 400.
Antennule. x 400.
Antenna. x 400.
ARWN-
ANNALS OF THE SOUTH AFRICAN MUSEUM
60
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 61
Fig. 3
fe
oe ee ee
Ilyocypris australiensis Sars, 1889; SAM-A11; adult 9; detail anterior margin right valve
showing radial pore canals and muscle scars. x 160.
Antennule, six terminal segments. x 400.
P III, terminal segments. x 400.
PI. x 400.
Mandible palp. x 400.
Furca. x 400.
Physocypria capensis (Sars, 1896); SAM-A11136; adult 3; internal view right valve.
x 160.
ANNALS OF THE SOUTH AFRICAN MUSEUM
62
ws 4 ZN | d ry \
Des e// 2 rT
Be ||) t/a) || eee
zane = <8
os Sekt | RL
OW :
fe
ys
if e
&
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 63
Fig. 4
Physocypria capensis (Sars, 1896); SAM-A11136; adult ¢; Zenkers Organ. x 400.
Mandible palp, detail 8 and y bristles. x 400.
Antennule, six terminal segments. x 400.
P III, terminal segment. x 400.
Cypridopsis viduella Sars, 1896; SAM-A11171; adult 2; PI. x 400.
Rake-like organ. x 400.
P III, terminal segment. x 400.
Antennule, six terminal segments. x 400.
Mandible palp, detail «, 8 and y bristles. x 400.
Internal view right valve. x 160.
Sie EON EA ee eS Seo
—
G4) ) (WO Sea), GOOPANINIATE S OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 65
Fig. 5
1. Oncocypris cf. voeltzkowi Miller, 1898; SAM—A11374; ovigerous 2; maxillule palp and
lobes, third lobe with some Zahnborsten. x 400.
2. PI. x 400.
3. Anterior margin of left valve. x 160.
4. Mandible palp, detail of «, B and y bristles. x 400.
5. Antennule, six terminal segments. x 400.
6. Cyprilla arcuata Sars, 1924; SAM-A11158; ovigerous 2; antennule, six terminal segments.
x 400.
7. Internal view left valve. x 160.
8. Maxillule palp. x 400.
9. PII. x 400.
0. Antenna, first endopod segment. ~ 400.
1. PII. x 400.
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 67
Fig. 6
1. Kapcypridopsis barnardi McKenzie, 1977; SAM-A15451; adult 2 (yellowish form);
internal view left valve. x 160.
2. Antennule segments. x 400.
3. Antenna, with ‘natatory’ setae. x 400.
4. Rake-like organ. x 400.
5. Mandible palp, detail of «, 8 and y bristles. x 400.
6. Maxillule palp and lobes. x 400.
7. PIII, terminal segments. x 400.
sae IT. >< -400.
Ss Burca. < 400.
0.
—"
PI. x 400.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 69
Fig. 7
Ile
NDAR WH
Kapcypridopsis barnardi McKenzie, 1977; SAM-—A15451; adult ¢ (greenish form); internal
view right valve. x 160.
P I right palp. x 400.
PII. x 400.
Zenkers Organ. x 400.
Eye. x 400.
Antennule, six terminal segments. x 400.
Hemipenis. x 400.
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
. Lee
@
aneeeBane =anOe =
1 =
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 1
Fig. 8
1. Sarscypridopsis striolata (Sars, 1924); SAM-—AI1201; ovigerous 9; internal view right
valve. x 160.
2. Sarscypridopsis ochracea (Sars, 1924); SAM-A11220; adult 3; mandible palp detail
-a, B and y bristles. x 400.
3. Antennule, six terminal segments. x 400.
4. Antenna, first endopod segment with sensory bristle. x 400.
5. Posteroventral right valve. =< 160.
6. PII. x 400.
7. Hemipenis. =< 400.
8. Maxillule third lobe with Zahnborsten. x 400.
9. Muscle scars. x 160.
10. PI, left palp. x 400.
- 11. PIII, terminal segment. = 400.
12. Maxillule palp. x 400.
13. PI, right palp. x 400.
ANNALS OF THE SOUTH AFRICAN MUSEUM
2
a
Th
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 73
Eig, 9
I.
eg
3g
4.
GN
Plesiocypridopsis cf. aldabrae (Miller, 1898); SAM-A5940; adult ¢; P I right palp.
x 400.
P I left palp. x 400.
Antenna, sensory bristle first endopod segment. x 400.
Zonocypris cordata Sars, 1924; SAM-—A11144; ovigerous 2; anterior margin left valve.
x 160.
Plesiocypridopsis cf. aldabrae (Miller, 1898); SAM-A5940; adult 2; ventral view of labia
with rake-like organs. x 400.
Mandible palp, detail of «, 68 and y bristles. 400.
Antennule, six terminal segments. x 400.
P III, terminal segment. x 400.
Adult 3; head and oral region. x 160.
Anteroventral right valve. ~ 160.
Hemipenis. x 400.
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA Is)
Fig. 10
. Cypretta cf. turgida (Sars, 1895); SAM—A11159; ovigerous 9; anterior margin right valve
with radial septa. = 160.
. Antenna endopod segments with sensory bristle. 400.
Antennule six terminal segments. x 400.
Mandible palp, detail of «, 8 and y bristles. x 400.
Maxillule palp and third lobe with Zahnborsten. x 400.
P I, terminal segments. x 400.
Distal furca. x 400.
. Paracypretta ampullacea Sars, 1924; SAM-A11278; ovigerous 2; sensory bristle first
endopod segment of antenna. x 400.
P III, distal terminal segment. x 400.
Mandible palp, detail of «, 8 and y bristles. x 400.
. Maxillule palp and third lobe with Zahnborsten. x 400.
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 77
Fig. 11
I
—
SO WIA
Paracypretta ampullacea Sars, 1924; SAM—A11278; ovigerous @; internal view left valve.
x 40.
2. Chitin support. = 160.
3.
4. Bradycypris intumescens (Brady, 1907); SAM-A11299; ovigerous 2; internal view right
Detail anterior margin radial pore canals left valve. x 160.
valve, showing radial septa. x 40.
Rake-like organ. x 400.
Antennule, six terminal segments. x 160.
Antenna, sensory bristle first endopod segment. x 400.
Maxillule palp and lobes, third lobe with Zahnborsten. x 160.
Mandible palp, detail of «, 8 and y bristles. x 400.
Mandible coxale. x 160.
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 719
Fig. 12
1. Paracypretta ampullacea Sars, 1924; SAM-A11278; ovigerous 9; furca. x 160.
2 Bye, < 400:
3. Antennule, terminal six segments. x 160.
4. Antennule, detail second segment. x 400.
5. Bradycypris intumescens (Brady, 1907); SAM-A11299; ovigerous 2; furca. x 160.
6. Chitin support. x 160.
7. Cypricercus cuneatus Sars, 1896; SAM-~A11140; adult 3; antennule, detail first and second
segments with gland and chemo-receptor. x 400.
Chitin support. x 160.
Antenna, sensory bristle of first endopod segment. x 400.
. Cap of Zenkers Organ. x 400.
. Mandible palp, detail of «, 8 and y bristles. x 400.
. PI, right palp. x 400.
. Furca. x 160.
. PI, left palp. x 400.
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 8]
Fig. 13
1. Isocypris priomena Miller, 1908; SAM-A11177; ovigerous 2; antennule second segment.
x 400.
Internal view left valve. x 40.
P II, three terminal segments. x 160.
Antenna endopod segments. x 160.
Anterior margin, detail radial pore canals. x 160.
Mandible palp, detail of «, 8 and y bristles. x 160.
P III terminal segment. x 400.
Maxillule palp. x 160.
Maxillule third lobe with Zahnborsten. x 160.
Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous 2; antennule second and
third segments. x 160.
11. Antennule second segment chemo-receptor. x 400.
12. Rake-like organ. x 160.
13. PIII, terminal segment. x 160.
14. Internal view left valve. x 40.
SESE ON a S
—
ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 83
Fig. 14
1. Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous 2; maxillule third lobe.
x 400.
2. Chitin support. x 160.
3. PI, terminal segments and part of claw. x 160.
4. Mandible palp, detail of «, B and y bristles. x 400.
5. Parastenocypris declivis (Sars, 1924); SAM-A3797; ovigerous 2; internal view, left valve.
x 40.
6. Distal furca. x 160.
7. Mandible palp, detail of «, 8 and y bristles. x 160.
8. Antennule second segment with chemo-receptor. x 400.
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 85
Fig. 15
igh se oe ee
Parastenocypris declivis (Sars, 1924); SAM-A3797; adult 2; chitin support. x 160.
Detail anterior margin left valve. x 160.
Detail posteroventral margin left valve. x 160.
Maxillule third lobe with Zahnborsten. x 400.
Parastenocypris hodgsoni (Sars, 1924); SAM-—A11126; ovigerous 9; internal view left
valve. x 16.
Furca, detail of posterior margin. x 160.
Detail anteroventral margin left valve. x 160.
Furca. x 40.
Furca, other ramus. x 160.
Antennule, detail of second segment chemo-receptor. 400.
. Parastenocypris pardalis (Sars, 1924); SAM-—A11167; internal view right valve. x 16.
Internal view left valve. x 16.
. Antenna, sensory bristle of basal endopod segment. x 400.
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 87
Fig. 16
1. Parastenocypris pardalis (Sars, 1924); SAM-—-A11167; adult 3; detail anterior margin left
valve. x 160.
2. PI, left palp. x 160.
3. PI, right palp. x 160.
4. Antennule second and third segments. x 160.
5. Parastenocypris olivacea (Sars, 1924); SAM-A3797; adult 3; internal view left valve.
x 16.
6. Muscle scars. x 160.
7. Detail anterior margin left valve. =< 160.
8. Antenna terminal segment, detail of comb bristle. x 400.
9. Mandible palp, detail of « and f bristles. =< 400.
10. Mandible palp, detail of y bristle. x 400.
11. Terminal rake-like organ. = 160.
12. Antenna basal endopod segment, showing exopod and sensory bristle. < 400.
13. Hemipenis. x 160.
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 89
Rig. 17
We
Parastenocypris olivacea (Sars, 1924); SAM-—3793; adult 3; distal furca. <= 160.
2. Frontal head region and labrum. x 160.
8.
?Chrissia fascigera (Sars, 1924); SAM-A11361; adult 9; internal view posteroventral left
valve. <x 160.
Anteroventral margin left valve, showing simple nature of radial pore canals. < 400.
Homocypris conoidea Sars, 1924; SAM-A11170; adult 9; detail anterior margin right
valve. x 160.
Muscle scars and detail of shell microstructure. x 160.
P III, terminal segments. x 160.
Chitin support. =< 160.
Internal view right valve. x 40.
Detail anteroventral margin right valve. x 160.
. Furca. x 160.
Antenna basal endopod segment, showing exopod, sensory bristle and part of the ‘natatory’ —
setae. < 160.
ANNALS OF THE SOUTH AFRICAN MUSEUM
90
11
10
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Dp
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queer ar’.
“see,
NY 4 :
Ns ~~
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 9]
Fig. 18
ii:
Oe ee
Homocypris conoidea Sars, 1924; SAM—A11170; adult &; mandible palp, detail of «, £
and y bristles. x 400.
Rake-like organs. = 400.
Antennule, detail second segment. x 400.
Maxillule third lobe with Zahnborsten. » 400.
P I, endopod and exopod, several bristles not shown. x 160.
Heterocypris aurea (Sars, 1896); SAM-A11310; ovigerous 2; antennule, terminal six
segments. x 160.
Mandible palp, detail of «, 8 and y bristles. x 160.
P III, terminal segment. x 160.
Internal view right valve. = 40.
Maxillule palp and lobes, third lobe with Zahnborsten. x 160.
meliurca,. <x, 160.
. PII, four terminal segments. x 160.
Afrocypris barnardi Sars, 1924; SAM-A11291; ovigerous 2; maxillule palp, second
segment. x 160.
Detail, anterior margin right valve. x 40.
. Internal view right valve. = 16.
92 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 93
Fig. 19
1. Afrocypris barnardi Sars, 1924; SAM-—-A11291; ovigerous 2; antennule, detail of second
segment. x 160.
2. Antenna, endopod segments. x 40.
3. Antenna, detail of sensory bristle on basal endopod segment. x 160.
4. Rake-like organ. x 160.
5. Mandible palp, detail of «, 8 and y bristles. =< 160.
6. PIII, terminal segment. x 40.
7. Distal furca. x 40.
8. Antennule, five terminal segments. x 40.
9. Chitin support. x 40.
10. PII, three terminal segments. x 40.
11. Eucypris purcelli Sars, 1924; SAM—A11287; adult 2; PII. x 160.
12. Distal furca. x 160.
13. Maxillule palp and third lobe showing Zahnborsten. x 160.
14. PI, exopod bristles not shown. x 160.
15. Antennule, detail of chemo-receptor. 400.
16. Antennule, six terminal segments. x 160.
ANNALS OF THE SOUTH AFRICAN MUSEUM
94
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA
Fig. 20
. Afrocypris barnardi Sars, 1924, SA\1 A11291; ovigerous ~; maxillule third lobe.
95
x 160.
Pseudocypris triquetra Sars, 1924; SAM-A11211; adult 3; maxillule palp and lobes.
<- 160:
P I, right palp. x 160.
Pislett palp: > 160.
P II, two terminal segments and claw. » 160.
Outline hemipenes. 40.
Chitin support. < 160.
Distal furca. = 160.
Antenna, sensory bristle. = 160.
P III, distal segment. = 160.
. Antennule, six distalmost segments. = 160.
. Maxillule third lobe showing Zahnborsten. 160.
Rake-like organ. x 160.
Mandible palp, detail x, 8 and y bristles. < 160.
ANNALS OF THE SOUTH AFRICAN MUSEUM
96
©
i
li
4
a,
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 97
Fig. 21
1. Sclerocypris clavularis Sars, 1924; SAM-A11118; ovigerous °; antennule, six distalmost
segments. x 160.
Rake-like organ. x 160.
P I, four terminal segments. x 160.
Chitin support. < 160.
Maxillule palp and third lobe showing Zahnborsten. x 160.
Carapace, muscle scars. = 40.
Mandible palp, detail «, 8 and y bristles. x 160.
Anterior margin left valve. =< 160.
Anterior margin right valve. = 160.
Internal view right valve. x 16.
Distal furca. x 160.
FHSS SNIDARYWNY
pay
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
—_*
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA
Fig. 22
1.
—a
ee aS
99
Liocypris grandis Sars, 1924; SAM-6281; adult $; carapace, detail anterior margin
reticulate valve microstructure (in part). x 40.
Internal view right valve. x 16.
Furca, distal detail. « 160.
Chitin support. x 40.
Antennule, detail of second segment showing gland and bristle. < 160.
Maxillule third lobe with Zahnborsten. x 160.
Maxillule palp. x 160.
PI palp. x 40.
Rake-like organ. 160.
Mandible palp, indicating complexity of setation. < 160.
Mandible epipod. x 160.
[ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 101
Fig. 23
1. Megalocypris hispida Sars, 1924; SAM-A11845; adult 2; mandible palp, detail of «, p
and y bristles. x 160.
2. SAM-A11844; adult $; antennule, six distalmost segments. x 40.
3. SAM-A11845; adult 2; mandible coxale, detail of toothbrush bristles between first and
second teeth. x 400.
4. Megalocypris sp. (smooth form with hemipenis similar to M. Hiberelitatt Sars, 1924);
SAM-A11849; adult 3; antenna, endopod segments. x 40.
and 6. Rake-like organs. x 160.
Mandible palp, detail of « and f bristles. Note that this male carries only one y bristle.
x 160.
8. Megalocypris princeps Sars, 1898; SAM-1479; adult ¢; rake-like organ. x 160.
9. and 10. Megalocypris tuberculata Sars, 1924; SAM-A11273; adult 3; rake-like organs.
x 160.
11. PI palp. x 40.
12. Furca. x 40.
13. Chitin support. x 40.
14. PII, three terminal segments. x 160.
15. Maxillule palp. x 160.
16. Maxillule third lobe with Zahnborsten. x 160.
17. Antenna, sensory bristle of first endopod segment. x 160.
18. Carapace, muscle scars. x 40.
19. Outline of hemipenis. x 40.
apes
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 103
Fig. 24
Megalocypris tuberculata Sars, 1924; SAM—A1i1273; adult $; mandible palp, detail of «,
B and y bristles. Note that the specimen may be a freak since it carries two y bristles
instead of the normal complement of one only. x 160.
Antennule, detail of second segment showing gland, chemo-receptor and bristle. x 160.
Apatelecypris brevis Sars, 1924; SAM-—A11369; adult ¢; carapace, internal view right
valve, muscle scars not shown. x 16.
Detail anteroventral radial pore canals. = 160.
Detail anterior margin right valve. x 40.
Antenna, detail ‘natatory’ setae. =< 160.
P I, detail of left palp. x= 160.
P I, detail of right palp. x 160.
Antennule, detail of second segment showing chemo-receptor and bristle. x 160.
Mandible palp, detail of y bristle. = 160.
Mandible palp, detail of « and f bristles. = 160.
. Tips of rake-like organs. x 160.
Furca. x= 40.
Maxillule, palp and third lobe showing Zahnborsten. = 160.
. Chitin support. x 40.
. PII, three terminal segments. x 160.
Dat Poke
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’
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, €.g. gen. NOV., Sp. Nov., comb.
nov., syn. nov., etc.
‘An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed i in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus...’; ‘*. . . in C. namacolus (Fig. 10)...”
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L. du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
K. G. McKENZIE
ILLUSTRATED GENERIC KEY TO
SOUTH AFRICAN CONTINENTAL OSTRACODA
e SOU Wy a* ee
- VOLUME 74 PART 4 NOVEMBER 1977 | ISSN 0303-2515
-_ MUS. COMP. ZOOL: |
» LIBRARY
JAN 3 1 1978
HARVARD
UNIVERSITY
_ANNALS
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
(c) Table of contents giving hierarchy of headings and subheadings
(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 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of
legends for illustrations should be typed separately, their positions indicated in the text. All
pages should be numbered 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
legend); the reduction or enlargement required should be indicated; originals 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 legend; 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.
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, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 1960b. Spawning behaviour, eae masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4 ): 1.
THIELE, J. 1910. Mollusca: B. Dalene oubea Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
November 1977 November
Part 4 Deel
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 6
AMPHIPODA
By
CHARLES GRIFFITHS
Cape Town Kaapstad
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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 6
AMPHIPODA
By
CHARLES GRIFFITHS
Zoology Department, University of Cape Town
(With 10 figures)
[MS. accepted 4 August 1977]
ABSTRACT
Amphipoda recovered during the 1975, 1976 Meiring Naude cruises are listed and species
of note discussed. The collection includes two new genera —Jzinkala gen. nov. and Valettiella
gen. nov. (Lysianassidae)—represented by J. fihla sp. nov. and V. castellana sp. nov. Three
further species are recorded from southern Africa for the first time and Bathyamaryllis
conocephala (K. H. Barnard) is redescribed.
CONTENTS
PAGE
Introduction yi. ae ee LO>
IPiStrofaspeciesi 49 ate ee a LOO
Systematic discussion. . . 108
Acknowledgements . . . 123
References Aa ahaa hi eB) 128)
INTRODUCTION
The material described herein is derived from collections made by the staff
of the South African Museum during exploratory cruises by the R.V. Meiring
Naude in 1975 and 1976. These cruises are of particular interest since they have
investigated deeper waters off Natal which have previously received scant
attention, despite their potential zoogeographical interest. A full listing of
station data from both cruises has been presented elsewhere (Louw 1977), but
few of these samples in fact used gear suitable for the capture of benthic
amphipods. Although the number of amphipods recovered was limited, the
proportion of new or unusual species in the collection bears witness to the
potential for further exploration of this area.
All material described below has been deposited in the South African
Museum, Cape Town.
105
Ann. S. Afr. Mus. 74 (4), 1977: 105-123, 10 figs.
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
LIST OF SPECIES
SM Station Numberof
Number Individuals
Suborder GAMMARIDEA
Family Acanthonotozomatidae
Dikwa acrania Griffiths . ; ‘ : eg ae ; 86 2
Family Ampeliscidae
Ampelisca anomala Sars. : ; : : : P } 86 10
Ampelisca brevicornis (Costa) . A : : : ; : 69 1
Ampelisca byblisoides K. H. Barnard Wer oe Ee, Oe 53 4
103 1
Ampelisca palmata K. H. Barnard . es ee Me ee 53 3
Byblis anisuropus Stebbing . ; ; : , : ; : 86 1
Byblis gaimardi (Kroyer) .. ane an Fi eee 60 1
109 3
Family Amphilochidae
Hoplopleon medusarum K. H. Barnard . ; : ; =) 103 1
Unguja yaya Griffiths . ; ; : : : : : : 86 1
Family Corophiidae
Chevalia aviculae Walker . ; : : : : : ’ 86 77
103 2
Concholestes armatus Griffiths . . . . 2 Ts hae 31 3
78 31
86 4
103 34
109 13
Gammaropsis afra Stebbing ene Set ast Ly ean 86 4]
91 3
103 32
109 2
Photis uncinata K. H. Barnard . 5 , : : : : 86 26
Unciolella spinosa Griffiths ath Me alt , Neh Ml 60 1
69 1
86 8
109 1
Family Dexaminidae
Atylus homochir Haswell . , : ; ; E é J) £09 1
Lepechinella occlo J. L. Barnard at Pee ae i ecauh esc 60 1
Family Eusiridae : ns ie ee Pe es 86 9
_ Dautzenbergia mee (Chevreux) : . ‘ 5 : 60 1
107 4
Family Gammaridae
Maera inaequipes (Costa) i Oe ae 86 6
Maera mastersi (Haswell) . ; : . : ; : : 86 2
Family Haustoriidae
Urothoe elegans Bate . : : : : : : 53 1
Urothoides inops J. L. Barnard . . : : s ; : 86 1
Family Ischyroceridae
Cerapus tubularis Say . : , : ; ; : , 5-408 26
Family Leucothoidae 3
Leucothoe dolichoceras K. H. Barnard . $ 5 ; F 103 2,
Leucothoe spinicarpa (Abildgaard) . : E : : ; 86 12
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
Family Liljeborgiidae
Liljeborgia epistomata K. H. Barnard
Family Lysianassidae
Amaryllis macrophthalma Haswell
Bathyamaryllis conocephala (K. H. Barnard) .
Hippomedon longimanus (Stebbing) .
Hippomedon onconotus (Stebbing)
Izinkala fihla gen. et sp. nov.
Lepidepecreum clypeatum Chevreux .
Lepidepecreum twalae Griffiths .
Lysianassa ceratina
Valettiella castellana gen. et sp. nov.
Family Oedicerotidae
Perioculodes longimanus (Bate & Westwood) .
Family Phoxocephalidae
Heterophoxus opus Griffiths
Family Podoceridae
Laetmatophilus purus Stebbing
Family Stegocephalidae
Stegocephaloides australis K. H. Barnard
Family Stenothoidae
Parametopa grandimana Griffiths
Proboloides rotunda (Stebbing) .
Superfamily TALITROIDEA
Family Phliantidae
Plioplateia triquetra K. H. Barnard .
Suborder CAPRELLIDEA
Family Aeginellidae
Eupariambus fallax K. H. Barnard
Pseudoprotella phasma (Montagu)
Family Phtisicidae
Phtisica marina Slaber
107
SM Station Number of
Number
86
Individuals
y
SSS SS So Sa wwe seme
~] 0O — W
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
SYSTEMATIC DISCUSSION
Family Acanthonotozomatidae
Dikwa acrania Griffiths, 1974
Fig. |
Dikwa acrania Griffiths, 1974: 266, fig. 2.
Material
SM 86, 2 specimens
Remarks
This record is only the second of this interesting species and extends its
range considerably to the north. Further drawings of certain appendages are
provided here to supplement the original description. These reveal a number of
interesting features previously unnoticed. The surface of the mandibular molar
is strongly triturative and projects a considerable distance from the body of the
mandible, the apical margin is formed into a sharp chitinized ridge. Article 2 of
pereiopods 3 and 4 is posteriorly keeled on inner and outer margins, leaving a
Fig. 1. Dikwa acrania Griffiths, 1974.
Ovigerous female, 4,5 mm. A. Pereiopod 4 with dactyl enlarged. B. Medial view of pereiopod 5.
C. Mandible. D. Telson. ;
THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 109
deep groove posteriorly, article 4 is acutely produced distally and the dactyl
bears small chitinized teeth. The fifth perelopod was erroneously described as
missing in the original description. It is, in fact, grossly reduced with article 2(?)
apparently fused to, and partially covered by, an enlarged coxa 7; the remainder
of the appendage consists of two small articles bearing short anterior spines.
This remarkable feature is somewhat similar to that found in Tetradeion
(Stegocephalidae) and should be incorporated into the generic definition of
Dikwa.
Distribution
Endemic, Still Bay to Zululand 200-550 m.
Family Dexaminidae |
Lepechinella occlo J. L. Barnard, 1973
Fig. 2
Lepechinella occlo J. L. Barnard, 1973: 21, figs 8-9.
Material
SM 60, | specimen
SM 86, 9 specimens
Remarks
The present material differs only marginally from that described by
Barnard. Thus in South African specimens coxae | and 2 are more distinctly
bifid, the accessory teeth on pereon segments 6 and 7 and pleon segments |-3
somewhat more distinct, and the epimeral setae less marked. These differences
cannot be considered of taxonomic significance, particularly in the light of
similar size-related differences reported by Barnard (1973).
Distribution
New Zealand, 721-860 m; Natal 550-810 m. This record is the first from
southern Africa.
Family Eusiridae
Dautzenbergia grandimanus (Chevreux, 1887)
Fig. 3
Parapleustes megachir Walker, 1897: 230, pl. 18 (fig. 4).
Sympleustes megachir: Stebbing, 1906: 317. Stephensen, 1944: 5, fig. 1.
Sympleustes grandimanus: Sexton, 1909: 857, pl. 90 (figs 8-32).
Sympleustes (Dautzenbergia) grandimana: K. H. Barnard, 1937: 158.
Dautzenbergia grandimanus: J. L. Barnard, 1961: 106.
Material
SM 60, | specimen
SM 107, 4 specimens
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Lepechinella occlo J. L. Barnard, 1973.
Male, 6 mm. A. Lateral aspect. B. Accessory flagellum. C. Mandible. D-—E. Coxae |, 2.
F. Third pleonal epimeron. G. Uropod 3. H. Telson.
THE SOUTH AFRICAN MUSEUM'S ME/RING NAUDE CRUISES 11]
A
Fig. 3. Dautzenbergia grandimanus (Chevreux, 1887).
Ovigerous female, 8,5 mm. A. Upper lip. B. Lower lip. C. Mandible. D. Maxilla 1.
E. Maxilliped. F—G. Gnathopods 1, 2. H. Uropod 3 and telson. I. Telson.
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
The genus Dautzenbergia has recently been revived by J. L. Barnard (1961)
to accommodate species of ‘“Sympleustes’ with a partially cleft telson. Of the
three species in the genus D. grandimanus and D. megacheir appear to be
indistinguishable. The present material correlates closely with the descriptions
given by Sexton (1909) and Stephensen (1944).
Distribution
North Atlantic, Indian Ocean. This is the first record from southern
Africa.
Family Haustoriidae
Urothoides inops J. L. Barnard, 1967
Fig. 4
Urothoides inops J. L. Barnard, 1967: 23, figs 8-10.
Material
SM 86, | specimen
Remarks
There can be no doubt that the present material is synonymous with that
illustrated by J. L. Barnard (1967). This unusual species can readily be dis-
tinguished by its broad, flat, down-turned rostrum, which covers the anterior
end of the body and deflects antenna 2 laterally. Pereiopods 3-5 are strongly
developed and have article 2 thickened by glandular tissue. The mouthparts, and
particularly the mandibles, are extremely large relative to the body, which is
almost as broad as long.
Distribution
California, 2 700m; Natal 550m. This is the first record of this species
from the southern African region.
Family Lysianassidae
Bathyamaryllis conocephala (K. H. Barnard, 1925)
igi)
Amaryllis conocephalus K. H. Barnard, 1925: 324.
Bathyamaryllis conocephala: K. H. Barnard, 1940: 441.
Material
SM 61, 2 specimens (including neotype SAM-—A13657)
SM 78, | specimen
SM 109, | specimen
SM 103, fragment
SM 60, 2 specimens
THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 113
Fig. 4. Urothoides inops J. L. Barnard, 1967.
Female, 2 mm. A. Ventral view of rostrum. B. Mandible. C. Lower lip. D. Maxilliped.
E. Gnathopod |. F-H. Pereiopods 3, 4, 5. I. Uropod 3. J. Telson.
114 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Bathyamaryllis conocephala (K. H. Barnard, 1925).
Female, 5 mm. A. Lateral aspect. B. Head and gnathopod 1. C. Mandible. D. Maxilla 1.
E. Maxilliped. F. Gnathopod 2. G-—H. Uropods 2, 3. I. Telson.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES HUES)
Remarks
This species has hitherto been known only through K. H. Barnard’s
somewhat scanty and unfigured description of the holotype female. When an
attempt was made to obtain this specimen from Barnard’s type collection it was
found to have completely decomposed. A neotype has hence been selected from
the present material and is shown in Figure 5. The bevelled off antero-lateral
angles of the head clearly identify this species and distinguish it from the allied
B. rostrata (Chevreux), in which the lateral lobes of the head are large and
vertically truncated. The unusual form of the head in these two species is quite
distinct from that in the other members of the genus—B. haswelli (Stebbing) and
B. perezii Pirlot.
Distribution
Endemic, Cape Peninsula to Natal, 680-1 300 m.
Izinkala gen. nov.
Diagnosis
Head and coxa | both very small, largely concealed by enlarged coxa 2;
epistome and upper lip inconspicuous; mandible with distinct non-dentate
incisor, palp attached distally; maxillae poorly developed; maxilliped lacking
plates, palp with minute article 4; gnathopod | slender, minutely subchelate,
articles 3 and 5 elongate; gnathopod 2 chelate; uropod 3 reduced, enclosed
within urosomite 3, outer ramus with small article 2; telson entire.
Type species
Izinkala fihla sp. nov.
Relationships
This remarkable genus appears quite discrete from any known form. The
greatly reduced mouthparts are of particular note. A comparable degree of
reduction occurs in Kerguelenia Stebbing, but here the mandible lacks both
incisor and molar, while the maxilliped retains small plates and a normal palp
of four articles. The two genera, however, differ grossly in other respects, such
as the form of gnathopod | and relative prominence of coxa |, although other
notable similarities with certain species of Kerguelenia occur, for example the
enclosure of uropod 3 within urosomite 6 in K. borealis Sars. Izinkala gen. nov.
also bears a certain resemblance to various genera in the Lepidepecreum group.
Thus species such as L. clypodentatum J. L. Barnard show carinate peduncular —
segments in antenna |, similar gnathopods | and 2 and enlarged article 2 of
pereiopod 5. Clepidecrella J. L. Barnard lacks certain of these features but has
reduced maxillipedal plates, a short uropod 3 and entire telson. Despite these
features, /zinkala gen. nov. remains quite distinct with its grossly reduced coxa |
and forward projecting, enlarged coxa 2.
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
Izinkala fihla sp. nov.
Figs 6-7
Description (of ?female 3,2 mm)
Head shorter than pereon segment |, post-antennal angles acutely produced;
peduncle of antenna | large, article | acutely produced, article 2 overhanging
remainder of antenna as a broad lobe with lateral flanges between which distal
articles originate, flagellum 4-articulate, article 1 broad, posteriorly fringed with
long aesthetascs, accessory flagellum 3-articulate; antenna 2 with article 1 sub-
circular, apparently fused to head, gland cone prominent, flagellum 3-articulate;
epistome and upper lip inconspicuous; mandible with non-dentate incisor, spine
row absent, molar apparently large and smooth; maxillae not satisfactorily
resolved, minute; maxilliped completely lacking both inner and outer plates,
palp article 4 reduced.
Coxa | small and thickened by muscle fibres, completely concealed by large
forward-projecting coxa 2, margins of coxae 2-4 of the two sides abutting
ventrally, the small gap below the head closed by projecting lobe of article 2 of
antenna 1; gnathopod | slender, elongate, articles 3 and 5 each as long as 2,
palm slightly oblique, defined by a single strong spine which projects into a
pocket formed within the thick, highly chitinized dactyl, which bears sharp
transparent lateral flanges; gnathopod 2 chelate; article 2 of pereiopod 3
rotund, article 4 greatly widened posteriorly; pereiopod 4 similar in structure to
3 but longer; pereiopod 5 with article 2 elongate and produced distally as far as
tip of article 4, articles 4 and 5 acutely lobed posteriorly.
Pleonal epimera | and 2 smoothly rounded, 3 rounded-quadrate; pleon
segment 4 with a rounded dorsal carina overhanging pleon segment 5; uropods |
and 2 strongly spinose dorsally, rami equal; pleon segment 6 overhanging
uropod 3 and telson above and to the sides; uropod 3 very small, outer ramus
with a spinose article 2 overhung by dorsal projection of article 1, inner ramus
shorter, evenly tapering; telson elongate, entire, bearing strong dorsal and
terminal spines.
Holotype
SAM-A13659, ?female, 3,2 mm.
Type locality
SM 86, 27°59,5’S 32°40,8’E, 550 m, 22 May 1976.
Material
SM 86, 4 specimens
SM 103, 1 specimen
Valettiella gen. nov.
Diagnosis
Antenna | geniculate, articles 2 and 3 slender and elongate; article | of
antenna 2 greatly enlarged; upper lip and epistome inconspicuous; mandibular
THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES LF
Fig. 6. Izinkala fihla gen. et sp. nov.
Female 3,2 mm. A. Lateral aspect. B. Antennae | and 2. C. Mandible. D. Maxilliped
E-F. Unidentified mouthparts, probably maxilla 1.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
G
Fig. 7. Izinkala fihla gen. et sp: nov.
Female, 3,2 mm. A. Gnathopod | with palm enlarged. B. Gnathopod 2. C-—D. Pereiopods 3, 5.
E, Lateral view of urosome. F. Uropod 3. G. Telson.
THE SOUTH AERICAN MUSEUM'S MEIRING NAUDE CRUISES 119
molar triturative, incisor strongly toothed, palp attached level with molar;
inner plate of maxilla | setose medially; coxae | and 2 rectangular and equally
elongate; gnathopod |! weakly chelate; gnathopod 2 subchelate; telson short,
cleft.
Type species
Valettiella castellana sp. nov.
Relationships
The strongly dentate mandibular incisor and unspecialized gnathopods of
this genus place it in a distinct group presently comprising Valettia Stebbing,
1888 and Valettiopsis Holmes, 1908.
The new form, however, shows features intermediate between those
defining Valettia and Valettiopsis, as well as a number of distinct characteristics
of its own. Thus Valettia has the outer plate of the maxilliped apically produced
and coxae | and 2 similarly short and broad, while Valettiopsis has a normal
maxilliped but coxa 2 elongate and almost covering a reduced coxa |. Valettiella
gen. nov. not only combines a normal maxilliped with elongate and equal
coxa | and 2, but also differs from both the above genera in its unusual geniculate
antenna i with elongate peduncle, enlarged article | of antenna 2, widened and
castelloserrate article 2 of posterior pereiopods and shortened uropod 3 with
reduced inner ramus.
Valettiella castellana sp. nov. |
Figs 8-9
Description (of ovigerous female, 4 mm)
Head as long as first two pereon segments, ocular lobes acute, eyes weak;
antenna | with article | elongate, article 2 almost as long as 1, normally folded
back into ventral groove of article |, flagellum 9-articulate, accessory flagellum
uni-articulate; article 1 of antenna 2 acutely produced into an enlarged shield
projecting just below ocular lobe of head, gland cone prominent, flagellum
5-articulate; mandible with 3-articulate palp, incisor with 6 strong teeth, lacinia
mobilis with 4 teeth, spine row of 8 spines, molar large and strongly ridged;
maxilla | with bi-articulate palp projecting beyond tip of outer plate, palp with
7 terminal spine teeth, outer plate bearing 8 strong serrate spines, inner plate
setose medially; inner plate of maxilla 2 strongly setose medially; maxilliped
with powerful 4-articulate palp, outer plate apically rounded, bearing eight
medial spine teeth.
Coxae 1-3 equally elongate, | and 2 with small postero-distal tooth;
gnathopod | weakly chelate, article 5 considerably larger than 6; gnathopod 2
with article 3 only slightly elongate, articles 5 and 6 subequal, palm oblique;
pereiopods 3-5 progressively longer, article 2 greatly lobed posteriorly, anterior
margin spinose, posterior margin strongly castelloserrate.
120 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Valettiella castellana gen. et sp. nov.
Ovigerous female, 4mm. A. Lateral aspect. B. Antenna | with accessory flagellum enlarged.
C. Antenna 2. D. Mandible. E. Lower lip. F. Maxilla 1.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES U2
B
Fig. 9. Valettiella castellana gen. et sp. nov.
Ovigerous female, 4 mm. A. Maxilla 2. B. Maxilliped. C. Gnathopod 1. D. Gnathopod 2
with palm enlarged. E. Pereiopod 5. F—H. Uropods 1, 2, 3. I. Telson.
| DD ANNALS OF THE SOUTH AFRICAN MUSEUM
Pleon segments |, 2, 4 and 5 each bearing a single medio-dorsal tooth,
pleonal epimera quadrate postero-distally; rami of uropods | and 2 equal,
strongly spinose dorsally and apically; uropod 3 considerably shorter than |
and 2, outer ramus with a small article 2, inner ramus about half size of outer
and closely appressed to its medio-ventral surface; telson short, cleft, each lobe
bearing a large terminal spine and three small setae.
Holotype
SAM-A13658, ovigerous female, 4 mm, unique.
Type locality
SM 86, 27°59,5’S 32°40,8’E, 550 m, 22 May 1976.
Family Stenothoidae
Parametopa grandimana Griffiths, 1974
Fig. 10
Parametopa grandimana Griffiths, 1974: 324, fig. 18.
Material
SM 86, 13 specimens
SM 103, |! specimens
Fig. 10. Parametopa grandimana Griffiths, 1974.
Male, 4,5 mm. A. Gnathopod 1. B. Gnathopod 2.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES IZ
Remarks
The above material includes adult males of up to 4,5 mm which show
gnathopods somewhat different to those described by Griffiths (1974). In
terminal males the palm of gnathopod | is defined by two small spines and the
dactyl bears three setae. Article 6 of gnathopod 2 is somewhat more elongate
than is typical of smaller males and the palm is undefined, while the dactyl
becomes greatly thickened terminally.
Distribution
Endemic, Still Bay to Natal 200-680 m.
ACKNOWLEDGEMENTS
This project was made possible through the generosity of the staff of the
South African Museum, who not only lent the author the material but also
provided excellent library facilities. Iam indebted to the South African Council
for Scientific and Industrial Research for financial support. My thanks also to
Mrs S. Hardman who kindly typed the manuscript.
REFERENCES
BARNARD, J. L. 1961. Gammaridean Amphipoda from depths of 400 to 6000 meters.
Galathea Rep. 5: 23-128.
BARNARD, J. L. 1967. Bathyal and abyssal gammaridean Amphipoda of Cedros Trench, Baja
California. Bull. U.S. natn. Mus. 260: 1-205.
BARNARD, J. L. 1973. Deep-sea Amphipoda of the genus Lepechinella (Crustacea). Smithson.
Contr. Zool. 133: 1-31.
BARNARD, K. H. 1925. Contributions to the crustacean fauna of South Africa. 8. Further
additions to the list of Amphipoda. Ann. S. Afr. Mus. 20: 319-380.
BARNARD, K. H. 1937. Amphipoda. Scient. Rep. John Murray Exped. 1933-34 5: 131-201.
BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. 12. Further
additions to the Tanaidacea, Isopoda, and Amphipoda with keys for the identification of
hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543.
GRIFFITHS, C. L. 1974. The Amphipoda of southern Africa, Part 4. The Gammaridea and
Caprellidea of the Cape Province east of Cape Agulhas. Ann. S. Afr. Mus. 64: 251-336.
Louw, E. 1977. The South African Museum’s Cruises on the R/V Meiring Naude off the east
coast of southern Africa. 1. Station data, 1975, 76. Ann. S. Afr. Mus. 8: 147-159.
STEBBING, T. R. R. 1906. Amphipoda 1. Gammaridea. Tierreich 21: I-XXXIX, 1-806.
SEXTON, E. W. 1909. Notes on some Amphipoda from the north side of the Bay of Biscay:
Families Pleustidae and Eusiridae. Proc. zool. Soc. Lond. 1909: 848-879. .
STEPHENSEN, K. 1944. Crustacea Malacostraca, VIII: (Amphipoda IV). Danish Ingolf-Exped.
3: 1-51.
WALKER, A. O. 1897. On some new species of Edriophthalma from the Irish Seas. J. Linn. Soc.
26: 226-232.
6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must.be placed in parentheses if a
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Synonymy arrangement should be according to chronology of names, i.e. all published
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order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845
Figs 14-15A
Nucula (Leda) bicuspidata could, get: O Sil
Leda plicifera A. Adams, 1856:
Laeda bicuspidata Hanley, 1859: ae: pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
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In describing new species, one specimen must be designated as the holotype; other speci-
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Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
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Biological Abstracts.
CHARLES GRIFFITHS
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 6
AMPHIPODA
4 \/ =a
VOLUME 74 PART 5 JANUARY 1978 | ISSN 0303-2515
MUS. adit ZOOL.
LIPRARY
APR 2 281078
HARVARD
UNIVERSI ry
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Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Koun, A. J. 1960a. Ecological notes on Genus (Mollusca: Gastropoda) i in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, eee masses and feral development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 51.
THIELE, J. 1910. Mollusca: B. Calveieechece? Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band .
January 1978 Januarie
Rant ee Deck
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 7
MARINE ISOPODA
By
BRIAN KENSLEY
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 7
MARINE ISOPODA
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 21 figures and 1 table)
[MS. accepted 9 August 1977]
ABSTRACT
Excluding the Anthuridea, twenty-one species of isopods are recorded from deep water
off the Natal coast, including fourteen new records. The latter include one new genus,
Spinarcturus, and ten new species. These are Microarcturus biserialis, M. youngi, Neastacilla
longispina, Spinarcturus natalensis, Excirolana bicornis, Cirolana caeca, Serolis brinki,
Haploniscus gernekei, Eurycope glabra, and Ilyarachna wolffi. Five endemic South African
species are also recorded, as well as Bathycopea typhlops, premouyy only known from the
coast of Ireland.
CONTENTS
PAGE
Introduction : : ; : VTS
Species list . , : é Rape Me
Systematic disenssion! ; : ; 128
General discussion . : : Wee | 'S)5)
Acknowledgements . ; 4 ee boy)
References . 4 : ‘ : geri,
INTRODUCTION
The present paper forms part of a series, based on material collected by
the South African Museum during the 1975, 1976 cruises off the Natal coast on
the R/V Meiring Naude. For the scope of, and the background to this pro-
gramme, as well as for all station data, the reader is referred to Louw (1977).
The Anthuridea, of which about seven species were collected, are not dealt
with, and will form the subject of a future paper.
The following abbreviations are used throughout this paper: SAM—South
African Museum catalogue number; SM—Meiring Naude Station numbers;
TL—total length; ovig.—ovigerous; juv.—juvenile.
125
Ann. S. Afr. Mus. 74 (5), 1978: 125-157, 21 figs, 1 table.
126
ANNALS OF THE SOUTH AFRICAN MUSEUM
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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Ail
SPECIES LIST
Material not identified to specific level is either damaged or immature.
SM
Station 3d ojo) ovig. 22 Juvs
SUBORDER VALVIFERA
Family Arcturidae
* Microarcturus biserialis sp. nov. ; ; , OY 3 — <= =
Microarcturus oudops (Barnard) . : : . 103 2 4 4 —
* Microarcturus youngi sp. Nov. . : ; an 86 12 5 3 +50
103 8 10 — 37
* Neastacilla longispina sp. nov. . ; : 5 &© 2), — 3 3
* Spinarcturus natalensis gen. et sp. nov. . 5 ti) 4 DD, 3 D
103 ol — — 2
SUBORDER FLABELLIFERA
Family Cirolanidae
*Excirolana bicornis sp. nov. ‘ : : 5 68 4 6 7
*Cirolana caeca sp. nov. ‘ ; ; : eS 2 4 4 —
Cirolana imposita Barnard . ; ‘ : AG 1 — —
23 1 — — —
Family Sphaeromatidae
*Bathycopea typhlops Tattersall . ' : 5 108 2 3 1 —
Family Serolidae
*Serolis brinki sp. nov. . Sy Pe a, : 5» 108 2 3 1 —
Family Cymothoidae
Nerocila sp. . : : : : : : 5 OF — — — 1
SUBORDER GNATHIIDEA
Family Gnathiidae
Gnathia sp. . : : : : ‘ : 5 sh) +20 + 20 +20 +20
SUBORDER ASELLOTA
Family Stenetriidae
*Stenetrium abyssale Wolff . : : : . 86 1 — 1 —
Stenetrium dalmeida Barnard : : ; a SO 1 —- — —
Stenetrium dagama Barnard
Family Haploniscidae
* Haploniscus gernekei sp. nov. . : 3 . 86 3 3 — —
103 9 4 — —-
Family Ischnomesidae
Ischnomesis sp. j : i : : ; 4 6 2 fragments
Family Eurycopidae
*Furycope glabra sp. nov. . , : ‘ ny 60) — 1 = —
103 1 — — —
Family Dendrotionidae
* Acanthomunna spinipes (VanhOffen) . P eX) 4 8 3 —
103 y) 1 — —
Family Ilyarachnidae
*Ilyarachna wolffi sp. nov. . : : : TP SO 4 _- _ ==
103 2 — — —
Ilyarachna sp. : ; : é f 3 . 103 — Z, — —
*New Record
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
SYSTEMATIC DISCUSSION
Family Arcturidae
Microarcturus biserialis sp. nov.
Fig. 2
Description
Body elongate, slender, widest at pereionites II and III; head fused with
pereionite I, anterior margin concave, anterolateral corners acute, eyes lacking,
one pair small submedian tubercles anterior to submedian pair dorsal spinose
processes; lateral margin of head incised. Coxae of pereionites II and III acute,
of pereionites III to VII with two or three smaller points. Each pereionite
possessing dorsal submedian pair of spinose processes and broader apically
acute lateral process. Pleon consisting of three fused segments plus pleotelson,
former each marked by a pair of submedian spines; pleotelson apically acute,
with acute lateral tooth at about midpoint.
Antennule not reaching midpoint of second antennal peduncle segment.
Three distal segments of antennal peduncle becoming progressively longer.
Mandibles, maxillae and maxilliped typical of the genus.
Pereiopod I shorter than following pereiopods, with numerous serrate
spine-like setae.
Pereiopods II to IV slender, all segments with elongate simple setae on
ventral surface.
Pereiopods V to VII more robust, dactyli, propodi, carpi, meri and distal
part of ischia with dense ventral pile of short setules, propodi and carpi with few
fringed ventral spines.
Penial process with distal third of rami separate, tips truncate.
Pleopod 1 with endopod about half length of exopod, distally rounded,
bearing simple setules; exopod apically curved, with only two distal fringed
setae; about twenty simple spines on outer margin, broad furrow on anterior
face. Pleopod 2 rami similar, distally rounded/truncate, stylet on endopod only
slightly longer than ramus, proximally stout, tapering distally, with small spines
on inturned distal flanges.
Material
Holotype SAM-A15467 1 3 TL5,9 mm SM 109 28°41,0’S 32°36,8’E
1 300 m
233 TL5,9 mm SM 109
Remarks
The structure of the male pleopods | and 2, the mouthparts, and pereional
appendages leave no doubt that the present species belongs to the genus
Microarcturus, in spite of the lack of eyes and the relatively short exopod of
pleopod 1. The dorsal sculpturing resembles that found in the males of
M. ornatus Kensley, 1975, and M. similis (Barnard), but neither of these has
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 129
Boe
F
Fig. 2. Microarcturus biserialis sp. nov.
A. 3 dorsal view. B. @ lateral view. C. Pleopod 1 g. D. Maxilliped. E. Pleopod 2 3.
F. Pereiopod I. G. Pereipod II. H. Pereiopod VII. I. Penis.
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
the double row of spines as well developed on the last three pereionites.
The specific name ‘biserialis’ is derived from the double row of prominent
dorsal spines on the pereion and pleon.
Microarcturus oudops (Barnard)
Fig. 3
Neoarcturus oudops Barnard, 1914: 214, pls 18c, 19b; 1920: 397; 1940: 508. Nordenstam,
[YB Ils.
Previous records
Off Cape Point.
Material
SAM-A15472 2 gg 492 4 ovig. 22 SM 103
Remarks
Barnard (1914) based the new genus Neoarcturus on several features,
including the non-geniculate body, the non-elongate fourth pereionite, distinct
coxae, pleon of four fused segments, and the antennal flagellum of three articles.
All these features apply equally to Microarcturus. Kensley (1975) characterized
Fig. 3. Microarcturus oudops (Barnard) Pleopod 1 3.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 131
Microarcturus by incised lateral margins of the head, dorsolateral eyes, first
pereionite completely fused to head, two or three pleonal segments anterior to
the pleotelson, and the first pleopod of the male with the inner ramus more than
half the length of the outer. All these features are found in M. oudops. Further,
although Barnard characterized Neoarcturus as being blind and possessing three
pairs of oostegites, four pairs of oostegites are present, as are eyes; these latter
are, however, very weakly pigmented. There is thus no justification for placing
oudops in a genus separate from Microarcturus. The relative length of the inner
ramus of the first pleopod of the male cannot be used as a criterion for generic
separation as this shows a range from short in M. biserialis to fairly long in
M. youngi. The first pleopod of this species is refigured.
Microarcturus youngi sp. nov.
Fig. 4
Description
Female. Body with numerous small tubercles; widest at pereionites II
and III. Head with anterolateral corners subacute, with submedian ridges.
Eyes large, lateral. Coxae indistinct, of pereionite I ventrally directed, tridentate;
coxae of pereionites II and III expanded laterally, convex in outline; coxa of
pereionite IV rectangular. Pereionites V to VII shorter and narrower than
preceding ones. Pleon consisting of one indistinct and two distinct segments
plus pleotelson. Latter broad, terminally subacute, with rounded lateral tooth.
Antennule reaching to midpoint of second antennai peduncle segment.
Antennal peduncle 4-segmented, two basal segments together equal in length to
third segment, latter slender, slightly shorter than fourth segment; flagellum of
two articles with slender terminal spine.
Mandibles, maxillae and maxilliped typical of the genus.
Pereiopod I shorter than following pereiopods, with numerous serrate
spine-like setae.
Pereiopods II to IV relatively slender, with numerous slender setae.
Pereiopods V to VII more robust, propodi, carpi and meri with dense very
short setae on ventral surface plus several widely spaced spines, those on
propodus barbed.
Male. Body parallel-sided, cylindrical, pereionites varying in sculpture
from granular to almost smooth. Pleon with pair of large subconical processes
at base.
Penial rami fused for three-quarters of length, tips diverging. Pleopod 1
basis with rounded proximal flange, and fourteen peg-like processes; inner
ramus shorter than outer, distally rounded/truncate; outer ramus with apex
produced and bent, numerous spines on outer margin, broad furrow on anterior
face, ten plumose setae distally.
Pleopod 2 with stylet of endopod stout, cylindrical, slightly convoluted
apically.
ANNALS OF THE SOUTH AFRICAN MUSEUM
1132
Li,
wit”
Fig. 4. Microarcturus youngi sp. nov.
A. 2 dorsal view. B. 3 lateral view. C. Penis. D. Maxilliped. E. Pleopod 1 g. F. Antennule.
G. Pereiopod VII. H. Pereiopod II. I. Pleopod 1 ¢.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 133
Material
Holotype SAM-A15465 1 ovig. 2 TL 5,8 mm SM 86 = 27°59,5’S
32°40,8’E 550m
Allotype SAM-A15465 1¢ TL6,0mm SM 86
12 gs 592 3ovig. 92 SO0juvs SM 86
SAM-A15466 8 gg 1092 37juvs SM 103
Remarks
M. youngi is obviously closely related to the group of species including
M. ornatus Kensley, M. dayi Kensley, and M. similis (Barnard), in which the
basic plan and sculpture of the males and females are similar. Each, however,
may be distinguished by details of the sculpture. The most distinctive feature of
M. youngi is the very prominent pair of bosses at the base of the pleotelson in
the male.
The species is named for Mr D. Young, first officer of the R/V Meiring
Naude.
Neastacilla longispina sp. nov.
Figs 5-6
Description
Female. Body slender, elongate. Head and first pereionite fused, fusion
indicated by shallow furrow; eyes large, lateral; pair of dorsal spines and few
Fig. 5. Neastacilla longispina sp. nov.
A. Ovigerous @ lateral view. B. ¢ lateral view.
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
small tubercles present. Pereionites I and II with small dorsal spine and spinose
lateral process. Pereionite II with single very strong dorsal spine, smaller spine
between dorsal spine and spinose lateral process. Pereionite IV elongate, with
pair of strong submedian dorsal spines at midpoint, and single slightly hooked
median dorsal spine on posterior margin. Pereionite V with few small tubercles
and spinose coxa. Pleon consisting of three fused pleonites plus pleotelson,
segments indicated by shallow furrows; spinose lateral process on third pleonite.
Pleotelson with distal half tapering, apically acutely rounded. Antennule
consisting of 3-segmented peduncle, basal segment with small rounded dorsal
process; flagellum of single article reaching beyond basal antennal segment.
Antenna longer than the total body length, basal segment short, with
spinose distal process; following three segments very elongate, slender, flagellum
Vines
Fig. 6. Neastacilla longispina sp. nov.
A. Pereiopod VII. B. Pereiopody (Ge Pereiopod II. D. Pleopod 2 g. E. Antennule 3.
F. Pleotelson 3.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 135
short, of three articles, terminal article very short.
Mouthparts typical of the genus.
Pereiopod I shorter than following pereiopods, dactylus distally rounded/
truncate, with strong terminal serrate spine; all segments bearing numerous
simple setae; serrate spines also present on dactylus and propodus.
Pereiopods II to IV slender, elongate, dactyli absent, remaining segments
bearing elongate simple setae.
Pereiopods V to VII becoming shorter posteriorly, more robust than
preceding pereiopods, dactyli with two short apical claws, numerous fine setules
on all segments except basis, especially concentrated on ventral surfaces.
Male. Pereionite IV more elongate than in female; head and pereionites
with few scattered tubercles; no elongate dorsal spines as in female. Antennular
flagellum relatively longer than in female.
Antenna with second peduncular segment with small scattered tubercles.
Pleotelson with slight point at lateral midpoint.
Pleopod | with rami of equal length, with elongate distal plumose setae;
exopod with three elongate proximal setae, endopod with proximal indentation.
Pleopod 2 rami of equal length, with distal elongate plumose setae;
endopod with stylet at base of inner margin extending well beyond rami,
apically acute and slender.
Material
Holotype SAM-A15459 1 3 TL10,5 mm SM 86 27°59,5’S 32°40,8’E
550 m |
Allotype SAM-A15459 19° TL8,3mm SM 86
Ig 292) 3 yuvs SM 86
Remarks
The present species bears little resemblance to the three species of Neastacilla
hitherto recorded from South African waters, but most closely resembles
Astacilla attenuata Hale, 1946, from New South Wales. N. Jongispina differs
from the Australian species especially in possessing dorsal elongate spines on
the pereion, and in having more elongate antennae.
The specific name is derived from the dorsal spines of the female.
Spinarcturus gen. nov.
Diagnosis
Arcturid possessing a geniculate body; first pereionite not fused with the
head; dorsolateral eyes; antennae shorter than the total body length; fourth
pereionite not elongate; anterior four pairs of pereiopods unspecialized, not
differentiated from the posterior three pairs; pleopod | of the male relatively
unspecialized, not furrowed on anterior face; stylet of pleopod 2 of the male
apically simple.
Type species of the genus Spinarcturus natalensis.
The generic-name is derived from the numerous spines of the body.
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
Spinarcturus natalensis sp. nov.
Figs 7-8
Description
Male. Body elongate, geniculate, with numerous short fine setules
entrapping debris. Head with concave anterior margin, not fused with
pereionite J. Latter bearing pair of slender spines. Pereionites II to IV with two
pairs of dorsolateral spines. Pleon consisting of three fused segments, first
pleonite unarmed, second pleonite with one pair dorsolateral spines, third
pleonite with pair strong proximolateral spinose processes, rest of segment
smooth, convex, terminally bluntly rounded. Coxae not dorsally visible. Eyes
dorsolateral.
Antennule 4-segmented, terminal article elongate, bearing aesthetascs.
Antenna shorter than body, peduncle consisting of two short proximal
segments and two more elongate distal segments, plus flagellum of three articles,
slender terminal spine present; all segments with scattered setae. Mandible with
tridentate incisor, tridentate lacinia, four setae in row, broad truncate molar
process with four stout spines on ventral margin. Maxilla | with three fringed
setae on inner ramus, several simple spines on outer ramus.
Maxilla 2 with both lobes of outer ramus tipped with two sparsely fringed
Fig. 7. Spinarcturus natalensis gen. et sp. nov.
A. 3 dorsal view. B. Pleopod 1 g. C. Pleopod 2 3. D. Penis.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 137
setae, inner ramus with about ten fringed setae.
Maxilliped with several fringed setae on four distal palp segments and on a
endite.
Pereiopod I shorter than following pereiopods, dactylus with strong
terminal spine, smaller secondary spine, and several fringed spines, propodus
with many fringed spines; carpus, merus, and ischium with few ventral fringed
spines and numerous fine short setules.
LIZZ Ss
Za.
Fig. 8. Spinarcturus natalensis gen. et sp. nov.
A. Pereiopod VII. B. Maxilla 2. C. Mandible. D. Uropod apex. E. Pereiopod I. F. Maxilla 1.
G. Maxilliped. H. Antennule. I. Antenna.
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pereiopods II to IV essentially similar to pereiopods V to VII, latter with
more elongate propodi and carpi. Perelopods more or less covered with fine
short setules and adhering debris.
Rami of penis united almost to apex, latter tapered, finely setose.
Pleopod | relatively unspecialized, with proximal indentation in endopod,
both rami apically rounded, with numerous plumose setae.
Pleopod 2 both rami apically rounded, with plumose setae; stylet of
endopod sabre-shaped, almost twice length of ramus, apically acute.
Uropod with strong ridge near outer margin, outer ramus small, triangular,
inner ramus less than half width of outer, with two subapical spines.
Female. Similar to male, but pereionites II to IV wider; broodpouch
formed by four pairs of oostegites.
Material
Holotype SAM-A15473 1 3 TL 6,0 mm SM 86 27°59,5’S 32°40,8’E
550 m
Allotype SAM-A15473 1 ovig. 9 TL 8,2 mm SM 86
SOS Z2ZLQ Lowe 2ws SMS
SAM-A15474 1g 2juvs SM 103
Remarks
Deciding on the generic position of the present species gives rise to some
difficulty. Pereionite IV is not elongate, which rules out Astacilla, Neastacilla,
Arcturella, Arcturina, Arcturopsis, Parastacilla and Arcturinoides. The head is
not fused with the first pereionite, which eliminates Pleuroprion, Antarcturus
and Jdarcturus. The antenna is shorter than the body, and has a flagellum of
three articles, which rules out Antarcturus, Arcturus, and Dolichiscus. Pleopod 1
in the male is not specialized or furrowed as in Austroarcturus, Microarcturus,
Antarcturus and Holidotea. The elongate stylet of the second pleopod of the
male is reminiscent of Arcturinoides and Arcturina but does not have a trifid
apex as in these latter. The first pleopod with its proximal indentation again
resembles that of Arcturina and Arcturinoides. The pereiopods, however, are not
differentiated as they are in these genera. The creation of a new genus for this
material would thus seem to be warranted.
The specific name is derived from the Province of Natal.
Family Cirolanidae
Excirolana bicornis sp. nov.
Figs 9-10
Description
Male. Body width about half length, widest at pereionites II and VI. Head
with dorsolateral eyes; two elongate slightly curved apically rounded ‘horns’
anterior to eyes, length of ‘horns’ varying with total length. Frontal lamina
distally rounded, dorsally visible between antennal bases, proximally narrowed.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 139
—————>——.
}
\)
Ze
Fig. 9. Excirolana bicornis sp. nov.
A. 6 dorsal view. B. Mandible. C. Maxilla 1. D. Maxilla 2. E. Maxilliped.
F. Head of ¢ in lateral view.
Clypeus acutely pointed in lateral view. Pereionites unsculptured except for
faint, impressed line. All coxae pointed, those of pereionites IV to VII being
elongate-acute. Pleotelson triangular, distal margin dentate, apically acute.
Antennal peduncle 3-segmented, third segment elongate; flagellum of eight
to nine articles.
Antenna longer than antennule, peduncle 4-segmented, terminal segment
longest, flagellum of fourteen articles.
Mandible with 3-segmented palp, middle segment nearly three times length
- of basal segment; incisor 4-dentate; setal row of fourteen to sixteen short
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
spine-like setae; molar process with numerous small teeth on upper surface.
Maxilla 1 with three setae on inner ramus, ten dentate and simple spines on
apex of outer ramus.
Maxilla 2 with several plumose setae on inner ramus, inner lobe of outer
ramus with five setae, outer lobe with three setae.
Maxillipedal endite narrow, armed with five plumose setae and single
coupling hook; palp 5-segmented, third segment longest and broadest, all
segments setose.
Pereiopod I with sensory spines on ventral margin of propodus, carpus and
merus, carpus very short.
Pereiopod VII armed with clusters of serrate spines on propodus, carpus,
merus and ischium.
Fig. 10. Excirolana bicornis sp. nov.
A. Antenna. B. Antennule. C. Pereiopod I. D. Pereiopod VII. E. Pleopod 2 d.
_ F. Apex of telson. G. Uropod.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 14]
Pleopod 2 with stylet attached to base of endopod, extending well beyond
both rami, apically tridentate.
Uropod with inner ramus broader than outer, margins serrate, apically
narrowed, with several setae and few spines on outer margin.
Female. Similar to male, but lacking ‘horns’ on head.
Material
Holotype SAM-A15469 1 3 TL (excl. horns) 5,6mm SM 86 27°59,5’'S
32°40,8’E 550 m
Allotype SAM-A15469 1 @ (with oostegites) 5,5imm SM 86
Paratypes SAM-A15469 3 gg 592 7juvs SM 86
Remarks
Sexual dimorphism in Excirolana has been recorded only in one other
species, E. kumari Bowman, 1971, from Malaysia. In this species the male bears
a pair of triangular horns at the dorsal posterolateral corners of the head, two
slender lateral horns on pereionites I and II, plus a mediodorsal spine on
pereionite II.
The distinctive pair of horns anterior to the eyes on the head of the male
(from which is derived the specific name), immediately separates E. bicornis
from E. kumari. ;
Cirolana caeca sp. nov.
Figs 11-12
Description
Body width slightly more than half length, strongly convex dorsally, widest
at pereionites IV and V, coxae well developed, leaf-shaped; no dorsal sculpture
other than impressed line on pereionite VII. Pleon consisting of five segments
plus pleotelson, lateral margin of first pleonite overlapped by coxae of last
pereionite; lateral margin of fifth pleonite overlapped by fourth; pleotelson
wider than long, distally rounded, distal margin crenulate.
Eyes absent.
Antennule shorter than antenna, third peduncular segment longest,
flagellum of five articles.
Antenna with terminal peduncular segment longest, flagellum of eleven
articles.
Mandibular palp 3-segmented, middle segment longest, bearing simple and
plumose spines; incisor of four broad teeth; setal row of ten curved spine-like
setae; molar process with marginal row of teeth.
Maxilla 1 inner ramus short, with three apical plumose setae; outer ramus
with several dentate and simple spines.
Maxilla 2 inner ramus with five plumose setae, inner lobe of outer ramus
with four simple setae, outer lobe with three.
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxillipedal endite short, with one coupling hook and four plumose setae;
palp 5-segmented, middle segment broadest and longest.
Pereiopod I shorter than following pereiopods, carpus short and triangular,
ventral margins of propodus, carpus, and merus bearing few sensory spines.
Pereiopod VII with serrate spines on ventral and distal margins of segments.
Penial processes digitiform.
Pleopod 2 male with stylet on inner ramus longer than rami, apically with
slender spine and subterminal blunt tooth.
Uropod with both rami distally narrowly rounded, inner ramus broad,
with sparse plumose setae.
Fig. 11. Cirolana caeca sp. nov.
A. ¢ dorsal view. B. Maxilliped. C. Maxilla 1. D. Maxilla 2. E. Mandible.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 143
Material
Holotype SAM-A15468 1 ¢ TL 3,9 mm SM 78 27°31,6’S 32°50,0’E
VSO) tom
Allotype SAM-A15468 1 ovig.2 4,2 mm SM 78
Paratypes SAM-A15468 1 3 TL 3,2 mm 5 99 TL 2,8 mm-4,0 mm
3 ovig. 22 TL 3,6 mm-4,0 mm SM 78
Remarks
This very distinctive species differs markedly in body shape from other
blind species of Cirolana, such as C. californiensis Schultz and C. poissoni
Monod.
The specific name is derived from the lack of eyes.
Fig. 12. Cirolana caeca sp. nov.
A. Antenna. B. Antennule. C. Pleopod 2 3. D. Uropod. E. Apex of telson.
F. Pereiopod I. G. Pereiopod VII.
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Sphaeromatidae
Bathycopea typhlops Tattersall
Fig. 13
Bathycopea typhlops Tattersall, 1905: 12, 65, pl. 3. Loyola e Silva, 1971: 216, figs 2-3. Schultz,
IO7Bs LIS
Previous record
Off County Kerry, Ireland.
Material
SAM-A15461 2 33 399 lovig. 2 3,9-5,3 mm SM 103
Remarks
This is only the second record of this remarkable species. Tattersall (1905)
recorded B. typhlops from five stations off the Irish coast, in depths ranging from
about 400 to 830 metres, and in all cases the bottom substrate was fine sand.
The length of the adult male and female from Ireland was 5,0 mm, agreeing well
with the present material. No differences in proportions or in the appendages
could be detected between Tattersall’s and Loyola e Silva’s figures and
descriptions, and the present specimens. The huge distance between these two
records can only be explained by a lack of sampling of the appropriate substrate.
Fig. 13. Bathycopea typhlops Tattersall, dorsal view.
Family Serolidae
Serolis brinki sp. nov.
Figs 14-15
Description
Male. Body longer than wide, lacking spines and tubercles on dorsum.
Head with anterolateral angles acute, slightly produced, anterior margin
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 145
Fig. 14. Serolis brinki sp. nov.
medially emarginate, eyes entirely absent. Coxal plates marked off by sutures
on pereionites II to IV, coxal plate of pereionite VII not extending beyond
pleural plates of second and third pleonites. Pleotelson with very slight rounded
median ridge, more strongly indicated proximally, becoming obsolete distally;
small lateral point at uropodal insertion. Antennular peduncle longer than
flagellum, latter of twenty articles.
Antennal peduncle about equal in length to entire antennule, flagellum of
fourteen articles.
Left mandible with lacinia tridentate, right mandible with incisor somewhat
expanded, with two strong spines and rounded tubercles; lacinia process distally
dentate.
Maxillipedal palp 3-segmented, distal segment well developed.
Pereiopod I, palm of propodus with alternating closely packed slender bifid
and broad finely setulose setae; distal margin of carpus with two short setae and
few fine setules.
Pereiopod II with double row of eight stubby, bristled setae on palm, distal
margin with several simple setae.
Pereiopod VII slender, elongate, with setae on all segments except basis.
Pleopod 2 with distal portion of endopodal stylet about three times longer
than proximal portion.
Uropod with exopod well developed, oval, shorter than endopod.
Colour pattern (when alive): only anterior half of dorsum pigmented
red-grey; posterior half white. Pigment present on antennal and antennular
peduncles, with white dapples on head and terga. Head and anterior four
pereionites with broad hyaline margins.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
Holotype SAM-A15460 1 3 TL 8,6mm SM 103 28°31,7’S 32°34,0’E
680 m
Remarks
This blind species of Serolis closely resembles two previously recorded
deep-water species, viz. S. vemae Menzies, 1962, from the North and South
Atlantic, and S. menziesi Hessler, 1970, from the South Atlantic off Brazil, but
differs from both.
Fig. 15. Serolis brinki sp. nov.
A. Inner view of mandibles. B. Maxilliped. C. Pleopod 2 ¢. D. Pereiopod I.
E. Spines on palm of propodus, pereiopod I. F. Pereiopod II. G. Pereiopod VII.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 147
S. menziesi does not have a lateral spine on the pleotelson, the uropodal
exopod is minute and the pleotelson possesses a triangular flattened proximal
area.
In S. vemae the uropodal exopod is also much smaller than in S. brinki, the
pleotelson also possesses a flattened proximal area, but lacks a lateral spine.
S. brinki is the first species of the genus recorded in South African waters,
and is named for Dr C. van der Merwe Brink, President of the South African
Council for Scientific and Industrial Research.
Family Stenetriidae
Stenetrium abyssale Wolff
Stenetrium abyssale Wolff, 1962: 25, figs 1-5.
Previous records
Kermadec Trench (north-east of New Zealand) and Tasman Sea, 4510-
4540 m.
Material
SAM-A15470 1 ovig.2 5,3mm 13 3,0mm SM 86
SAM-A15471 12 5.4mm 4 g¢ 5,0 mm-5,8 mm SM 103
<
Remarks
Wolff (1962) gives the dimensions of 9,9 mm TL ¢ and 8,6 mm TL Q.
The present specimens, including an ovigerous female, are all less than 6 mm in
length. This is the only difference from the Kermadec material, the appendages
and body shape agreeing exactly with Wolff’s description.
Family Haploniscidae
Haploniscus gernekei sp. nov.
Fig. 16
Description
Body about twice longer than wide. Anterolateral margins of pereionites
and pleotelson with broad hyaline border. Lateral margin of pleotelson
continuous with pereion. Distal margin of pleotelson convex, distolateral
corners not produced beyond medial margin. Rostrum acute, slightly upturned,
with small tubercle on each side of base; frontal margin excavate on either side
of rostrum. Lateral margin of pereionite IV squarely truncate. Pleopod | in
male with rami fused, distal margin straight, with seven setae on each side,
distolateral portion tapering to narrowly rounded apex. Uropods dorsally
visible, extending almost to level of medial pleotelsonic margin.
Material
Holotype SAM-A15454 1 ¢ TL 2,8 mm SM 103 28°31,7’S 32°34,0’E
680 m
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
Allotype SAM-A15454 192 SM 103
8 gg 3 22 SM 103
SAM-A15455 3 33 3°22 SM 86
Remarks
H. gernekei differs from H. tricornis Menzies, 1962, which it closely
resembles, in the antennular flagellum (six articles in H. tricornis, five in
H. gernekei, and the relative lengths of the articles differing); the first pleopod
of the male, the distolateral angles being more produced than in H. tricornis;
in the uropod extending at least to the level of the medial pleotelsonic margin
or slightly beyond, and in lacking a lateral ridge on the dorsum of the pleotelson.
The species is named for Mr D. Gerneke, previously of the South African
Museum, for his invaluable assistance on both the Meiring Naude cruises.
E
Fig. 16. Haploniscus gernekei sp. nov.
A. Holotype dorsal view. B. Maxilliped. C. Antennule. D. Apex of pleopod 1 ¢.
E. Pleopod 2 3.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 149
Family Eurycopidae
Eurycope glabra sp. nov.
Figs 17-18
Description
Body glabrous, head and first four pereionites narrower than rest of body.
First four pereionites narrow, anterolateral angles acute; pereionite V broader
than VI, lateral corners rounded; pereionite VII wider than VI, distal margin
slightly concave. Pleotelson of single segment, broader than long, distally
broadly rounded, uropods just visible dorsally. Rostrum distally rounded, with
eight setae.
Basal antennular segment broad, with inner distal angle produced, second
segment narrow; flagellum of about fifteen articles.
Mandibular palp with third segment strongly rounded, with several setae
and fringed spines; molar process broad, truncate, with five marginal setae.
Maxilla | with several simple curved spines distally, inner ramus with single
strong distal seta.
Maxilla 2 slender, with several elongate distal setae. Epipodite of maxilliped
with strong rounded process on outer margin; three basal palp segments broad,
second segment longest, with two spines on outer distal angle; two distal
segments narrow, setose; endite narrower than basal palp segments.
Pereiopod I very slender, remaining pereiopods missing.
B
Fig. 17. Eurycope glabra sp. nov.
A. Holotype dorsal view. B. Pleopod 1 g. C. Maxilliped. D. Epipod of maxilliped.
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 18. Eurycope glabra sp. nov.
A. Left mandible. B. Maxilla 1. C. Maxilla 2. D. Antennule. E. oe 2 3. F. Uropod.
G. Rostrum. H. Pereiopod I.
THE SOUTH AFRICAN MUSEUM’S MEIJRING NAUDE CRUISES 15]
Pleopod | in male narrow, elongate, rami distally rounded, with sub-
terminal process on outer margin.
Pleopod 2 with outer margin strongly convex.
Uropod biramous, inner ramus four and a half times longer than outer.
Operculum of female with strong median crest, forming spine-like process
proximally.
Material
Holotype SAM-A15452 1 ¢ TL 3,6 mm SM 103 28°31,7’S 32°34,0’E
680 m
Allotype SAM-A15453 192 TL 3,3 mm SM 60 27°09,6'S 32°58,2’E
800-810 m
Remarks
From Wolff’s key (1962: 144) to the species of Eurycope the present species
would seem to resemble £. antarctica Vanhoffen, 1914, but this species does not
have acute antero-lateral corners of the first four pereionites, and the pleotelson
is not distally as broadly rounded; further, the uropodal exopod is relatively
longer, the rostrum not as narrow as in the present material.
The specific name derives from the completely smooth dorsum, entirely
lacking in sculpture.
Family Dendrotionidae
Acanthomunna spinipes (Vanhoffen)
Fig. 19
Mormomunna spinipes Vanhoffen, 1914: 569, figs 100, 101.
Acanthomunna spinipes: Menzies, 1962: 174. Wolff, 1962: 65.
Previous records
Antarctic (Gauss Station 8.1I.1903) 385 m.
Material
SAM-A15463 4 g¢ TL2,5mm 8 99 3 ovig. 92 TL4,4mm SM 86
SAM-A15464 2 gg 12 SM 103
Remarks
The presence of a uropodal peduncle, pereionites I to III separate, the head
not fused with pereionite I, maxilliped palp segments all slender, the presence
of a mobile lacinia in the left mandible, an apically truncate molar process,
coxae on pereionites II to VII, an antennular flagellum of at least six articles,
pereiopods with a single claw, and dorsolaterally inserted uropods, all place
the present specimens in the family Dendrotionidae. The presence of eyes
indicate the genus Acanthomunna (Wolff, 1962: 65). The present material agrees
almost exactly with Vanh6ffen’s description of A. spinipes, especially in the
structure of the first and second pleopods of the male. While the generally
spinose pereion and pleon, and the elongate antennae and pereiopods also
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 19. Acanthomunna spinipes (Vanhoffen).
A. Dorsal view. B. Maxilliped. C. Pleopod | 3. D. Pleopod 2 3. E. Right mandible.
F. Maxilla 1. G. Maxilla 2. H. Operculum 9.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 153
agree well, unfortunately the spectacularly spinose uropods of A. spinipes are
absent in all the Natal material.
In one specimen, the base of the uropodal peduncle was still in situ. The
relatively large insertion for the uropods and the sturdy remnant mentioned
indicate a massive uropod of the type figured by Vanhoffen.
Family Ilyarachnidae
Ilyarachna wolffi sp. nov.
Fig 20
Description
Body glabrous, tapering posteriorly. Head with transverse convex portion
and broad lateral areas. Pereionite I not as broad as II, both laterally acute.
Pereionites II to IV increasing in length, coxae of pereionites posterior to II
rounded; posterior margin of pereionite V concave; VI triangular, VII narrower.
Pleotelson of two segments, terminal segment longer than broad, distally
narrowly rounded.
Antennular basal segment broad, outer distal angle somewhat produced,
second segment narrow; flagellum of about fifteen articles.
Mandibular palp slender, 3-segmented, middle segment with two fringed
spines, shorter terminal segment with four spines; incisor process a single
rounded tooth; molar process narrow, with three terminal setae; strong ridge
on inner face of mandible.
Maxilliped with second palp segment very broad and long, third segment
triangular, two distal segments short and narrow; endite less than half width of
second palp segment, epipodite broad, distally rounded.
Pleopod | in male curved, rami distally narrowly rounded, setose.
Uropods missing.
Material
Holotype SAM-A15457 1 3 TL 2,9 mm SM 86 27°59,5’S 32°40,8’E
550 m
3 g3 SM 86
SAM-A15458 2 33 SM 103
Remarks
Three species closely resemble present material. (See Wolff 1962: 94.)
I. triangulata Menzies, 1962, is very similar in body proportions but lacks the
acute coxae of the first two pereionites, the mandibular palp is also very different.
I. affinis Barnard, 1920, has acute coxae on the first four pereionites, rather
than just the first two.
I. crassipes Barnard, 1920, has rounded coxae of the first two pereionites,
while the pleon is about as long as wide.
The species is named for Dr Torben Wolff, for his contributions to isopod
biology and taxonomy.
154
ANNALS OF THE SOUTH AFRICAN MUSEUM
ZZ =
Sree
m—
Fig. 20. I/yarachna wolffi sp. nov.
A. Holotype dorsal view. B. Pleopod 1 g. C. Maxilliped. D. Antennule. E. Maxilla 1.
F. Maxilla 2: G. Mandible. H. Pleopod 2 dg.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 155
TIlyarachna sp.
le, ZI
Description
Body markedly spinose. Head with eight spines. Pereionites I to IV with
two rows of small spines and several lateral spines; pereionites V to VII with
single row of spines. Pleotelson of two segments, terminal segment with several
small granules plus single strong pair of spines.
Material
SAM-A15462 299 TL3,0 mm SM 103
Remarks
As the uropoda and all the pereiopods are lacking from both specimens,
specific status is not given, even though a species of //yarachna with a double
row of small spines on the anterior four pereionites does not seem to have been
recorded. This species is obviously related to such spinose species as
I. spinosissima Hansen, I. argentinae Menzies, I. gurjanovae Menzies, and
I, multispinosa Menzies
GENERAL DISCUSSION
Of the 21 species of isopod recorded, 15 were taken from either station
SM 86 or SM 103, 7 species being common to both stations. The nature of the
bottom at both stations was very similar, and as 9 of the 15 species caught are
new, it may fairly be said that this is a habitat not previously sampled. A
Fig. 21. Ilyarachna sp. °.
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
comparison of the two stations (Table 1) and the isopods caught at them will
give some idea of the characteristics of this ecotype. The differences in the catch
between the stations can to some extent be explained by the difference in fishing
ability of the gear used. The heavy dredge, with its 100 cm by 30 cm mouth,
would be able to pick up much larger organisms and rocks than could the light
biological dredge (see Menzies 1962, fig. 1) with its narrow 100 cm by 10 cm
mouth. Thus Alcyonaria and Gorgonacea with attendant isopods such as
Neastacilla would be caught only by the heavy dredge, while some of the smaller
debris-dwelling isopods of the upper few centimetres of substrate such as
Eurycope, Serolis and Microarcturus would be caught by the light dredge, but
perhaps lost through the wider mesh of the heavy dredge.
TABLE |
A comparison between two stations and the isopods caught at them.
SM 86 SM 103
Depth 550 metres 680 metres
Gear Heavy dredge Light biological dredge
Bottom Fine hard mud, overlain with hetero- Fine hard mud, overlain with hetero-
pod and pteropod shells, coral pod and pteropod shells, coral
fragments, numerous foraminifera, fragments, numerous foraminifera,
sponge fragments and spicules sponge fragments and spicules
Non-Isopod Hydroids, gorgonaceans, solitary Hydroids, small echinoderms, pycno-
organisms corals, glass sponges, pycnogonids, gonids, amphipods, small crabs,
caught small crabs, cumaceans, amphipods, _— small gasteropods and bivalves.
ostracodes, polychaetes, small
gasteropods and bivalves
Isopods Acanthomunna spinipes Acanthomunna spinipes
caught Anthurids Anthurids
— Bathycopea typhlops
Excirolana bicornis —
— Eurycope glabra
Gnathia sp. =
Haploniscus gernekei Haploniscus gernekei
Ilyarachna wolffi Ilyarachna wolffi
— Ilyarachna sp.
Microarcturus youngi Microarcturus youngi
= Microarcturus oudops
Neastacilla longispina —
— Serolis brinki
Spinarcturus natalensis Spinarcturus natalensis
Stenetrium abyssale Stenetrium abyssale
Stenetrium dalmeida —
Stenetrium dagama Stenetrium dagama
The affinities of this fauna are not easily apparent, especially with 52 per
cent being described as new. The presence of a Serolis, and Stenetrium abyssale
and Acanthomunna spinipes indicate possible Antarctic/subantarctic affinities,
while five endemic species (Microarcturus oudops, Stenetrium dagama, Cirolana
imposita, and two easily recognizable anthurids Leptanthura laevigata and
Mesanthura catenula) show the expected South African component. The
unexpected presence of Bathycopea typhlops can only be explained by the
paucity of collecting in this type of habitat, especially off the west African coast.
THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Sy
ACKNOWLEDGEMENTS
I should like to thank the crew of the R/V Meiring Naude and my colleagues
of the South African Museum who assisted in the collecting of the material
described here. I am grateful to Dr T. E. Bowman of the Smithsonian Institution,
and Prof. J. H. Day of the University of Cape Town, who read the manuscript
and made many useful criticisms and suggestions.
REFERENCES
BARNARD, K. H. 1914. Contributions to the Crustacean fauna of South Africa. 3. Additions
to the marine Isopoda, with notes on some previously incompletely known species.
Ann. S. Afr. Mus. 10: 325-442.
BARNARD, K. H. 1920. Contributions to the Crustacean fauna of South Africa. 6. Further
additions to the list of marine Isopoda. Ann. S. Afr. Mus. 17: 319-438.
BARNARD, K. H. 1940. Contributions to the Crustacean fauna of South Africa. 12. Further
additions to the Tanaidacea, Isopoda and Amphipoda, together with keys for the
identification of the hitherto recorded marine and freshwater species. Ann. S. Afr. Mus.
32: 381-543.
Bowman, T. E. 1971. Excirolana kumari, a new tubicolous isopod from Malaysia. Crustaceana
20: 70-76.
HALE, H. M. 1946. Isopoda—Valvifera. Rep. B.A.N.Z. antarct. Res. Exped. 4: 163-212.
HESsLeR, R. R. 1970. A new species of Serolidae (Isopoda) from bathyal depths of the
equatorial Atlantic Ocean. Crustaceana 18: 227-232.
KENSLEY, B. 1975. Marine Isopoda from the continental shelf of South Africa. Ann. S. Afr.
Mus. 67: 35-89.
Louw, A. E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station Data
1975, 1976. Ann. S. Afr. Mus. 72: 147-159.
LoyYoLa E SILVA, J. 1971. Sobre os generos Ancinus Milne Edwards, 1840 e Bathycopea
Tattersall, 1909, da colecao U.S. Nat. Mus. Archos Mus. nac., Rio de J. 54: 209-223.
MENZIES, R. J. 1962. The isopods of abyssal depths in the Atlantic Ocean. Vema Res. Ser. 1:
84-206.
NorDENSTAM, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae,
Arcturidae, Parasellidae, and Stenetriidae mainly from the South Atlantic. Further zool.
Results Swed. Antarct. Exped. 3 (1): 1-284.
SCHULTZ, G. A. 1973. Ancinus H. Milne Edwards in the New World (Isopoda, Flabellifera).
Crustaceana 25: 267-275.
TATTERSALL, W. M. 1905. The marine fauna of the coast of Ireland. Part V. Isopoda. Scient.
Invest. Fish. brch Ire. 1904 (2): 1-90.
VANHOFFEN, E. 1914. Die Isopoden der Deutschen siidpolar-Expedition 1901-1903. Dt. Siidpol.-
Exped. 14: 447-598.
Wo rr, T. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota.
Galathea Rep. 6: 1-320.
- nd
ed
7 _
ee
_
Sara ive
a auc
eee
haan!
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
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, it not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
BRIAN KENSLEY
THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PART 7
MARINE ISOPODA
VOLUME 74 PART 6 JANUARY 1978 | | ISSN 0303-2515
MUS. COMP. ZOOL.
| LIBRARY
APR 28 1978
HARVARD
UNIVERSITY
ug 22 ap NM
CAPE TOWN
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(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction
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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.:
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Note: no comma separating name and year
Dagination indicated by colon, not p.
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et al. in text for more than two joint authors, but names of all authors given in list of references.
(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
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For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
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number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 1960b. Spawning. behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
January 1978 Januarie
Part 6 ~~ Deel
Bb
owing NS
THE GEOGRAPHICAL DISTRIBUTION OF
SOUTHERN AFRICAN HYDROIDS
By
N. A. H. MILLARD
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town
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Price of this part/Prys van hierdie deel
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE GEOGRAPHICAL DISTRIBUTION OF
SOUTHERN AFRICAN HYDROIDS
By
N. A. H. MILLARD
South African Museum, Cape Town
(With 9 figures, 8 tables and 2 appendices)
[MS. accepted 9 August 1977]
ABSTRACT
The geographical distribution of 251 species of hydroid Cnidaria round the coast of
southern Africa is analysed by means of a system of radial sectors. It is concluded that there is
no distinct west coast fauna as such, but that a boundary zone exists at approximately 31°S
latitude between an east coast region with mainly tropical affinities and a temperate west—plus—
south coast region with a reduced tropical component and a large endemic population. The
main centre of the endemic population is the Agulhas Bank. The relationships of the tropical
species are mainly with the Western Indian Ocean Province and the Indo-West-Pacific Region.
Deep-water species (45) which occur below 400 m are of a mixed nature and show no clear
relationship with other parts of the world. Deep-water species are classified into stenobathic/
eurybathic and stenothermic/eurythermic categories, and their distribution relative to water
temperatures is discussed.
CONTENTS
PAGE
Introduction : ‘ : : ; i 5 ; sn LESS)
Methods . : ; ‘ : : t j E nel LOO
Faunal boundaries. é i ; : : . 164
The tropical component . ; ; 5 : ; 169
The temperate component : ; : : : wat tel ©
The endemic component . ; F ; ‘ G : 173
The cosmopolitan and scattered components ; ‘ 174
Distribution and depth . ; : ; K ; ae tS
Estuarine and brack-water species . : 2 : mel 9
Conclusion . : : 5 ; ‘ ; : ‘ : 182
References . : : ; ; k : ; oh LSS)
Appendix 1 F : : : : 5 4 : 4 185
Appendix 2 : ; : q : , i 5. IBS
INTRODUCTION
This paper is an attempt to analyse the distribution of the hydroid fauna of
the coast of southern Africa as documented in a recent monograph (Millard
1975). Since the latter went to press a few more taxonomic papers on, or lists of,
hydroids have appeared, namely Arnaud ef al. (1976), Beurois (1975), Calder
(1975, 1976), Cooke (1975), Cornelius (1975a, 1975b), Mergner & Wedler
(1977), Millard & Bouillon (1975), Millard (1977a) and Watson (1975). Among
159
Ann. S. Afr. Mus. 74 (6), 1978: 159-200, 9 figs, 8 tables, 2 appendices.
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
these are records which extend the range of some South African species and they
have been taken into account in the analysis which follows, as also have records
from Buchanan (1957), Michel (1974), Blanco (1967, 1968, 1973, 1974) and
Blanco & Bellusci de Miralles (1972a, 19726), which have only recently become
available to the author.
Within the borders of southern Africa, here taken to be the 20° parallel of
south latitude, several new distribution records from deep water off the Natal
coast and some new records for the country were published by Millard (19775).
There are also scattered identifications by the author, as yet unpublished, which
are listed in Appendix 1.
The following changes in name have been adopted here:
Antennella africana Broch, 1914, to A. quadriaurita Ritchie, 1909, fide Millard
CIS Vilo2 123).
Eulaomedea calceolifera (Hincks, 1871) to Campanularia calceolifera Hincks,
1871, fide Cornelius (1975a: 254).*
Lafoea fruticosa (M. Sars, 1851) to Lafoea dumosa (Fleming, 1820) fide Cornelius
(I97Sbe 385).
Obelia bicuspidata Clarke, 1875, to O. bidentata Clarke, 1875, fide Cornelius
(1975a: 260).
Any zoogeographical analysis is aimed at pinpointing population changes
within an area, and, if possible, relating them to physical or chemical factors
and determining relationships with other areas. Such an analysis is subject to
certain unavoidable weaknesses—the assignment of species to categories or
components is largely subjective; numbers of species within categories are
liable to change as more records are forthcoming (for instance some of the
South African endemics may in time be expected to turn up outside our
boundaries); and numbers of species may be influenced by different intensities’
of collecting in different areas or by misidentifications. The longer we wait the
more reliable will be the results, but results, if only interim ones, are needed
now. The larger the number of species involved the more meaningful will be the
conclusions. One must bear in mind that it is the main trends which are
significant rather than the details.
METHODS
The first step in the analysis was to remove all doubtful records and
incomplete identifications. This left a total of 251 species.
The first objective was an analysis of distribution within the borders of
southern Africa, and for this some system of comparison was necessary. When
dealing with the littoral zone it is a comparatively simple matter to convert the
distance along the coast to a straight line, a method with obvious practical
* Cornelius pointed out that the type species of Campanularia is Sertularia verticillata
Linnaeus, 1758, and not Sertularia volubilis Linnaeus, 1758, as previously thought (Millard
1975: 203). Since S. verticillata is a branched species the diagnosis of Campanularia given
previously (Millard 1975: 203) must be altered to include branching forms, and Eulaomedea
sunk in Campanularia.
16]
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS
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162 ANNALS OF THE SOUTH AFRICAN MUSEUM
advantages, and this has been done by Stephenson (1947), Stephenson &
Stephenson (1972), Day (1967) and others. However, when dealing with the sub-
littoral area as well, this is clearly not possible for a continent of the shape of
southern Africa. After much consideration the following arbitrary method was
devised, in which the coastal waters were divided into thirty-six sectors of
approximately equal area (Fig. 2).
The central point for the sectors was chosen as the intersection of the
latitude line of 20°S and the longitude line of 24°E, a point which is approxi-
mately mid-way between the west and east coastlines. From this central point
radii were drawn at 5° intervals, starting with one through the Cape Peninsula,
and then working outwards in both directions. The sectors thus constructed
were numbered 1-36. The reason for placing one radius (Number 14/15)
through the Cape Peninsula is that Cape Point is generally recognized as an
approximate zoogeographical boundary between the cold Benguela water on
the west and the warmer Agulhas water on the east, and might be expected to
have some significance. It so happens that another radius (Number 15/16)
conveniently passes through Cape Agulhas, the most southerly point of Africa,
and a third (Number 23/24) through the boundary between Transkei and Natal.
ae
DG
/
ES
Ox 15° 20° 25° 30° 35° 40
20)
35
40
Fig. 2. Map of southern Africa illustrating the sector system which was used to analyse the
distribution.
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 163
It must be emphasized that this is an arbitrary method, and obviously
could not be applied to all coastlines, but it is felt that it might form a useful
working method and basis for comparison with other southern African marine
groups. It has the advantage that each sector covers an approximately equal
area of sea and ‘straight-line’ length of coast and represents a cross-section
through all depths. Since the sectors are roughly normal to the coast and the
currents tend to follow the coast, each sector is affected in a similar way and
population differences related to water temperature should be highlighted. It is
felt that the advantages of this method at least outweigh the disadvantages (and
impracticabilities) of the ‘straightening-out’ method of Stephenson.
The second objective was an analysis of the world-wide distribution of the
southern African species, and for this the species were delegated to components
defined as follows:
TROPICAL
Circumtropical. Present in the tropics of all three oceans, but may spread
north or south into warm temperate seas as well.
Indo-West-Pacific Region. Stretching from the Suez Canal across the Indian
Ocean and the East Indies into the Western Pacific as far as Hawaii and
Easter Island. Includes northern Australia, Madagascar and East Africa.
May spread into warm temperate seas as well.
Western Indian Ocean Province. The western section of the Indo-West-
Pacific Region, bounded in the north by the Persian Gulf, and including
Madagascar and surrounding islands.
TEMPERATE. (In a trial analysis it was found impossible to differentiate between
cold temperate and warm temperate fauna when assessing world distribution.)
Antitropical (bipolar). Present in the temperate zones of all three oceans,
and present in both the Northern and the Southern hemisphere, but not
in the tropics except perhaps just at the edge.
Southern. Restticted to the temperate waters of the Southern hemisphere,
but often in the subantarctic islands as well, and rarely reaching the
Antarctic.
Atlantic. Restricted to the northern and/or southern temperate zones of
the Atlantic, but absent from the tropics. May spread further north into
the subarctic.
ENDEMIC
OTHER
Cosmopolitan. Present in all three oceans, and occurring in both tropical
and temperate seas.
Scattered. Present in various regions, but not sufficiently widespread to
be included in Cosmopolitan. Does not fit any other category.
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
A list of species, together with their components and distribution within
thirty-four sectors, is given in Appendix 2. (Sectors 35 and 36 have been omitted
as no records were available from this region.)
FAUNAL BOUNDARIES
It has long been recognized that the marine fauna of southern Africa
includes both tropical elements, predominating on the east coast, and temperate
elements, predominating on the south and west coasts. Most authorities,
following Stephenson (1947) and Stephenson & Stephenson (1972), have
recognized a separate east coast (tropical or subtropical) fauna, a south coast
fauna (warm temperate) and a west coast fauna (warm or cold temperate) for
the littoral region. These earlier views are summarized by Briggs (1974), who
includes the east coast in the Western Indian Ocean Province of the Indo-West-
Pacific tropical region; the south coast in the Agulhas Province of the South
African warm temperate region; and the west coast as a separate South West
Africa Province of the South African warm temperate region. Opinions have
differed as to the position of the boundary between the east and south coast,
and whether the west coast represents a separate faunal province, and if so
whether it should be regarded as cold temperate or warm temperate.
The distribution of the faunal components of the continental shelf and the
extent of the faunal regions is largely dependent on the ocean currents round
the South African coast, in general the warm southward-flowing Agulhas
Current being responsible for the southerly extension of tropical forms on the
east coast, and the cold northward-flowing Benguela Current being responsible
for the more northerly distribution of cool-water forms on the west coast. It is
not intended to discuss these current systems in any detail here since accounts -
are available in most textbooks of marine biology (a concise general account is
given by Stephenson & Stephenson (1972) and more recent contributions are
summarized by M. J. Penrith (1976)).
In order to determine whether there is any region of the coast where a
marked change in the hydroid fauna occurs, the range of the tropical, temperate
and endemic components was plotted separately (Fig. 3). (The range of species
was used in preference to presence/absence, in order to minimize differences in
collecting intensity.) This figure emphasizes the tropical nature of the east coast
fauna, for there is a strong tropical component at the northern end which
gradually drops away, until beyond St Helena Bay on the west coast (Sector 14)
only one species remains for a short distance. The rate of disappearance is
greatest at the boundary of Sectors 23/24, and is also high in the region of
Sectors 13/14/15. The temperate and endemic components are both highest on
the south coast, falling away fairly evenly to the north-east, but more sharply to
the north-west in the region of Sectors 13/14/15. A similar trend appears when
the range of species is plotted as a percentage of the total range (Fig. 4).
It is thus possible to distinguish two ‘zones of change’, though they are not
as marked as might be expected. The first (Sectors 23/24) provides a convenient
NUMBER OF SPECIES
20
10
50
40
30 4
20
10
80
70
60
50
40
30
20
10
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS
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TEMPERATE
ENDEMIC
TROPICAL
5 "10 15 20 “95
SECTORS
«I|nhaca
30
165
Fig. 3. The distribution of the temperate component (above), the endemic component (centre)
and the tropical component (below). From range of species.
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
boundary between the east and south coast fauna. It is located between 30 and
32°S latitude, i.e. the stretch of coast between the Bashee River in the south and
Durban in the north. For the discussions which follow, the boundary between
Sectors 23 and 24 is used (31°S). This is more or less in agreement with con-
clusions drawn from other faunal groups. Stephenson & Stephenson (1972)
suggest a boundary zone between Port St Johns and Qolora, 1.e. at about 32°S,
for the intertidal fauna (the form of the graph of the tropical component in
Figure 3 of this paper is very similar to that for the intertidal fauna in
Stephenson & Stephenson 11972, ie. 8:26); Je 1 Ba Smith G949) and
M.-L. Penrith (1970) suggest the Great Kei River at 32°42’S for the fishes, and
Day (1967) suggests the Bashee River at 32°15’S for the benthic Polychaeta.
TEMPERATE
20 owe
40 0-0
20 °
ENDEMIC :
RANGE
80
60
TROPICAL
PERCENTAGE
40
20
SECTORS
Fig. 4. The distribution of the temperate, endemic and tropical components, from range of
species as a percentage of the total range. Sectors 1-7 have been omitted since the total number
of species is less than twenty and the percentages unrealistic.
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 167
At the boundary of Sectors 23/24 the monthly minimum surface temperature
near the coast does not fall below 16°C, the maximum does not rise above 27°C
and the mean ranges from 21 to 24°C (U.S. Naval Oceanographic Office 1967).
This is approximately the region where the Agulhas Bank begins to widen and
force the warm Agulhas Current offshore. Further south the latter is separated
from the coast by a cooler counter-current flowing northwards.
The usefulness of the second ‘zone of change’ in the south-westerly corner
of the Cape Province (Sectors 13/14/15) is more debatable. It could be used to
divide the temperate region, which stretches from north of 20°S on the west
coast to 30-32°S on the east coast, into two provinces, if warranted by the
fauna.
To test this an arbitrary boundary was placed at Cape Point (Sector 14/15)
and the composition of species on the two sides of it compared with one another
and with the east coast (Table 1). (Cape Point was used as a boundary by
Stephenson (1947, fig. 3), Ekman (1953) and Briggs (1974).) It may be noted
from Figures 3 and 4 that there is no evidence of a change in fauna at Cape
Agulhas (Sector 15/16), the most southerly point of Africa).
TABLE |
Analysis of fauna of west coast, as compared with that of the south and east coasts
(actual records, not range).
East coast
(Sectors 24-36)
West coast South coast
(Sectors 1-14) | (Sectors 15-23)
Component
No. of he No. of VA No. of Sfp
species species species
Tropical 8 9,6 30 19,0 eZ 39,6
Temperate . 13 IS7/ 20 Ora 12 6,6
Endemic to area : : 2 2,4 D)\ 13,3 7 933
Endemic to more than one area 25 30,1 4| D9 30 16,5
21 1353 28 15,4
Tes) 15,8 23 12,6
Cosmopolitan
Scattered
Total
It is apparent from Table 1 that the west coast fauna consists mainly of
endemic, cosmopolitan and scattered species. The tropical species diminish in
number round the coast from east to west; only 8 species remain on the west
coast and these are all absent north of St Helena Bay. The temperate species of
the west coast are fewer in number than those of the south coast. Only 2 of them
(Clavopsella nayvis and Gonothyraea loveni, both from Table Bay harbour and
probably introduced by ships), and possibly a third (Sarsia eximia), are
restricted to the west coast. Only 2 rare species are endemic solely to the west
coast (Myriothela tentaculata and Eudendrium ritchiei), whereas 21 are endemic
to the south coast and 17 to the east coast. Of the cosmopolitan and scattered
components only 3 (Acryptolaria crassicaulis, Hybocodon unicus and Hydro-
dendron gracilis) have been recorded from the west coast alone.
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
In total, thus, the west coast has only 8 unique species (9,8 %), of which
only 2 (2,4°%) are endemic to the area. Briggs (1974) requires at least 10 per cent
endemic species for the designation of a separate province, and on this basis
the west coast of South Africa does not qualify, at least for the hydroids. The
hydroid fauna of the west coast seems to be mainly an impoverished south coast
fauna.
Although Day (1967) accepted a separate west coast element for the littoral
polychaets, he could find no marked difference between the south and west
coasts for the sublittoral ones, and ascribed this to the fairly uniform water
temperature on the bottom of the continental shelf (he gave figures of 12—14°C
at 100 m from Port Elizabeth to Liideritz, but the records collated by Hulley
(1972) show a somewhat lower range of 8-13°C at 100 m between 20°S on the
west coast and Port Elizabeth on the south coast). On the other hand there is a
difference in the surface water temperature on the two sides of Cape Point
(Day gives a range of 15-20°C for the south coast and 12—15°C for the west
coast, but under certain conditions and for certain periods the south coast
temperature may drop to as low as 11°C (during local upwelling) and rise as
high as 24°C, while that of the west coast may drop to 9°C (M. J. Penrith 1976)).
In this respect 1t might be noted that all hydroids are sublittoral in the sense
that they cannot withstand drying and survive in the littoral zone only in damp
overhangs or open runnels.
None of this really explains why there should be a diminution of hydroid
species on the west coast and practically no endemic population of cold-
temperate forms. Ekman (1953) associates the poverty of the benthos of the
‘Namaqua coast’ with the hydrogen sulphide resulting from frequent upwelling
and red tides. Briggs (1974) suggests that certain species are prevented from”
spreading westwards round Cape Point by the absence of high summer
temperatures for reproduction (he gives a February mean of 16—-18°C for South
West Africa as against one of 20—22°C for the south coast, but compare remarks
in parenthesis above). M.-L. Penrith & Kensley (1970a) suggest that a region of
minimal temperatures between Hondeklip Bay and Liideritz (i.e. north of
St Helena Bay) acts as a barrier to the establishment of southern warm-
temperate intertidal species further north. The paucity of hydroid records on the
west coast may be partly a reflection of the difficulty of collecting on the exposed
and inhospitable Skeleton Coast. However, M.-L. Penrith (pers. comm.), who
has collected in this area, feels that the poverty is a real one, and perhaps linked
to the absence of suitable habitats such as sheltered coves, rock pools, etc., and
the preponderance of surf beaches and sandy floors. The harsh conditions and
poverty of the rocky-shore fauna are also emphasized by M. J. Penrith (1976).
There is no evidence at all of a southward spread of species from tropical
west Africa. The southern boundary of the west African tropical region is too
far north for consideration in this paper (14 or 15°S: Briggs 1974; 18-22°S:
Hulley 1972). M.-L. Penrith & Kensley (19706) found the first of the west
African littoral species at Rocky Point (18°59’S).
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 169
In conclusion the south and west coast sublittoral areas are considered to
represent a single temperate province for the hydroid fauna, which might be
termed the Agulhas/Namaqua Province, the centre of the endemic population
being on the Agulhas Bank.
THE TROPICAL COMPONENT
Of the total number of 251 species, 77 (30,7 °%%) can be classified as tropical.
Of these, 20 are circumtropical, 24 Indo-West-Pacific and 33 confined to the
Western Indian Ocean. In addition there are 17 endemic species which are
confined to the South African east coast, and therefore with tropical tendencies
(these were termed ‘subtropical endemics’ by Day (1974)). (Appendix 2.)
Ekman (1953) and Briggs (1974) consider that the main centre of origin of
the tropical fauna is the Indo-Malayan Region. From here species tend to
spread west to the Indian Ocean (and may penetrate into the Red Sea or round
southern Africa), and to a lesser extent east to the Eastern Pacific.
Species have been classified here as Indo-West-Pacific or Western Indian
Ocean even though a few have one or two records outside these areas. Thus,
2 have penetrated into the Mediterranean (Corydendrium parasiticum, Thyro-
scyphus fruticosus), 4 have reached the tropical west coast of Africa (Abietinaria
laevimarginata, Campanularia africana, Corydendrium parasiticum, Sertularia
ligulata), | has reached tropical South America (Dentitheca bidentata), and | has
reached southern Australia (Dynamena obliqua). Further spread would produce
a circumtropical distribution, but of the species classified as such no less than
13 have failed to reach the east coast of the Pacific, and this applies also to 4 of
the cosmopolitans (Table 2). There is therefore support for Briggs’s contention
(1974) that the ‘East Pacific barrier’ is one not easily crossed.
There is some evidence for a secondary centre of origin in the Western
Indian Ocean, for in addition to the seventeen South African subtropical
endemics (i.e. present on the east coast only), there are seven Western Indian
TABLE 2
List of circumtropical and cosmopolitan species which
do not occur on the east coast of the Pacific.
Circumtropical Cosmopolitan
Diphasia digitalis Antennella secundaria
Dynamena quadridentata Cladocoryne floccosa
Gymnangium hians Coryne pusilla
Halecium dyssymetrum Halecium sessile
Hydrodendron caciniformis
Idiellana pristis
Monostaechas quadridens
Plumularia strictocarpa
Scandia mutabilis
Sertularella diaphana
Sertularia distans
S. marginata
S. turbinata
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ocean species which, though absent from the Indo-Polynesian area, have yet
penetrated into the Red Sea (C/ytia latitheca, Cytaeis nassa, Diphasia heurteli,
Halopteris glutinosa, Hydractini kaffraria, Plumularia wasini, Solanderia
minima).
Some communication between southern Australia and the Western Indian
Ocean is indicated, since there are five species common to the two, namely
Amphisbetia maplestonei, Crateritheca acanthostoma, Halopteris glutinosa,
Sertularella arbuscula and Synthecium dentigerum. These occur nowhere else
(except for H. glutinosa which also occurs in the Red Sea) and the likelihood is
that the movement has been from west to east with the West Wind Drift.
THE TEMPERATE COMPONENT
The total temperate component includes 28 species (11,2%), of which
11 are antitropical (bipolar), 10 are restricted to the Southern hemisphere and
7 are restricted to the Atlantic (Appendix 2). These are species which cannot
survive the high temperatures of the tropics, although some can spread into the
cooler waters of the subpolar regions. The southern species must have evolved
in the south, but the antitropical and Atlantic species may have evolved in
either the south or the north and must at some time have crossed the tropics.
This may have been achieved by equatorial submergence, for Briggs (1974)
points out that the tropical surface water is only about 30-40 m deep in the
Eastern Atlantic. It may also be due to transportation on the hulls of ships.
In all, twenty-one species have been recorded on ships’ hulls or floating objects
such as buoys. The fauna of ships’ hulls is a characteristic group of hardy
species (Millard 1952), most of them cosmopolitan. Three of the species (all.
belonging to the temperate component) are known only in harbour areas
in South Africa, and it is suggested that these are recent introductions by
ships:
Gonothyraea loveni is well established in Cape Town docks, occurring
abundantly on pylons, cables, etc., as well as on hulls of ships. It has not been
reported from any other part of South Africa. Elsewhere it is known from the
North Atlantic, New Zealand, Tasmania and the Argentine.
Campanularia calceolifera has been found, and reproducing, in Cape Town
docks on a barge and on a raft, neither of which had left the harbour. It must,
therefore, be established, though not common. It is not known for certain
elsewhere in South Africa, a record from False Bay by Stechow (1925) being of
infertile material and thus subject to doubt. The species is well known on
European and Mediterranean coasts and also occurs on the east coast of North
America.
Clavopsella navis is known from a ship’s hull in Cape Town docks, and also
from the Kiel Canal (Thiel 1962 as C. quadranularia). The latter is probably the
original locality since the colonies there were more abundant and better
developed.
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 17]
Since all these three species occur both in the South and North Atlantic
they must have traversed the tropics, though they have not established them-
selves in tropical latitudes so far as is known. They may have been hardy
enough to survive a brief passage through warmer waters on ships’ hulls, or
they may have died back during transport, leaving the hydrorhiza to regenerate
again in suitable temperature conditions. Ralph (1961) also found evidence of
the introduction of hydroids by ships’ hulls to New Zealand harbours.
It is interesting to compare the relations of the temperate species with the
other two southern continents. From Table 3 it can be seen that South Africa
has 9 species in common with southern Australasia alone, but only 3 species in
common with South America alone. There are 7 species common to all three.
TABLE 3
Temperate species common to the southern continents.
South Africa and South South Africa and temperate Temperate part of all
Australasia South America three southern
continents
Kirchenpaueria triangulata Phialella turrita Amphisbetia minima
Nemertesia ciliata Sertularella striata _ Filellum antarcticum
Parascyphus simplex Symplectoscyphus paulensis Gonothyraea loveni
Plumularia obliqua Nemertesia cymodocea
P. spinulosa Plumutaria filicaulis
Sarsia eximia Staurocladia vallentini
Sertularella annulaventricosa Tubularia larynx
Stereotheca elongata
Symplectoscyphus macrogonus
(Total: 9) (Total: 3) (Total: 7)
This would suggest dispersal eastward from the Agulhas Bank. Beurois (1975)
emphasized the role of eastward drifting kelp in the dispersal of attached forms
to St Paul and Amsterdam Islands, and Arnaud et al. (1976) discussed the
transport of South African fauna to St Helena on kelp. If the list in Table 3 is
extended to include the cosmopolitan and scattered components and those
tropical forms which have spread south of the Tropic of Capricorn, the relation-
ship between South Africa and Australasia is more marked (34 species in
common with Australasia alone, 10 with South America alone, and 26 in all
three).
The distribution of those South African species which have antarctic or
subantarctic affinities is given in Table 4. In all, 41 species are listed, a relatively
small number (16,3 % of the total), and of these half (21) are cosmopolitan and
this tends to mask the relationships. Without the cosmopolitan species only 20
(8,0°%) have antarctic affinities and of these 11 are classified as temperate. The
absence from South Africa of certain common antarctic genera, such as
Staurotheca, Tulpa, Grammaria and Silicularia, is very striking. On the whole
there is very little evidence of relationship with the Antarctic.
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE 4
Distribution of species with antarctic/subantarctic affinities. The Magellan Region includes
Gauss Station, the Falklands and South Georgia. The Kerguelen Group includes Marion,
Prince Edward and Crozet Islands.
Be 2
ae 3 2
| Seen
Species Component 2 e=| c = §
gee se 8
=e) ps) ae 5 An
a 24 ee ees
Acryptolaria conferta Cosmopolitan x x
A. crassicaulis Scattered x x
Aglaophenia p. pluma Cosmopolitan < x
Amphisbetia minima Temperate (Antitr.) < x
A, operculata Cosmopolitan x x x
Antennella quadriaurita Scattered x
A. secundaria Cosmopolitan x
Bimeria vestita Cosmopolitan x
Bougainvillia macloviana Scattered x <
Campanularia crenata Cosmopolitan x
C. integra Cosmopolitan x
Clytia hemisphaerica Cosmopolitan x x
C. paulensis Scattered x x
Coryne pusilla Cosmopolitan x
Dynamena cornicina Cosmopolitan x<
Filellum antarcticum Temperate (Southern) x XK
F. serratum Cosmopolitan x x
Halecium beanii Cosmopolitan x
H. delicatulum Cosmopolitan x x x
H. tenellum Cosmopolitan x x x x
Hincksella echinocarpa Temperate (Southern) x
Hybocodon unicus Scattered x
Kirchenpaueria triangulata Temperate (Southern) x
Lafoea benthophila Scattered x x x x
L. dumosa Cosmopolitan x x x
Modeeria rotunda Cosmopolitan x x
Nemertesia cymodocea Temperate (Southern) x
N. ramosa Scattered x
Obelia bidentata Cosmopolitan x x x
O. dichotoma Cosmopolitan x x < x
O. geniculata Cosmopolitan x % x x
Parascyphus simplex Temperate (Antitr.) s< x
Phialella turrita Temperate (Antitr.) x
Plumutlaria pulchella Seattered x
P. setacea Cosmopolitan x x
Sertularella leiocarpa Temperate (Southern) ~ S<
S. p. polyzonias Cosmopolitan x s<
S. striata Temperate (Southern) x
Staurocladia vallentini Temperate (Southern) X< <
Symplectoscyphus paulensis Temperate (Southern) S x
Zygophylax armata Scattered Ne
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS WB,
THE ENDEMIC COMPONENT
The number of endemic species strictly belonging to the South African
coast south of 20°S is 76. However, there are 2 species which also occur
immediately north of 20° on the west coast (Gattya humilis and Salacia
articulata). 4 species occur only in South Africa and the Vema Seamount
(Halopteris pseudoconstricta, Salacia articulata, Sertularella flabellum and
S. megista), and it was shown by Millard (1966) that the hydroid fauna of the
Vema Seamount is essentially South African (21 of the 23 species also occur in
South Africa). If these 6 species are included, the endemics number 82 (32,7 °%
of the total of 251). It might be noted, however, that Mergner & Wedler (1977)
have recorded Eudendrium deciduum and Sertularella natalensis from the
Red Sea with a query.
As a comparison the following percentages of endemics have been given
for other large marine groups in southern Africa. The figures are not strictly
comparable, however, since the boundaries used for the “Southern African
Region’ are not always the same.
Polychaeta . 107, (Deyn oy)
Echinoderms, other than Holothuria 48% (Clark & Courtman-Stock 1976)
ASCidians |=) eres | 60 7 ra Mullan nlO7)
Bishi sy ee ee 25s A (ME Ivin Snntinel 970; 380Vontvot
1 500 species)
The south coast of southern Africa has more endemic species of hydroid
than any other region, and the Agulhas Bank seems to be the centre of distri-
bution of the endemic population (Figs 3-5).
There are only three genera endemic to South Africa, Bicorona, Hydro-
corella and Uniscyphus, and all of these are monotypic. However, certain genera
have a larger proportion of endemic species than others (Table 5). These are
probably genera in which speciation is actively occurring in the South African
area. They are also genera in which many of the species are not easily dis-
tinguished from one another, and thus pose constant problems to the systematist.
Similarly, species with several subspecies are ones likely to be undergoing active
speciation. Thus, Thecocarpus flexuosus has four subspecies; the nominal one is
TABLE 5
List of South African hydroid genera with the largest number of endemic species and/or
subspecies.
Number of endemic species Total number of species
Genus or subspecies or subspecies in South Africa
Cladocarpus 9 13
Corhiza 5 5)
Halopteris 4 6
Hydractinia 3 3
Sertularella 14 23
Zygophylax 4 7
NUMBER OF SPECIES
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
WEST SOUTH EAST
West only
West + south
AO South only
30
20
10 South+ east
O
East only
West + south+ east
30
SECTORS
Fig. 5. An analysis of the distribution of the endemic species, with the total number of species
distributed between six components, as in Stephenson (1972). From range of species.
tropical Indo-West-Pacific and the other three are endemic to southern Africa.
Sertularella polyzonias also has four subspecies; the nominal one is cosmopolitan
and two of the others are endemic to southern Africa.
Several common genera, on the other hand, have no southern African
endemics: Aglaophenia, Amphisbetia, Dynamena, Lytocarpus, Nemertesia,
Obelia and Sertularia.
THE COSMOPOLITAN AND SCATTERED COMPONENTS
The cosmopolitan species number 28 (11,2°%%). They owe their wide
distribution to the fact that they are eurythermic, hardy and adaptable. The
wide distribution possibly indicates a very old history, in which there has been
plenty of time to disperse. |
Species classified as scattered or disjunct number 36 (14,3 %). Most of them
NUMBER OF SPECIES
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 175
have odd distributions which do not fit any of the other categories. There are
two possibilities. Either these are newly evolved eurythermic species which are
in the process of spreading rapidly and likely to become cosmopolitan, or they
are ‘peripheral relicts’ of fairly primitive forms which were once more widely
distributed.
Among the scattered species three are interesting because outside southern
Africa they are limited to the west and east coasts of the tropical Atlantic
(Clytia hummelincki, Zyzzyzus solitarius) or to the western tropical Atlantic
only (Cladocarpus tenuis). Three others have a similar distribution but also
extend further north and/or east (Aglaophenia latecarinata, A. pluma dichotoma,
Symplectoscyphus amphoriferus).
Cosmopolitan and scattered species with antarctic/subantarctic affinities
are included in Table 4.
DISTRIBUTION AND DEPTH
Figure 6 shows that there are more species present in the shallow water
(0-100 m) than at any other level, and although there have been fewer samples
from deep water it is not expected that more extensive collecting will alter the
picture to any great extent. The shallow water is obviously the optimum zone
for hydroids. Most of the species in the littoral area are at the upper edge of
180
160
140
120
100
80
60
40
20
© 100.2250 500 750 1000 1250 1500
DEPTH (m)
Fig. 6. The distribution of species according to depth. From range of species.
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
their depth range and able to survive only under overhangs and in rock pools
and crevices. Only 24 of the 98 littoral species have not been found below low
water of springs.
Deep-water species are those which occur over the edge of the continental
shelf, and Ekman (1953) places an arbitrary upper boundary for the deep-sea
fauna at 400 m. The number of hydroids known from below this depth round
the South African coast is relatively small (45 = 17,9%), and only eight of these
reach | 000 m (abyssal). The species are listed in Table 6.
These species can be divided into two arbitrary categories on the basis of
their depth range both in South Africa and the rest of the world:
Stenobathic: those that are restricted to water deeper than 200 m.
Eurybathic: those that occur above and below 200 m.
These two categories are distributed between the different geographical
components as follows:
Stenobathic Eurybathic Total
bropicall, smc psihae pes we ee Cae 5 5 10
Temperate . l 4 5
Endemic 5 6 1]
Cosmopolitan 0 2) 9
Scattered I 9 10
Mkovtalt nee Feu? Poe eh teenie i 12 33 45
All the (12) stenobathic species are in reality cold-water species frequenting
water below about 16°C (Ekman 1953, table 42), and this includes five so-called
‘tropical’ species. These species are thus also stenothermic. Their limiting factor
for distribution may be depth or temperature or a combination of both.
Some of the eurybathic species are also stenothermic. This applies to the
following tropical and scattered species whose records within the tropics are
never shallow, and which are thus restricted to cold water:
Most shallow record in tropics (m)
Acryptolaria rectangularis (tropical) 228
Cladocarpus tenuis (scattered) 185
Cryptolarella abyssicola (scattered) 4 560
Symplectoscyphus amphoriferus (scattered) 161
Zygophylax sibogae (scattered) Only one tropical record:
depth not given.
To the same category of eurybathic/stenothermic species might be added
the endemic and temperate eurybathic species, at any rate those restricted to the
south-west coast of South Africa and the east coast below 200 m:
Cladocarpus sinuosus (endemic): minimum east coast record 400 m.
Hincksella echinocarpa (temperate): minimum record Kerguelen 41 m, minimum
east coast record | 610 m.
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 177
TABLE 6
South African deep-water species.
SB—stenobathic (never recorded above 200 m); EB-—eurybathic (occurring at all depths);
ST—stenothermic (cold-water species); ET—eurythermic (occurring at all temperatures).
Depth range Depthrange Component
Species Component in South outside South _ of depth|
Africa (m) Africa (m) temperature
Acryptolaria conferta Cosmopolitan 64-1200 0-4400 EB/ET
A, crassicaulis Scattered 2835 109-767 EB/ET
A. rectangularis . Tropical 110-1200 228-600 EB/ST
Antennella quadriaurita Scattered 0-825 0-400 EB/ET
Branchiocerianthus imperator Tropical 730 200-5290 SB/ST
Campanularia hincksii Cosmopolitan 9-450 0-811 EB/ET
Cladocarpus distomus . Tropical 292-2200 55-1362 EB/ET
C. dofleini Tropical 425-550 1019 SB/ST
C. inflatus Endemic 495 — SB/ST
C. millardae Tropical 200-595 5020 SB/ST
C. natalensis Endemic 400-900 — SB/ST
C. sinuosus . Endemic 183-680 — EB/ST
C. tenuis Scattered 495 185-550 EB/ST
Clytia gravieri Tropical 0-528 04243 EB/ET
Cryptolarella abyssicola Scattered 100-2740 745-4970 EB/ST
Filellum serratum Cosmopolitan 24—900 0-650 EB/ET
Garveia crassa Tropical 625-900 741 SB/ST
Halecium beanii . Cosmopolitan 0-780 0-1134 EB/ET
H. sessile Cosmopolitan 425-430 0-752 EB/ET
H. tenellum . : Cosmopolitan 0-900 0-730 EB/ET
Peat glutinosa . Tropical 0-550 1-80 EB/ET
H. polymorpha Tropical 0-900 0-89 EB/ET
H. tuba Endemic 11-550 — EB/ET
Hincksella echinocarpa Temperate 1610-2200 41-109 EB/ST
Hydrocorella africana Endemic 0-500 — EB/ET
Kirchenpaueria triangulata Temperate 111-1207 550-585 EB/ST
Lafoea benthophila Scattered 425-430 672-3237 SB/ST
L. dumosa Cosmopolitan 60-920 0—2200 EB/ET
Modeeria rotunda Cosmopolitan 70-550 0-1240 EB/ET
Nemertesia antennina Scattered 425-430 0-1779 EB/ET
N. ramosa Scattered 11-700 0-872 EB/ET
Plumularia prfonbritni Endemic 440 SB/ST
P. mossambicae . Endemic 110-550 — EB/ET
P. setacea Cosmopolitan 0-430 0-752 EB/ET
Sertularella ieecanoas Temperate 200-900 183-672 EB/ST
Stegolaria geniculata . Tropical 775-850 253-910 SB/ST
Symplectoscyphus amphoriferus Scattered 550-900 161-1256 EB/ST
S. arboriformis Endemic 10-420 = EB/ET
S. paulensis . Temperate 347-1200 400-672 SB/ST
Uniscyphus fragilis Endemic 360-420 — SB/ST
Zygophylax africana . Endemic 137-850 — EB/ST
Z. armata Scattered 48-440 183 EB/ET
Z. brownei . Temperate 400-550 20-752 EB/ET
Z. inconstans Endemic 360-450 =< SB/ST
Z. sibogae Scattered 88-900 146-550 EB/ST
Summary: Cosmopolitan . 9
Scattered 10
Tropical 10
Temperate 5
Endemic 11
Total . 45
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
Kirchenpaueria triangulata (temperate): minimum record Agulhas Bank 111 m,
minimum east coast record 350 m.
Sertularella leiocarpa (temperate): minimum record Tristan 183 m, minimum
east coast record 280 m.
Zygophylax africana (endemic): minimum east coast record 400 m.
For the (10) species in this eurybathic/stenothermic category the limiting
factor is obviously temperature rather than depth.
The remaining (23) species are truly eurybathic and eurythermic, for they
can occur in any temperature and at any depth. As might be expected, the
majority (13) are cosmopolitan or scattered.
Hedgpeth (1957) suggested that all the deep-sea benthic population will
eventually prove to be cosmopolitan, but it is clear from this analysis that
geographically cosmopolitan species are not necessarily eurythermic.
The data for a selected group of 21 species were examined to see whether
the depth distribution round the coast bore any relation to changing water
temperatures. These were the species recorded from a minimum of 10 sectors
and from depths extending below 100 m. Only 7.of the species so tested showed
indications of an increasing minimum depth as one passes round the coast from
west to east and then north; only 5 showed indications of an increasing maximum
depth (Fig. 7, Table 7). This might be interpreted as an inability in a few species
to survive in the warmer surface and shallow waters of the east coast. It is not
surprising that none of these species belongs to the tropical component.
Cladocarpus 1s a characteristic deep-water genus, and 12 of the 13 South
African species extend down below 100 m, 9 below 200 m, 3 below 500 m and
7 do not occur in water shallower than 200 m.
TABLE 7
The relationship between depth and distribution.
A. The minimum recorded depth in metres; B. The maximum recorded depth in metres.
Sectors
Species ee ——— Component
0-5 6-10 11-15 16-20 21-24 25-30 31-35
A. Antennella quadriaurita -— 35 0 9 50 ~=100 — Scattered
Halecium delicatulum . — — 0 4 AP MNO 55 Cosmopolitan
Halopteris tuba. : — —— 22 11 Di 70 110 Endemic
Salacia articulata . é 0 25 0 10 0 18 110° Endemic
Sertularella flabellum . — — PS 10 27 124 — Endemic
Sertularella megista . — —— 0 10 DBT 49 110 Endemic
Symplectoscyphus
arboriformis . . 0 10 11 11 40 DA — Endemic
B. Antennella quadriaurita — 40 Te Os Ae SZS — Scattered
Salacia articulata . y 0 23 SO CSS) ses EVaxeleinnvle
Sertularella flabellum . — — 99 200 Dilieuene oe — Endemic
Sertularella megista . — — 139 6 200) N64 219% 347 SEndennic
Symplectoscyphus
arboriformis . ; — 20 35.1120 90 420 — Endemic
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 179
WEST SOUTH EAST
Salacia articulata
DEPTH (m)
Sertularella megista
20 25 30
SECTORS
Fig. 7. The distribution according to depth of three species which show tendencies towards
greater depths in warmer waters. Actual records.
—_
1 ) 10
ESTUARINE AND BRACK-WATER SPECIES
Although many hydroids penetrate into the mouths of estuaries, especially
on rocky substrata, only two species are truly estuarine and have not been found
in the open sea. These are Bimeria fluminalis, which grows on mangrove roots
in Richards Bay and St Lucia estuaries, and Hydractinia kaffraria, which is a
commensal on the shell of Nassa kraussiana and occurs in estuaries from the
Breede River mouth on the south coast to Durban on the east coast.
The Limnomedusan, Ostroumovia inkermanica, is known from several
brack-water lakes on the east coast, the hydroid from Lagoa Poelela, Mozam-
bique, in a salinity of 8 parts per thousand, both hydroid and medusa from
Nhlange Lake, Zululand, in 3—4 parts per thousand, and the medusa alone from
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
A
@ Lovenella chiquitita
@ Cladocarpus leloupi
A Sertularella natalensis
B
Corhiza scotiae
Salacia articulata
Symplectoscyphus macrogonus
Sertularella leiocarpa
Fig. 8A-—D. Distribution of six representative endemic species: A. Lovenella chiquitita
(west coast + False Bay), Cladocarpus leloupi (south coast) and Sertularella natalensis (east
coast). B. Corhiza scotiae (west + south coast). C. Synthecium hians (south + east coast).
D. Salacia articulata (west + south + east coast).
E-F. Distribution of two representative temperate species: E. Symplectoscyphus macrogonus
(west + south coast). F. Sertularella leiocarpa (west + south + east coast), also steno-
thermic.
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 181
A B
Lytocarpus philippinus Dynamena crisioides
D
Sertularia turbinata Sertularella arbuscula
E
Hebella scandens Amphisbetia operculata
Fig. 9A-D. Distribution of four representative tropical species from east coast: A. Lytocarpus
philippinus (to just north of Durban). B. Dynamena crisioides (to East London). C. Sertularia
turbinata (to Mossel Bay). D. Sertularella arbuscula (to Saldanha Bay).
E-F. Distribution of two representative cosmopolitan species: E. Hebella scandens (St Helena
Bay to Mozambique). F. Amphisbetia operculata (Lideritz to Natal).
182
ANNALS OF THE SOUTH AFRICAN MUSEUM
Lake St Lucia North in 10,4-10,8 parts per thousand.
CONCLUSION
The composition of the southern African hydroid fauna is summarized in
Table 8, whence it can be seen that there is a mixture of endemic, tropical,
temperate and cosmopolitan species in that order of abundance.
The presence of a large endemic population, the particular distribution of
some of the Western Indian Ocean tropical species (p. 169), and the apparent
proliferation of subspecies in certain genera (p. 173), all point to an active
evolutionary centre in the South African area.
It is clear that there has also been a heavy invasion of tropical species from
the Indo-West-Pacific region, and these species form the major component of
the fauna of the east coast.
Regarding the temperate and cosmopolitan species, and considering
(1) the low relationship of the South African temperate fauna to that of the
antarctic and subantarctic regions on the one hand, and to that of South
America and Australasia on the other,
(11) the practically certain role of ships in the introduction of at least three
species out of 251 (1,2 %) to South Africa within a period of some 300 years
(p. 170),
TABLE 8
Summary of analysis of geographical components of South African hydroid fauna (from actual
records, not range).
West-plus-south East coast: Whole of
coast: South Africa:
Sectors 1-23 Sectors 24-34 Sectors 1-34
Component (171 species) (182 species) (251 species)
No. Percentage |No. Percentage | No. Percentage
TROPICAL
Circumtropical 10 5,8 18 9,9 20 8,0
Indo-West-Pacific . 7 ARI SS a 23) 12,6 +39,6 | 24 9,6 >30,7
W. Indian Ocean 16 9,4 31 17,0 33 Sil!
TEMPERATE
Antitropical 10 5,8 5 27 11 4,4)
Southern . 8 4,7 ~13,4 5 2 Os 10 4,0 -11,2
Atlantic 5 2,93) 2 ips f 2,8
ENDEMIC
To onearea only .. Bye OSS 17 9,3 e520 4 D0STe)
To the whole of South 38,0 25,8 325]
Africa 20" S15 30F J oliG.s 30 12,0
UNCLASSIFIED
Cosmopolitan Phe NDING 28: PANY Boe wiles
Scattered . 28 - 16.4 (72 wa) | 23) ON ees eee
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 183
(iii) the fact that drifting weed and flotsam has probably performed a similar
role throughout the ages,
it is hardly necessary to evoke the Theory of Continental Drift to explain the
distribution. :
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185
THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS
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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, ‘e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
ae Figs 14-15A
>» Nueula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87. _
Leda‘bicuspidata: Nicklés, ‘1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
: § » < comma/separates author’s name and-year
' » ‘semicolon separates’ more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; ‘. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
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“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
N. A. H. MILLARD
THE GEOGRAPHICAL DISTRIBUTION OF
SOUTHERN AFRICAN HYDROIDS
——
if Fa 0) |
~ VOLUME 74 PART in DECEMBER 1977 ISSN 0303-2515
i
iM . MUS. COMP. ZOOL.
l Lime a — i
NOV 4 94cC°N
A FIV A P)
WMIWaReSi ry
ANNALS
} OUTH AFRICAN
OF THE : MUSEUM
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:
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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)’
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‘As described (Haughton et al. 1927)...’
Note: no comma separating name and year
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author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
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number (only if independently paged) in parentheses, Paeieee (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FIscHER, P.-H. , DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. Zen. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 ~ Band
December 1977 Desember
Part 7 Deel
PAST AND PRESENT DISTRIBUTION OF SOME
RODENT AND INSECTIVORE SPECIES IN THE
SOUTHERN CAPE PROVINCE, SOUTH AFRICA:
NEW INFORMATION
By
D. M. AVERY
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
Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
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OUT OF PRINT/UIT DRUK
il, AO, GO), SID, ZG, 8, hoot), Ses, S, FD),
6(1, t.-p.i.), 771-4), 8, 9(1-2, 7), 10(1-3),
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Price of this part/Prys van hierdie deel
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Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum
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ISBN 0 908407 28 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
PAST AND PRESENT DISTRIBUTION OF SOME RODENT AND
INSECTIVORE SPECIES IN THE SOUTHERN CAPE PROVINCE,
SOUTH AFRICA: NEW INFORMATION
By
D. M. AVERY
South African Museum, Cape Town
(With 3 figures and 2 tables)
[MS. accepted 11 August 1977]
ABSTRACT
Micromammalian material from archaeological sites and modern owl roosts in the Cango
valley and Gansbaai area of the southern Cape Province provides new information concerning
the past and present distribution of certain species of rodents and insectivores. Modern
material from the Cango valley provides evidence of a link between widely separated popula-
tions of several species and shows Praomys natalensis (multimammate mouse) to be co-existing
with P. verreauxi (Verreaux’s mouse). The archaeological evidence indicates that Saccostomus
campestris (Cape pouched mouse) and Steatomys pentonyx (Cape fat mouse) arrived in the
southern Cape only at the beginning of the Holocene about 10000 years BP and that
P. natalensis perhaps entered the Cango valley with the advent of farming in the area around
200 BP. On the other hand, Tatera afra (Cape gerbil), Gerbillurus paeba (pygmy gerbil) and
possibly Otomys unisulcatus (bush Karoo rat) disappeared from the Cango valley some time
after about 1 600 BP.
CONTENTS
PAGE
Introduction : ; sc 20h
Present distribution . ae DOS
Past distribution ; Lo DOF
Conclusion . : : e208
Acknowledgements . . 208
References . i : ware 209
INTRODUCTION
The evidence put forward below became available during the course of a
study of micromammalian remains from archaeological sites and comparable
modern owl roosts in the southern Cape Province. Two areas are considered,
the Cango valley (3322AC) near Oudtshoorn, and the Gansbaai area (3419CB)
on the Walker Bay coast. For the Cango valley (Fig. 2) material was available
from the archaeological site Boomplaas A (33°23’S 22°11’E) which represents
well in excess of 40 000 years of accumulated deposit during the Upper Pleisto-
cene and Holocene (Deacon & Brooker 1976: 211) and from five modern owl
roosts. Two of these, Boomplaas B and C which are adjoining rock-shelters, are
within 100 m of Boomplaas A and in the same cliff. Two others, Nooitgedacht A
201
Ann. S. Afr. Mus. 74 (7) 1977: 201-209, 3 figs, 2 tables.
202 ANNALS OF THE SOUTH AFRICAN MUSEUM
B Praomys verreauxi C Steatomys pentonyx
P. natalensis 5
F Otomys laminatus G Dasymys incomtus
Fig. 1. Distribution of some rodent and insectivore species showing the relationship of the
Cango valley (C) and Gansbaai (G) to existing ranges. A is based on Meester & Lambrechts
(1971) and B and D-G are based on Davis (1974).
DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 203
Fig. 2. The position of the sites in the Cango valley. BPA, BPB and BPC = Boomplaas A,
B and C; NGA and NGB = Nooitgedacht A and B; OG = Osgat. Metric approximations
of 200 ft contours are given.
and B (33°22’S 22°10’E) which are also adjoining rock-shelters, are situated
2,5 km north-west of Boomplaas, whilst the fifth site, Osgat (33°22’S 22°11’E),
lies 1,3 km north-east of Nooitgedacht and 5 km north of Boomplaas. For the
Gansbaai area (Fig. 3) archaeological material was available from Die Kelders 1
(34°32’S 19°22’E), covering a period from about 80000 BP to 45000 BP
(years before present) during the Middle Stone Age (MSA) and from about
2000 BP upwards during the Late Stone Age (LSA) (Tankard & Schweitzer
1974: 367; Tankard 1976: 155), and from Byeneskranskop | (34°35’S 19°28’E)
which was occupied from about 12000 BP upwards (F. R. Schweitzer pers.
comm.) and which lies some 9,5 km south-east of Die Kelders. The modern
sample was taken from Byeneskranskop 2, a rock-shelter situated just below the
archaeological site and set in the same cliff. Total minimum numbers of
individuals represented in the various samples are given in Table 1.
It appears from Davis’s (1974) distribution maps that little collecting or
recording has been undertaken in the two areas discussed here, although
Grindley et al. (1973) collected owl pellets from near Stanford some 15 km north
of Gansbaai. For this reason the recent information has proved useful both for
ANNALS OF THE SOUTH AFRICAN MUSEUM
204
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DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 205
consolidating known ranges and for filling in gaps in others less well known.
Remains from archaeological contexts provide some indication of when various
species arrived in or, in some cases, disappeared from the two areas.
PRESENT DISTRIBUTION
The presence of many of the species listed in Table 1 was predictable and
in these cases the evidence merely adds another locality to an already clearly
established range. In some cases, however, the sparseness of the existing record
adds significance to the present information. Acomys subspinosus (Cape spiny
mouse), Saccostomus campestris (Cape pouched mouse), Dendromus melanotis
(grey pygmy climbing mouse), D. mesomelas (chestnut climbing mouse),
Crocidura flavescens (red musk shrew), C. cyanea cyanea (reddish-grey musk
shrew), Suncus varilla (lesser grey dwarf shrew) and, to some extent, Praomys
verreauxi (Verreaux’s mouse) fall into this category. The evidence confirms the
ranges of A. subspinosus and S. campestris and also the eastward extent of the
range of P. verreauxi as given in Davis (1974: 158, 163, 160 respectively). In the
case of D. melanotis, D. mesomelas (Davis 1974: 166, 168) and C. flavescens
(Meester 1963: 34) confirmation is provided for the existence of links between
populations in the south-west and others in the south-east. For S. varilla the
present information provides not only confirmation of similar linking but also
constitutes a southwards extension of the known range given by Meester &
Lambrechts (1971: 11) (Fig. 1A).
Further to this, and even more important, is another group of species
which has been discovered in the Cango valley. For this group evidence of links
between isolated populations had not previously been forthcoming. Dasymys
incomtus (shaggy swamp rat), Mystromys albicaudatus (white-tailed rat),
Otomys laminatus (laminate vlei rat), and O. saundersae (Saunders’s vlei rat)
comprise this group (Fig. 1D-G). It is interesting to note that Davis (1974:
166 et seq.) predicted that the gap was more apparent than real in several cases.
Most of these species are not frequently caught by owls (Vernon 1972: 119).
Assuming this to represent roughly the actual proportions of animals present,
it is likely that still more records will become available with more intensive
work. Steatomys pentonyx (Cape fat mouse) was also unexpected in the Cango
valley but the situation regarding Steatomys spp. is complicated by the fact that
this genus has long been in need of revision (Davis 1962: 73). Currently the
south-western Cape population is referred to S. pentonyx and the more northern
material to at least two other species. It is thought most likely that the present
material represents a considerable eastward extension of the range of S. pentonyx
(Fig. 1C).
Praomys natalensis (multimammate mouse) has not been thought to occur
further west than about 24°E in the southern Cape, the more westerly area being
occupied by P. verreauxi (Verreaux’s mouse) (Fig. 1B). Indeed, Davis (1974:
160) doubted Pocock’s identification of P. natalensis from Oudtshoorn,
considering it more likely to be referable to P. verreauxi. However, material
ANNALS OF THE SOUTH AFRICAN MUSEUM
206
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par = == =a a = == ame == x x [Iqios AWIsAg pqgand snanjjiqsay
x x x x = — —= — = x x [Iqses oded DAfO DAIDI,
x Xx x x = x a os x x Xx JEL PojIe}-SY MA snjopnorigip sAuossSA Jy
x x x = == == — as x x = osnow yey odeD xduojuad skwuojoajs
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x am = = = x x x x x x Molys jueYyds[q ‘ds snjnjunydayq
x = = = oa = — rae = = —= QJOU! Uspfos 10}U9}}0H Snjojuajjoy snuosdjqup
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(VSD (VSI) (H) (dn)
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— ZYNA pur DINE {BV auoIg 0107) pue (e8y BUOIS OIPPUN) I SIOPIEM AIC = (VSDIAC Pur (VSIN)IAG +7880 = DO +d PUP W yyoepes
MOON = FON Puke WON {0 pure g serjduioog = Odd Pur dda ‘(eusdo]oH) Pue (euEd0IsI=1q Joddq) Vv seEIcHOOT = (H) pue (dn)Vvda
“pole IeeqsUey oY} pue Aa][eA OSURD 9U} UI juasoid 18 pue ysed oY} Ul SUIIINIDO S}USPOI PUB SOIOATIOSUT
J a1avVL
DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 207
collected from the Cango valley by the writer contains examples of both
P. natalensis and P. verreauxi. Not only does this confirm Pocock’s findings, but
it appears to be the first record of the two species’ co-existing. It therefore goes
some way towards establishing the boundary between the two species, for which
Davis (1974: 160) saw a need. This boundary now lies along a line between
Oudtshoorn and Plettenberg Bay but it seems likely that P. natalensis will yet
be found further west in view of its liking for agricultural land, discussed below,
and its highly successful nature.
Davis (1962: 62) remarked that P. natalensis is ‘semi-domestic and present
distribution is possibly dependent on having followed early human population
movements’. It was probably early agricultural activity in particular which
attracted it. Roberts (1951: 472) and Shortridge (1942: 60) note that it is common
near cultivated land and the situation in the Cango valley seems to support this
suggestion. At Boomplaas A microfauna has only been recovered from the
pre-farming levels, the topmost of which is dated to 1 630 + 50 BP (Deacon
et al. 1976: 142). P. natalensis does not occur at all in these levels. It must,
therefore, have arrived subsequently, possibly in the wake of European farmers
who settled there around 200 BP. Not only does P. natalensis occur in significant
numbers in the modern sample, but also there is a change in proportion between
it and P. verreauxi in different parts of the valley (Table 2). At Boomplaas B
TABLE 2
Differences in the proportion of Praomys natalensis and P. verreauxi in the Cango valley.
See Table 1 for explanation of abbreviations.
BPB BPC Total NGA NGB Total
Praomys natalensis. : : : 94 99 193 8 28 36
Praomys verreauxi . ‘ : ; 13 9 22 Di DD 49
and C, which are situated on the hillside above the intensively farmed valley
floor, there is a preponderance of P. natalensis, whereas at Nooitgedacht, in an
uncultivated area, P. verreauxi tends to predominate. Whether the two species
are in competition or whether a state of equilibrium has been reached remains
to be seen. What is clear, however, is that P. natalensis is a recent newcomer to
the area, apparently attracted by cultivated land.
PAST DISTRIBUTION
It is evident from Table | that the majority of species was present through-
out the period of deposition in the caves. This presence was, however, inter-
mittent in some cases, a fact which could well be due to a paucity of remains
especially in the case of normally rare species such as Dasymys incomtus and
Otomys laminatus. The former, for instance, has so far been found only from
the lowest Holocene levels downwards at Boomplaas A but it is nevertheless
present at Nooitgedacht; likewise at Byeneskranskop it occurs in one of the
middle levels and then not again until the present. Several other species are not
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
represented in the Holocene levels at Boomplaas A, but none of them is ever
numerous and the present sample is not large. In this situation presence is
obviously more significant than absence and further sampling may prove the
fluctuation to be artificial.
In other cases, however, the situation is much clearer. P. natalensis does not
occur at all in the archaeological sample, as was mentioned above, whereas the
opposite is true of Tatera afra (Cape gerbil), Gerbillurus paeba (pygmy gerbil)
and Otomys unisulcatus (bush Karoo rat). The first two species occurred
intermittently and the latter consistently throughout the Upper Pleistocene and
Holocene levels at Boomplaas A, but are not represented in any of the modern
samples collected at various times during the last two years from the Cango
valley. The deserted nests of O. unisulcatus have, however, been found in the
vicinity of Oudtshoorn (C. G. Coetzee pers. comm.) so it is possible that they
may yet be found in the Cango valley. Two further species, Saccostomus
campestris and Steatomys pentonyx, occur only in the Holocene and modern
samples from the Cango valley. S. campestris does not occur in the Gansbaai
area, but S. pentonyx has been found in the Die Kelders 1 LSA levels and at
Byeneskranskop | and 2, but not in the Die Kelders 1 MSA levels. The evidence
suggests, therefore, that these two species migrated into the southern Cape at
the beginning of the Holocene some 10 000 or 12 000 years BP.
CONCLUSION
Increases in the record have been established for most of the species found.
In some cases where previous records were widely scattered the present results
provide significant confirmation of range. In others where the known pattern of
distribution is markedly discontinuous the discovery of linking occurrences in
the Cango valley shows that the disjunction is artificial. The occurrence of
Praomys natalensis with P. verreauxi in the Cango valley is also of considerable
interest since it establishes, at least provisionally, the boundary between the two
species.
Some general information has emerged concerning the movements of
certain species. The evidence seems secure for the arrival at the beginning of the
Holocene of Saccostomus campestris and Steatomys pentonyx and for the more
recent arrival of Praomys natalensis, and for the recent departure of Tatera afra
and Gerbillurus paeba, while Otomys unisulcatus has possibly retreated from the
valley to the plain. Other species have been present, at least intermittently, from
the Upper Pleistocene to the present.
ACKNOWLEDGEMENTS
Thanks are due to the following for making material available: Dr H. J.
Deacon, University of Stellenbosch (Boomplaas A), Mr F. R. Schweitzer,
South African Museum (Die Kelders 1 and Byeneskranskop 1), and Mr G. Avery
and Miss K. Scott, South African Museum (Byeneskranskop 2). Mr A. Botha
kindly pointed out owl roosts in the Cango valley.
DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 209
REFERENCES
Davis, D. H. S. 1962. Distribution patterns of southern African Muridae, with notes on some
of their fossil antecedents. Ann. Cape prov. Mus. 2: 56-76.
Davis, D. H. S. 1974. The distribution of some small southern African mammals (Mammalia:
Insectivora, Rodentia). Ann. Transy. Mus. 29: 135-184. :
DEACON, H. J. & BRooKER, M. 1976. The Holocene and Upper Pleistocene sequence in the
southern Cape. Jn: Proceedings of the Southern African Society for Quaternary Research.
Ann. S. Afr. Mus. 71: 203-214.
DEACON, H. J., DEACON, J. & BROOKER, M. 1976. Four painted stones from Boomplaas Cave,
Oudtshoorn District. S. Afr. archaeol. Bull. 31: 141-145.
GRINDLEY, J., SIEGFRIED, W. R. & VERNON, C. J. 1973. Diet of the Barn owl in the Cape
Province. The Ostrich 44: 266-267.
MEESTER, J. 1963. A systematic revision of the shrew genus Crocidura in southern Africa.
Transy. Mus. Mem. 13: 1-127.
MEESTER, J. & LAMBRECHTS, A. VON W. 1971. The southern African species of Suncus
Ehrenberg (Mammalia: Soricidae). Ann. Transv. Mus. 27: 1-14.
Roserts, A. 1951. The Mammals of South Africa. Johannesburg: The Trustees of ‘The
Mammals of South Africa’ Book Fund.
SHORTRIDGE, G. C. 1942. Field notes on the first and second expeditions of the Cape museums’
mammal survey of the Cape Province: descriptions of some new subgenera and subspecies.
Ann. S. Afr. Mus. 36: 27-100.
TANKARD, A. J. 1976. The stratigraphy of a coastal cave and its palaeoclimatic significance.
Palaeoecol. Afr. 9: 151-159.
TANKARD, A. J. & SCHWEITZER, F. R. 1974. The geology of the Die Kelders Cave and environs:
a palaeoenvironmental study. S. Afr. J. Sci. 70: 365-369.
VERNON, C. J. 1972. An analysis of owl pellets collected in southern Africa. The Ostrich 43:
109-124.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
‘An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name ‘(and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
- Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: ee pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861:
Leda bicuspidata: Nicklés, 1950: 165, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
- not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. °.. . the Figure depicting C. namacolus ...’; *. .. in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
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book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
D. M. AVERY
PAST AND PRESENT DISTRIBUTION OF
SOME RODENT AND INSECTIVORE SPECIES
IN THE SOUTHERN CAPE PROVINCE,
SOUTH AFRICA: NEW INFORMATION
- VOY FAZQ .
_ VOLUME 74 PART 8 JANUARY 1978 ISSN 0303-2515
| MUS. COMP, ZOOL,
LIBRARY
APR 28 1978
I ARV Are
UNIVERSITY
ANNALS
OF THE SOUTH AFRICAN
~ MUSEUM
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2. LAYOUT should be as follows:
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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 i of publication given in text, e.g.:
‘Smith (1969) describes .
‘Smith (1969: 36, fig. 16) ‘gettnes:
‘As described (Smith 1969a, 19696; roa ae
‘As described (Haughton & Broom 1927) .
‘As described (Haughton et al. 1927)...’
Note: no comma separating name and year
Dagination indicated by colon, not p.
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et al. in text for more than two joint authors, but names of all authors given in list of references.
(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
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number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOuGH, W. S. 1960. Practical invertebrate anatomy. 2nd aa London: Macmillan.
FISCHER, P. —H. 1948. Données sur la résistance et de Je vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Konn, A. J. 1960a. Ecologicai notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, eee! masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4):
THIELE, J. 1910. Mollusca: B. Polyplaeenharas Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
January 1978 Januarie
Parties) Deel
CRETACEOUS FAUNAS FROM ZULULAND AND
NATAL, SOUTH AFRICA
A FLICKIA FROM THE CENOMANIAN OF
NORTHERN ZULULAND
By
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
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ISBN 0 908407 29 7
Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
A FLICKIA FROM THE CENOMANIAN OF NORTHERN ZULULAND
By
WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford
&
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
(With 2 figures)
[MS. accepted II August 1977]
ABSTRACT
The Lower Cenomanian Mzinene Formation of the Ndumu area of northern Zululand
has yielded specimens of the peculiar cryptogenic dwarf ammonite Flickia Pervinquiere, 1907
—type genus of the Family Flickiidae Pervinquiére, 1910, a group characterized by simplified,
goniatitic sutures. The family was previously known almost exclusively from pyritic clay
facies, and the present occurrences in a nearshore clastic sequence is unusual. The specimens
are referred to Flickia quadrata Collignon, 1964, a species previously known only from
Madagascar. They represent the first record of the family from Africa south of the Sahara.
CONTENTS
PAGE
Introduction Q : : : 3 ee
Systematic palaeontology ; te?) 4
Discussion . 5 : ? d 2 216
Acknowledgements . j jf BS FO
References . : : ; i , alk
INTRODUCTION
In 1907 Pervinquiere introduced the new genus and species Flickia simplex
for a group of diminutive (S-15 mm diameter) moderately evolute compressed,
smooth ammonites with narrowly arched venters and extraordinarily goniatite-
like sutures. Pervinquiére compared his specimens with the Triassic ceratites
Nannites Mojsisovics, 1881, and Lecanites Mojsisovics, 1882, as well as con-
sidering that they might perhaps be juveniles of Neolobites Fisher, 1882, a
view which he rejected on the basis of fundamental differences in sutures and
shell form. He did, however, regard Flickia as a close ally within the “Hoplités’.
Thirteen years later, W. S. Adkins, working on the pyritic ammonite
faunas of similar clay facies occurring in Texas (Adkins 1920) described two
further species, Flickia boesi Adkins (Adkins 1920: 85, pl. 1 (figs 1-3)) and
Flickia? bosquensis Adkins (Adkins 1920: 87, pl. 1 (fig. 4)). The latter, with
211
Ann, S. Afr. Mus. 74 (8), 1978: 211-217, 2 figs.
PND, ANNALS OF THE SOUTH AFRICAN MUSEUM
prominent umbilical bullae, Adkins believed to represent a new genus, subse-
quently named Adkinsia by Emile Bose, with Adkinsia adkinsi as type species
(Bose 1928: 237, pl. 8 (figs 3-14)). Bose named three other species of Adkinsia
in the same publication, all from the same horizon (the Del Rio Clay), all,
however, probably being no more than intraspecific variants (A. sparcicosta
Bose, 1928: 238, pl. 8 (figs 15-20); A. tuberculata Bose, 1928: 240, pl. 87
(figs 21-26) and A. semiplicata Bose, 1928: 246, pl. 9 (figs 7-12)). In the same
year Adkins (1928) introduced the Family Flickiidae (as Flickidae) to
accommodate these two genera, without commenting on their affinities.
With Collignon’s (1928 onwards) monographic studies of the ammonite
faunas of Madagascar, further examples of these curious genera were described,
again as minute individuals in pyritic clay facies (Flickia pervinquierei Collignon,
1928: 4, pl. 2 (figs 15, 15a); Flickia costellata Coliignon, 1964: 23, pl. 322
(fig. 1427) and Flickia quadrata Collignon, 1964: 23, pl. 322 (figs 1428-1429)).
The group thus show a strong facies-linked distribution, but unlike some
other pyritic micromorphs from clay facies which have proved to be no more
than nuclei of large limestone individuals (e.g. many Submantelliceras Spath,
1923 and Neopulchellia Collignon, 1929; see Casey (1965) and Kennedy &
Hancock (1971, 1977) for discussions) they appear to be genuine micromorph
inhabitants of offshore clay depositing environments.
The present Zululand specimens, from lignitic nearshore siltstone sequences,
are thus a unique occurrence which extend both facies and geographic ranges
of these curious ammonites, as well as providing the first adult individuals to be
described which retain all of the body chamber and the aperture.
SYSTEMATIC PALAEONTOLOGY
Phylum MOLLUSCA
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order AMMONITIDA Hyatt, 1889
Superfamily ACANTHOCERATACEAE Hyatt, 1900
Family Flickiidae Adkins, 1928
Genus Flickia Pervinquiere, 1907
Type species |
Flickia simplex Pervinquiere, 1907 by original designation.
Flickia quadrata Collignon, 1964
Figs 1-2
Flickia quadrata Collignon, 1964: 23, pl. 322 (figs 1428-1429).
J\\3)
CRETACEOUS FAUNAS FROM SOUTH AFRICA
Both
I-K. BMNH C79841.
A-H. BMNH C79976.
Fig. 1. Flickia quadrata Collignon.
from the Mzinene Formation, Cenomanian I, Locality 181 of Kennedy & Klinger (1975).
E-H, J x 1.
b)
A=D, I,K x 2
214 ANNALS OF THE SOUTH AFRICAN MUSEUM
Holotype
The specimen figured by Collignon (1964, pl. 322 (fig. 1428)), from the
Lower Cenomanian of Beraketa sur Sakondry (Manera), Madagascar.
Material
Two specimens, BMNH C79841 and BMNH (C79976, both from
Locality 181 of Kennedy & Klinger (1975: 304), the Mzinene Formation
exposed in hill slopes east of the road, 1 km south-east of the store at Ndumu,
northern Zululand, 26°55’51”S_ 32°18’29”E. C79841 is from Bed 5 and C79976
from Bed 3 of this section (Kennedy & Klinger 1975, fig. 12, inset) and are of
basal Cenomanian age (Cenomanian I).
Dimensions
All dimensions are in millimetres.
D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilicus.
Figures in parentheses are dimensions expressed as a percentage of over-all
diameter.
D Wb Wh Wb:Wh U
BMNH 79976 23,8(100) 8,2(34) —_—_7,5(32) 1,09 7,8(33)
18,7(100) 6,7(36) —-6,3(34) 1,06 6,5(35)
Diagnosis
A slightly depressed to slightly compressed Flickia ornamented by low,
broad irregular prorsiradiate fold-like ribs separated by variably developed
constrictions, the shell surface being covered in coarse striae.
Description
Both specimens are adult and retain complete body chambers extending for
between half and two-thirds of the last whorl. They are preserved as internal
moulds, with traces of the original nacreous aragonitic shell present.
Coiling is evolute with a shallow, fairly wide umbilicus which comprises
33-35 per cent of the diameter. The whorls expand rather slowly, and are
slightly compressed on the inner whorls, becoming slightly depressed on the
outer. The greatest breadth is a little below mid-flank; the umbilical wall is low,
the shoulder rounded, inner flanks somewhat flattened and subparallel (Fig. 2A),
outer flanks convergent with an arched, rounded venter.
Ornament consists of rather irregular, low, broad, prorsiradiate fold-like
ribs, which are narrow at the umbilical shoulder but widen across the flanks to
pass across the venter as a broad, rounded swelling. The interspaces are of
irregular width and depth, and some are accentuated into distinct constrictions.
The shell surface (where preserved) of both ribs, interspaces and constrictions
is covered by well-developed striae (Fig. 1D, G); ribs, striae and constrictions
all strengthen on the body chamber. The aperture (Fig. 1D) is simple, entire,
CRETACEOUS FAUNAS FROM SOUTH AFRICA DNS
slightly flared in BMNH C79976, and preceded by a narrow rib in BMNH
C79841 (Fig. 11).
Both specimens show the sutures (Figs 1K, 2B), which are very simple and
of basic Flickiid type, with entire lobes and saddles. There is a small median
element to the external lobe E, a rather narrow lateral lobe L, and a shallow,
broad U,. Saddle E/L is the largest, and has a flattened top; L/U, is smaller
and rounded. In both specimens, the last few sutures crowd, indicating the
specimens to be adult.
Comparisons with other species
Flickia simplex (Pervinquiere 1907: 214, pl. 9 (figs 2a—b, 3a—b, 4, 5a—b);
Collignon 1928-9: 28, pl. 2 (figs 14, 14a); Collignon 1964: 23, pl. 322
(fig. 1425) is a much more compressed species during middle, and presumably
later growth. It is ornamented only by fine, falciform striae.
Flickia pervinquierei (lectotype here designated is the original of Collignon
S29, ol, 2 Cass (5, 15a)3 see also Colltiynon MWSle Fo, ob 2 Chess Zl, wiley)
Collignon 1964, pl. 322 (fig. 1426)) is also flat-sided and compressed, completely
smooth and, according to Collignon (1929), distinctly carinate. In his original
description of this species Collignon also referred the original of Pervinquiére’s
(1907) plate 9 (fig. 5) to Flickia pervinquierei. This specimen is, in fact, the
holotype of Flickia simplex, being so designated by Pervinquiere in his
explanation of the plate.
Flickia costellata Collignon (1964: 23, pl. 322 (fig. 1427)), is also rather
compressed but resembles Flickia quadrata in being ribbed, although these ribs
are simple, blunt and narrow rather than broad folds with associated
constrictions.
All described Adkinsia species bear strong umbilical bullae, whilst Fischeuria
Pervinquiere, 1910 are very involute, globose and smooth when young with an
| oe
Fig. 2. Flickia quadrata Collignon. A whorl section and B suture of BMNH C79841.
216 ANNALS OF THE SOUTH AFRICAN MUSEUM
angular umbilical shoulder. At least one species, Fischeuria pusilla Matsumoto &
Inoma (1975: 290, pl. 42 (figs 4-5)), develops folds and constrictions on the
body chamber.
Representatives of neither of these genera are liable to be confused with
Flickia.
Occurrence
Low Lower Cenomanian of Madagascar and the Ndumu area, Zululand.
DISCUSSION
Flickia was first described from north Africa, where Pervinquiere (1907)
stated the same specimens to be of both ‘Vraconien’ (e.g. Upper Albian) and
‘Cénomanien’ age in his description. Flickia quadrata was originally described
from Collignon’s Zone of Mantelliceras martimpreyi (auctorum, non Coquand)
(fide Kennedy & Hancock 1971) at his locality 478, Beraketa sur Sakondry
(Manera), Madagascar.
From the same locality Collignon (1964) recorded a large fauna, all as
minute pyritic individuals, including many heteromorphs and the following
normally coiled ammonites: Desmoceras latidorsatum (Michelin), D. lemoinei
Collignon, Protokossmaticeras madagascariense Collignon, Forbesiceras
largilliertianum (d’Orbigny), “Neopulchella gignouxi Collignon, Euhystrichoceras
(Sakondryella) madagascariense Collignon, Prionocycloides proratus (Coquand),
P. besairiei Collignon, Flickia simplex Pervinquiere, F. pervinquierei Collignon,
Neosaynoceras. gazellae Pervinquiere, Mantelliceras aumalense (Coquand),
M. mantelli (J. Sowerby), M. tuberculatum (Mantell), M. suzannae (Pervinquiere),
M. decaryi Collignon and Acompsoceras waterloti Collignon.
This fauna is that which is well known elsewhere in Madagascar and also
north Africa, where it is variously referred to the ‘martimpreyi’ Zone or
Submantelliceras aumalense Subzone, whilst the closest North American faunas
are approximately those of the Graysonites adkinsi Zone.
The presence in South Africa of one of the taxa apparently restricted to this
level in clay facies elsewhere is thus of interest, as it allows correlation with a
facies where there are large ammonites. This is important in deciphering the
still poorly understood relationship between pyritic micromorph faunas and
those of limestone and sandstone facies in Europe, India and elsewhere, where
pyritic micromorphs are either absent or unrecognized as nuclei of much bigger
specimens (Kennedy & Hancock 1971, 1977).
Bed 1 at locality 181 near Ndumu, concretions 1,6 m below the first
specimen of Flickia quadrata, yielded abundant heteromorphs, especially
Hamites and Anisoceras of the A. pseudoelegans Pictet & Campiche—A. campichei
Spath group, indicative of the Upper Albian Stoliczkaia dispar Zone. Bed 2,
which yielded a single F. quadrata, contained abundant Sharpeiceras, including
specimens resembling S. falloti (Collignon), S. florencae Spath and S. vohipalense
CRETACEOUS FAUNAS FROM SOUTH AFRICA DN
Collignon, rare Utaturiceras vicinale (Stoliczka), submantellicerine nuclei, and
Mariella (Mariella) oehlerti (Pervinquiére) and varieties, and Ostlingoceras
(Ostlingoceras) spp. Bed 3, a little higher still, yielded a further Flickia, and is
crowded with the same M. (Mariella) and O. (Ostlingoceras). Some tens of
metres higher in the section, a more diverse fauna with Forbesiceras,
Mantelliceras and Hypoturrilites appears.
These observations provide a basis for correlation of pyritic and limestone
faunas at the base of the Cenomanian in the area, and suggest that Flickia
comes from what is virtually the base of the Cenomanian. Sadly, the associated
Zululand faunas are still insufficient to tell whether or not these levels are
absent in Europe or simply represented in a different biofacies (Kennedy &
Hancock 1977).
ACKNOWLEDGEMENTS
We thank the following colleagues who have advised and helped in many
ways. Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural History),
Mr C. W. Wright (Oxford), Dr M. R. Cooper (Oxford), Dr J. M. Hancock
(London), General M. Collignon (Moirans, Isére), Dr C. Duerden and Professor
K. Young (Austin, Texas), and the staff of the Geological Collections, Oxford
University Museum.
One of us (W.J.K.) acknowledges with thanks financial support from the
Royal Society, N.E.R.C. and the Trustees of the Sir Henry Strakosh Bequest.
REFERENCES
ADKINS, W. S: 1920. The Weno and Pawpaw formations of the Texas Comanchean. Bull. Univ.
Tex. econ. Geol. Tech. 1856: 1-172.
ADKINS, W. S. 1928. Handbook of Texas Cretaceous fossils. Univ. Texas Bull. 2838: 1-303.
Bose, E. 1928. Cretaceous ammonites from Texas and northern Mexico. Bull. Univ. Tex. econ.
Geol. Tech. 2748: 143-312.
CASEY, R. 1965. A monograph of the Ammonoidea of the Lower Greensand, part VI.
Palaeontogr. Soc. (Monogr.): 399-546.
COLLIGNON, M. 1928-1929. Les céphalopodes du Cénomanien pyriteux de Diego—Suarez,
Paléontologie de Madagascar. Annis. Paléont. 17 (1928): 136-160 (1-24); 18 (1929):
1-56 (25-29).
COLLIGNON, M. 1931. La faune du Cénomanien a fossiles pyriteux du Nord de Madagascar.
Annls. Paléont. 20: 43-104 (1-64).
COLLIGNON, M. 1964. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XI
(Cénomanien). Tananarive: Service Géologique.
KENNEDY, W. J. & HANCOCK, J. M. 1971. Mantelliceras saxbii (Sharpe) and the horizon of the
Martimpreyi Zone in the Cenomanian of England. Palaeontology 14: 437-454.
KENNEDY, W. J. & HANcocK, J. M. 1977. Towards a correlation of the Cenomanian sequences
of Japan and western Europe. Trans. Proc. palaeont. Soc. Japan Spec. Pap. 21: 127-141.
KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South
Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315.
Matsumoto, T. & INomaA, A. 1975. Mid Cretaceous ammonites from the Shumarinai—
Soeushinai Area, Hokkaido. Mem. Fac. Sci. Kyushu Univ., Ser. D, Geol. 23: 263-293.
PERVINQUIERE, L. 1907. Etudes de paléontologie Tunisienne, 1. Céphalopodes des terrains
secondaires. Mém. Carte géol. Tunisie, 438 pp.
WRIGHT, C. W. 1957. In: ARKELL, W. J. et. al. Treatise on Invertebrate Palaeontology L.
Mollusca (4). Geological Society of America and University of Kansas Press.
y = ri
uy i 1 il
toh -
_) \ = is
Mw
cs F chal i
; i
4 =e
ie, Fr
7 aT i ia
: y : _
} 7 i] iE t - ah eh
, i v or t ; Dare: ie
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., Syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must_be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
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figures of plates are enclosed in parentheses to distinguish them from text-figures
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Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
~ not acceptable.
In describing new Species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
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must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
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by initials or full names
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
WILLIAM JAMES KENNEDY &
HERBERT CHRISTIAN KLINGER
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
A FLICKIA FROM THE CENOMANIAN OF
NORTHERN ZULULAND
a Chak | 0303-2515
VOLUME 74 PART 9 JANUARY 1978 | ISSN
LIBRARY
APR 2 8 1978
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(a) Author’s name and year of publication given in text, e.g.:
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Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. °
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHeER, P.-H., DuvAL, M. & RaFFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
January 1978 Januarie
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A NEW SPECIES OF TORTANUS
~ (CRUSTACEA, COPEPODA) FROM SOUTH AFRICA
By
JOHN R. GRINDLEY
Cape Town Kaapstad
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A NEW SPECIES OF TORTANUS
(CRUSTACEA, COPEPODA) FROM SOUTH AFRICA
By
JOHN R. GRINDLEY
School of Environmental Studies, University of Cape Town
(With 4 figures and 3 tables)
[MS. accepted 16 August 1977]
ABSTRACT
A new species of Copepoda Tortanus capensis (Calanoida, Tortanidae) is described from
lagoons and estuaries on the west and south coasts of South Africa. This species appears to
be most closely related to Tortanus recticauda Giesbrecht from the Red Sea and must be
placed in the subgenus Afortus.
CONTENTS
PAGE
Tortanus capensis sp. nov. algnete, ADO
Occurrence . : ; ‘ Bur ae
Types . i ; 3 j eal S)
Description : ; ? Sg elle
Juvenile stages . : : eae S
Remarks . : : : 226
Acknowledgements . , : Sate 22h
References . ; ; ; : rh ON
Tortanus capensis sp. nov.
(Figs 1-4)
Occurrence
Numerous specimens, comprising adult males and females and copepodite
stages of both males and females from estuaries and lagoons on the west and
south coasts of South Africa, in salinities between 34,5%, and 40,0%,.
Types ,
Type specimens from Langebaan Lagoon, 100 km north of Cape Town
have been deposited in the South African Museum, Cape Town. Holotype,
male, SAM-A12227, allotype, female, SAM—A12228, 15 paratypes, male,
female and juvenile, SAM—A12229, with dissected appendages mounted on
slides. Further paratypes also from Langebaan Lagoon have been deposited in
the British Museum (Natural History) and the United States National Museum.
Description
Female
Length 2,20—-2,40 mm.
Prosome (Fig. 1B—C) slender, viewed dorsally three times as long as wide,
characteristically shaped with a cervical groove prominent in lateral view.
DNS
Ann. S. Afr. Mus. 74 (9), 1978: 219-228, 4 figs, 3 tables.
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
Forehead shallowly sloped. Rostral projections without filaments. Eyes
prominent and reddish. All five pedigerous segments distinct but fourth and
fifth partly fused. Posterior angles of metasome produced but rounded.
Urosome (Fig. 1B—C) two-segmented, last segment partly fused with the
caudal rami. The genital segment is swollen.
Caudal rami (Fig. 1B—-C) asymmetrical, right ramus slightly wider and
longer, about four and a half times as long as wide distally but narrowed
proximally, furnished with fine hairs on the distal part of their medial margins.
Five plumose and one non-plumose caudal setae on each ramus, of which the
second inner plumose seta is twice as long as the neighbouring setae.
Antenna | (Fig. 1A) of fifteen separate segments, almost reaching the end
of the caudal rami when extended. Apparently segments 1-6, 8-12 and 24-25
are fused, but the segmentation of the proximal part of the antenna is somewhat
obscure. Some further fusions are apparent in immature stages in the proximal
segments and segments 23-25 are still fused in Stage IV. Aesthetasks appear on
a number of segments particularly on the proximal part of the antenna. Four
long setae are present on proximal segments and on segments 14, 16, 18 and 21
while six terminal setae arising from an expanded end give the antennae a
characteristic appearance. The terminal setae are all plumose and the four long
ones are asymmetrically constricted near their bases. Some specimens have one
or two small terminal non-plumose setae.
The second antenna and mouthparts appear to be identical in the female
and male except for the mandibular palp and are figured for the female
(Fig. 3D-H).
Antenna 2 (Fig. 3D) with basipod, bearing one lateral and one sub-
terminal seta and a medial fringe of fine hairs. Exopod one-segmented, bearing
six setae and a small sub-terminal fringe of bristles. Endopod two-segmented,
bearing four terminal setae and a small sub-terminal seta.
Mandible (Fig. 3E) with gnathal lobe heavily chitinized, bearing one large
and four smaller teeth. Each of the smaller teeth has a supporting rib bearing a
longitudinal row of short bristles on tiny transverse striations. Basipod of
mandibular palp unarmed. Exopod and endopod each one-segmented in the
male and bearing four and five terminal setae respectively. In the female the
exopod and endopod are each two-segmented and bear five and six setae
respectively.
Maxilla (Fig. 3F) with gnathobase, bearing seven serrated and plumose
spines, three plumose setae and three small non-plumose setae. Endopod one-
segmented, bearing three barbed terminal setae, one of which has widely spaced
plumules.
Maxilla 2 (Fig. 3G) stout, with three endites, and indistinct indications of
segmentation. Two of the three small lateral setae bear bristles. The three
different setae on the medial endite bear a variety of bristles. The remaining six
long terminal setae are flattened and bear lateral rows of spinules all medially
orientated (see enlarged figure of tip). Small subsidiary setae are situated at the
A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA DD)
Fig. 1. Tortanus capensis sp. noy. Female.
A. Antenna 1. B. Dorsal view. C. Lateral view. D. Fifth legs. E. First swimming leg.
F. Second swimming leg. G. Third swimming leg. H. Fourth swimming leg.
bases of the second and sixth long setae and the two barbed lateral setae.
Maxilliped (Fig. 3H) small and three-segmented, the basal segment bearing
two long, curvaceous setae ornamented with diverging pairs of bristles, and the
third segment bearing a small lateral seta and three strongly curved barbed
terminal setae.
oD) ANNALS OF THE SOUTH AFRICAN MUSEUM
Swimming legs 1-4 (Fig 1E-H) biramous with two-segmented basipodite,
two-segmented endopodite and three-segmented exopodite. They are similar in
both sexes. The ornamentation of the swimming legs is represented in Table 1.
TABLE |
Ornamentation of swimming legs.
Protopod Endopod Exopod
Leg ] 2 1 2 I 2) 3
Si Se Si Se Si Se Si St Se Si Se Si Se Si St Se
P, 1 O 0 O 3 0 SF ue 8 Le 1 O 4 I I
P, 1 0 O 9. 0 See rel ee | 1 5 Wl Wl
P, 1) 0 O JO Spee, oie i at ae | > ei
P, i 0 0 O 3) a eee | Nee 1 t eae UE |
Si, Se, St represent internal, external and terminal spines, or setae respectively. The number of
setae is shown in arabic numerals and spines in roman numerals.
The first leg has four less spines and one less seta than the remainder. The
fourth leg has a fringe of fine hairs on the medial margins of the second basipod
segment and the first exopod segment. All the setae on the swimming legs are
plumose, and the setae on the endopods are much more slender than the
remainder.
The fifth pair of legs of the female (Fig. 1D) are uniramous, three-segmented
and symmetrical. The second segment bears an outer seta and the third segment
bears a stout terminal spine and a subsidiary basal spine. The terminal spine is
not articulated but fused to the third segment and it is finely barbed distally.
Male
Length 1,80-1,85 mm.
Prosome (Fig. 2C, F) somewhat different from that of the female, with the
head more evenly tapered but with a cervical groove like that of the female.
Forehead shallowly sloped. Rostral filaments absent. Eyes not prominent and
dark brown. All five pedigerous segments distinct but fourth and fifth partly
fused. Posterior angles of metasome rounded and not produced.
Urosome five-segmented of which the first is largest. Caudal rami (Fig. 2E)
symmetrical and about five times as long as wide. The urosomes of several
males were characteristically twisted to the right.
Left antenna | (Fig 2A-B) fifteen-segmented and similar to that of the
female.
Right antenna | (Fig. 2D) geniculate, with fifteen separate segments, two
of which are beyond the articulation. Segments 1-6, 19-21 and 22-25 fused.
Prominent setae appear on proximal segments and on segments 14, 16, 18 and
21. The development of the specialization of the geniculate antenna may be
observed in the right antenna 1 of a stage V male copepodite (Fig. 4H).
A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 223
—,
=
S&
SS
WS 7
—jy
\
EEE EEE
=a
Fig. 2. Tortanus capensis sp. noy. Male.
A-B. Left antenna. C. Dorsal view. D. Right antenna. E. Caudal rami.
F. Lateral view.
Segments 11-12 and 23-25 are fused at that stage, but segments 18 to 22 are
free and there are prominent spines on segments 18 and 20. It would appear
that the spine on segment 18 forms the serrated lamella on that segment of the
geniculate antenna of the adult male.
Male fifth legs (Fig 3A—C) uniramous and asymmetrical with the right leg
three-segmented and the left leg four-segmented. The first segment of the right
leg is swollen and produced into a rounded projection at its medial distal corner.
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. Tortanus capensis sp. nov.
Male. A. Fifthlegs. B. Right fifth leg. C. Left fifth leg. Female. D. Antenna. E. Mandible
and palp. F. First maxillary. G. Second maxilla (end of one seta enlarged). H. Maxilliped.
The second segment bears a large bilobed medial projection near its distal end,
which bears two setae. The terminal segment is in the form of a sub-chelate
claw, strongly chitinized near the tip and bearing two medial setae, and two tiny
bristles near its tip.
The first segment of the left leg is short. The second segment is longer and
curved and bears a medial and an outer seta. The third segment is strongly
curved tapers distally, and has an angular swelling proximally bearing a medial
A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA DUDES
\ ea
Fig. 4. Tortanus capensis sp. nov. Juveniles.
A. Stage IV female, dorsal view. B. Fifth legs, stage IV female. C. Fifth legs stage V female.
D. Stage IV male, dorsal view. E. Fifth legs, stage IV male, tip enlarged. F. Stage V male,
dorsal view. G. Fifth legs, stage V male. H. Right antenna I, stage V male.
seta. The terminal segment is even more strongly curved and bears a finely
serrate terminal spine, two outer marginal spinules, two inner marginal setae,
and a large number of tiny bristles.
Spermatophore 300-380 pm long.
Juvenile stages (Fig. 4A—-H)
Copepodite stages including stages four and five of both males and females
were obtained. The changes that occur in the development of the urosome and
the fifth legs particularly are apparent in the figures. Lengths: Stage IV female
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
1,45 mm, Stage V female 1,75 mm, Stage IV male 1,12 mm, Stage V
male 1,50 mm. The ornamentation of the swimming legs of the Stage IV and
Stage V copepodites is represented in Tables 2 and 3.
TABLE 2
Copepodite Stage IV. Ornamentation of swimming legs.
Protopod Endopod Exopod
P, | 0 0 0 6 2 1 1 0 5 I I
P, 1 0 0) 0 6 2; 1 1 I 6. ALLY AE
P; 1 0 0 0 5 D I 1 I 5 i If
P, I 0 0 0 4 D 1 1 I 5) YT Ui
Si, Se, St represent internal, external and terminal spines, or setae respectively. The number
of setae is shown in arabic numerals and spines in roman numerals.
TABLE 3
Copepodite Stage V. Ornamentation of swimming legs.
Protopod Endopod Exopod
1 Ze 1 Z 1 2, 3
Si Se Si Se Si Se Si St Se Si Se Si Se Si St Se
P, I 0 O 3) 0) SN ART io ke O 2 etna Ug |
P, he 0 O 30 Sri De el Ua | eek eed UCP edt
P, LEO 0 0 3 O Bae hal ioe | 1 ean ef +. Tk SF
P, iL @ 0 O 3-00 Bi.) Wate al he ye List 53> lle cED
Si, Se, St represent internal, external and terminal spines, or setae respectively. The number
of setae is shown in arabic numerals and spines in roman numerals.
Remarks
Tortanus capensis is apparently the twenty-first species of this peculiar
genus of copepods to have been described. The genus Corynura was established
by Brady (1883) for two species obtained by the Challenger expedition in the
Philippine islands. This generic name which was preoccupied was replaced by
the name TJortanus (from tortus and anus) by Giesbrecht & Schmeil (1898). The
following species are now included in this genus: Tortanus barbatus (Brady),
1883 (=denticulatus Giesbrecht); Tortanus gracilis (Brady), 1883; Tortanus
forcipatus (Giesbrecht), 1889; Tortanus recticauda Giesbrecht, 1889; Tortanus
discaudatus (Thompson & A. Scott), 1897; Tortanus setacaudatus Williams,
1906; Tortanus murrayi A. Scott, 1909; Tortanus brevipes A. Scott, 1909;
Tortanus (Atortus sub. gen. nov.) tropicus Sewell, 1932; Tortanus derjugini
Smirnov, 1935 (see Brodsky 1950); “Tortanus sp.’ Colefax, 1940 Um Dakin &
A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA UE
Colefax 1940); Tortanus longipes Brodsky, 1948; Tortanus vermiculus Shen,
1955; Tortanus spinicaudatus Shen & Bai, 1956; Tortanus denticulatus Shen &
Lee, 1963; Tortanus compernis Gonzales & Bowman, 1965; Tortanus rubidus
Tanaka, 1965; Tortanus giesbrechti Jones & Park, 1968; Tortanus scaphus
Bowman, 1971; Tortanus lophus Bowman, 1971; and Tortanus capensis sp. nov.
(S77).
Tortanus capensis is the first species of the genus to have been found
anywhere round the coasts of Africa, south of the Red Sea. It appears to be
most closely related to Tortanus recticauda from the Red Sea (Giesbrecht 1889)
and Tortanus murrayi, scaphus and lophus from the East Indies (A. Scott 1909;
Bowman 1971), but it differs from each of them in many characters. Tortanus
capensis may be most readily distinguished by the fifth legs of either the male or
the female, which are quite distinct from those of any of the described species of
Tortanus.
Specimens of Tortanus capensis from the Swartkops estuary on the south-
east coast of South Africa showed a few minor differences in comparison with
the type material from Langebaan Lagoon.
In the two-segmented urosome of the female and the character of the male
fifth legs Tortanus capensis might be placed in Steuer’s group 3 which he
established in his 1926 revision of the genus Jortanus. This is equivalent to
Sewell’s (1932) subgenus Afortus (Bowman 1971).
ACKNOWLEDGEMENTS
I thank Mrs S. Grindley and Mr T. Wooldridge for assistance and the
South African National Committee for Oceanographic Research for financial
support.
REFERENCES
Bowman, T. E. 1971. Tortanus scaphus and Tortanus lophus, new Pacific planktonic copepods,
with notes on Tortanus murrayi (Calanoida, Tortanidae). Pacif. Sci. 25 (4): 521-528.
Brapy, G. S. 1883. Report on the Copepoda collected by H.M.S. Challenger during the years
1873 to 1876. Challenger Rep. 1873-76 8: 1-142.
Bropsky, K. A. 1948. T. Jongipes sp. nov. from the Sea of Japan. Proc. Pacif. Sci. Res. Inst.
Fish Oceanogr. 26: 80. (In Russian.)
Bropsky, K. A. 1950. Calanoida of the far eastern and polar seas of the U.S.S.R. Tabl. anal.
faune U.R.S.S. Zool. Inst. Acad. Sci. 35: 1-442. (In Russian.)
Dakin, W. J. & CoLeFAx, A. N. 1940. The plankton of the Australian coastal waters of New
South Wales. Publ. Univ. Sydney (Zool.) 1 (1): 1-215.
GIESBRECHT, W. 1889. Elenco dei Copepodi pelagici. Atti. R. Ac. Lincei (4) 5: 24-29.
GIESBRECHT, W. & SCHMEIL, O. 1898. Copepoda. 1. Gymnoplea. Tierreich 6: 1-169.
GONZALES, J. G. & BOwMan, T. E. 1965. Planktonic copepods from Bahia Fosforescente,
Puerto Rico, and adjacent waters. Proc. U.S. nat. Mus. 117: 241-303.
JONES, E. C. & Park, T. S. 1968. A new species of Tortanus (Calanoida) from Pago Pago
Harbour, American Samoa. Crustaceana (Suppl. 1): 152-158.
Scott, A. 1909. The Copepoda of the Siboga expedition. Siboga Exped. Monogr. 29 (A): 1-323.
SEWELL, R. B. S. 1932. The Copepoda of Indian seas. Calanoida. Mem. Indian Mus. 10:
400-402.
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
SHEN, C. J. 1955. On some marine crustaceans from the coastal water of Fenghsien, Kiangsu
Province. Acta. zool. Sin. 7: 75-100. (In Chinese & English.)
SHEN, C. & Bal, S. 1956. The marine Copepoda from the spawning ground of Pnueumatophorus
japonicus (Houttuyn) off Chefoo, China. Acta. zool. Sin. 8: 177-234.
SHEN, C. J. & Lee, F. S. 1963. The estaurine copepoda of Chiekong and Kaikong Rivers,
Kwangtung Province, China. Acta. zool. Sin. 15: 571-596.
STEUEUR, A. 1926. Revision der Copepodengattung Tortanus Giesbr. Boll. Soc. Adriatica Sci.
Nat. 29: 49-69.
TANAKA, O. 1965. The pelagic copepods of the Izu region, Middle Japan. Systematic account
XIII. Parapontellidae, Acartiidae and Tortanidae. Publ. Seto mar. biol. Lab. 12: 379-408.
THompson, I. C. & Scott, A. 1897. On the plankton collected continuously during the two
traverses of the North Atlantic in the summer of 1897. Trans. Liverpool biol. Soc. 12: 80.
WILLIAMS, L. W. 1906. Notes on marine Copepoda of Rhode Island. Amer. Nat. 40 (447):
639-660.
6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
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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:
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Polehvpe
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
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JOHN R. GRINDLEY
A NEW SPECIES OF TORTANUS
(CRUSTACEA, COPEPODA) FROM SOUTH AFRICA
| rt SA
VOLUME 74 PART 10 JANUARY 1978 | |
| | MUS. COMP. Oe ee
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & RAFFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
January 1978 Januarie
Part 10 #£4Deel
NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 3
SOME OBSERVATIONS ON THE DISTRIBUTION
AND BIOLOGY OF SCORPIONS ON
TABLE MOUNTAIN
By
FE. B. EASTWOOD
Cape Town Kaapstad
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NOTES ON THE SCORPION FAUNA OF THE CAPE
PAR 3
SOME OBSERVATIONS ON THE DISTRIBUTION AND BIOLOGY OF
SCORPIONS ON TABLE MOUNTAIN
By
E. B. EASTWOOD
South African Museum, Cape Town
(With 12 figures and 4 tables)
[MS. accepted 18 August 1977]
ABSTRACT
A survey of the scorpion fauna of Table Mountain on the Cape Peninsula was carried
out. Four of the six species recorded from this area were found. Distribution and habitat
preferences are discussed with reference to topography and vegetation. Some observations on
behaviour in the field and laboratory are recorded. A regional key is given.
CONTENTS
_ PAGE
Introduction : ; : : ; : : PS)
Description of the study area . : ; A an 230
Materials and methods . ; : ; 4 e723)
Distribution and habitat . ; ; : ; le 3 7)
Scorpion shelters : : : ; : : 3 236
Coexistence : : 3 : f : : eA)
Scorpion predators . é : : , : Fae Al
Some observations on behaviour . d : We
Key to the scorpions of Table Mountain . . 244
Conclusions 4 : , : : , : e246
Acknowledgements . : ‘ . , . 5 Vay
References . ; ; i 3 f 4 5 ot ead
INTRODUCTION
Six species in two families of scorpions have been recorded from Table
Mountain. Opisthophthalmus capensis (Herbst) was recorded by Pocock (1896),
Purcell (1899) and Lawrence (1955); Uroplectes insignis Pocock by Pocock
(1890, 1896), Kraeplin (1894, 1899), Purcell (1901) and Hewitt (1918); U. lineatus
(Koch) was reported by Purcell (1901) and Kraeplin (1908); U. variegatus (Koch)
by Pocock (1896) and Hewitt (1918), and Parabuthus capensis (Ehrenberg) by
Hewitt (1918) and Eastwood (19775). A juvenile specimen from Signal Hill was
identified as U. vittatus (Thorell) by Penther (1900) but it is very likely that this
record pertains to U. lineatus. The work on Cape scorpions by early taxonomists
was confined to morphological and geographical considerations. With later
descriptions of new forms, a reappraisal of scorpion taxonomy has become very
complex without the relevant background on ecology and behaviour.
A survey was carried out by regular collecting over a period of one year.
229
Ann. S. Afr. Mus. 74 (10), 1978: 229-248, 12 figs, 4 tables.
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
To complement field observations, live scorpions were kept in the laboratory.
Reports are given on diversity and distribution, coexistence, habitats and
shelters, and where possible notes are given on behaviour. A regional key was
constructed for field or laboratory use.
DESCRIPTION OF THE STUDY AREA
Location
Table Mountain forms the northern end of the mountainous ridge of the
Cape Peninsula and lies approximately 33°57’S 18°25’E.
Topography, geology and soils
Table Mountain forms a plateau of Table Mountain Sandstone, which is
surrounded by cliffs giving way to gentle slopes. The southern end is deeply
dissected by the Disa stream forming the Orangekloof subarea. The sandstone
overlies a layer of Malmesbury shale which in turn rests on Cape Granite. The
last two formations give rise to the gentle slopes (Moll & Campbell 1976), the
areas with which this study is primarily concerned. Soil types are dependent
on the parent material, and those derived from the sandstone are usually
acidic sands, the granite and shale derived soils contain more clay and are less
acidic. The lower slopes are covered with sandstone debris which give rise to
soils of mixed origin (Moll & Campbell 1976).
Climate
Table Mountain lies in the Winter Rainfall Region characterized by hot,
dry summers and cool, wet winters. The topography plays an important part
in the climate. Rainfall for the different subareas are as follows:
Summit and eastern slope . : . 1420-1780 mm
Western, northern slopes and Siena Hill RN: 889 mm
Southern subarea : ; ; 1227 mm
(from Moll & Campbell 1976)
Temperature differences over the whole area are not as pronounced.
Vegetation
The most extensive plant community is macchia or fynbos, consisting of
Proteaceae, Ericaceae, Restionaceae, Geraneaceae and other families. There
are also small areas of natural forest communities as well as extensively
cultivated alien forest vegetation consisting of Pinus spp., Eucalyptus spp.,
and Acacia spp. mainly on the eastern slopes and lower northern slopes.
There are also areas of Hakea sp. on the western slopes. The classification
of plant communities is given in Figure 1.
MATERIALS AND METHODS
Six study subareas were defined:
1. North-facing slopes 4. East-facing slopes
2. West-facing slopes 5. Plateau subarea
3. Southern subarea (Orangekloof) 6. Signal Hill subarea
231
NOTES ON THE SCORPION FAUNA OF THE CAPE
+7404
eae
ce
+t
fe:
2
oa
-
+
aoe
fg
2
eS
Dry proteoid
Oa
=)
—
LS
)
(eo)
—J)
Short restoid
Alien forest
Map of Table Mountain showing approximate boundaries of plant communities.
Big: 1.
Each subarea was divided into a number of transects, 0,5 km wide, extend-
ing from the base of the mountain to the sandstone cliffs. The table or plateau
not divided up in this way.
Was
soil
b)
logs and other debris, and search-
9
Collecting was carried out by moving up a transect and collecting samples
Records were kept of altitude
by turning over surface objects such as rocks
ing under the loose bark of dead trees.
types and vegetation.
D3) ANNALS OF THE SOUTH AFRICAN MUSEUM
Several areas were isolated for the more detailed studies of demes or popula-
tion aggregates of Opisthophthalmus and Uroplectes. In these areas burrow
morphology, run morphology, feeding and coexistence of species were studied.
In order to observe the feeding and burrowing behaviour of O. capensis, studies
were carried out in the field where possible. In addition a number of specimens
were kept in specially prepared boxes in the laboratory to complement field
observations of feeding and burrowing. The morphology of burrows was studied
by excavation and measurement. The numbers of observed specimens in relation
to vegetation types is given in Table 1.
TABLE |
Comparison of relative numbers of scorpions observed in each of the six study subareas and
seven vegetation types.
O. capensis U. lineatus U. insignis U. variegatus
Study subarea Vegetation type ©=£_ TT
oa 2 oF < § © 6 1g
North- and west- Dry proteoid OS) Zi Dap Bil S54:
facing slopes, and a
Signal Hill . . . Lowscrub WS) 3 DS
South-facing
slopes and
Plateau subareas . Tall proteoid ad Sy
Plateau subarea . Short restoid 3
East-facing Moist proteoid Dnwe'S
slopes subarea. . _- Se ———
Forest scrub-
forest Ye 1728
North- and east-
facing subareas . Alien forest I
Rotale 3 119 156 45 9
DISTRIBUTION AND HABITAT
One scorpionid species, viz. Opisthophthalmus capensis, was found on Table
Mountain, while other recorded species were buthids. O. capensis has also
been found on the lower-lying Cape Flats and recorded randomly in the Cape
Peninsula. This and allied species extend into Namaqualand (Eastwood 1977a).
Another species, O. macer Thorell, extends from False Bay eastwards to Port
Elizabeth (Lawrence 1955). Parabuthus capensis (Ehrenberg) extends from Clan-
william in the north to Cape Town, and eastwards to Tulbagh (Eastwood 19775).
Another widely distributed species, P. granulatus (Hemprich & Ehrenberg), is
found on the Cape Flats to the north and has recently been collected in the
suburban areas north of the city. Uroplectes lineatus is reported by Lawrence
(1955) to extend from Cape Town eastwards to Mossel Bay, and northwards
as far as Tulbagh and Worcester. U. insignis is confined to Table Mountain.
U. variegatus, another local species, is recorded from the Cape Peninsula and
its immediate neighbourhood, and Langebaan in the north (Lawrence 1955).
In the present survey of the Table Mountain study area, O. capensis was
located all along the west-facing subarea, the Signal Hill subarea and the
NOTES ON THE SCORPION FAUNA OF THE CAPE 233
@ U. lineatus
O U. insignis
@ U. variegatus
O O. capensis
0
V. BRANCO
Fig. 2. Map of Table Mountain showing distribution of scorpions collected during the survey.
western extremity of the north-facing subarea. There was a tendency for demes
to be found in fairly open areas of the dry proteoid communities, i.e. below the
400 m contour, where the slopes were not too steep and ground cover plentiful,
but they also occurred in the low scrub communities where the ground cover
and substratum were suitable (Fig. 3).
U. lineatus is the most widely distributed scorpion on Table Mountain
(Fig. 2). As well as occupying widely differing habitats it was found in all the
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
me
Fig. 3. Typical habitat of Opisthophthalmus capensis, Uroplectes lineatus and U. variegatus.
Inthe foreground is the dry proteoid plant community and further back the lowscrub community.
subareas under study, and at varying altitudes where ground cover was adequate.
The demes were not as obvious as those of O. capensis and in many cases only a
few specimens could be found in a large area. This could be accounted for by
the immovability of much of the ground cover, or thick scrub and grass, making
collection difficult. Thus U. /ineatus was found in the dry proteoid, low scrub,
tall proteoid, moist proteoid, short restoid and forest scrub-forest plant com-
munities. This species appeared to be most abundant in the dry proteoid
communities on the north- and west-facing slopes. Only one specimen was
found in alien forest. It will adapt to many situations as shown by the range of
habitats it occupies, and is often reported to be found in the homes of residents
near Table Mountain and even at the 600 and 700 m contours where access
for the collector is allowed only by traverses along rocky ledges.
Specimens of U. variegatus were found in the same areas as occupied by
O. capensis in the dry proteoid communities (Fig. 2).
U. insignis is taxonomically very near U. lineatus. Purcell (1902) regarded
it as a distinct species. He recorded specimens from the Constantia and Newlands
slopes of Table Mountain (east-facing subarea). In the present survey two
substantial demes were located (Fig. 2). These were situated in the forest and
scrub forest communities which dominate the east-facing subarea. Specimens
were collected from under stones as well as from under the dried bark of Pinus
pinaster and P. radiata stumps in areas which are being cleared of alien vege-
tation (Fig. 4).
NOTES ON THE SCORPION FAUNA OF THE CAPE py)
hil
Fig. 4. Typical habitat of Uroplectes insignis and U. lineatus in the forest scrub-forest
community of the east-facing study subarea.
Fig. 5. Run and burrow of Opisthophthalmus capensis. The antechamber is absent in this
particular run.
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
No specimens of Parabuthus capensis were found in the present survey.
There are several specimens in the collection of the South African Museum
collected from Devil’s Peak (north-facing subarea) which lies in the dry proteoid
community.
SCORPION SHELTERS
Opisthophthalmus capensis was always found to construct its burrow beneath
some surface object. The excavation leading from the opening under the stone
or log to the burrow entrance is the run. The structure of the run is dependent
on the size and shape of the ground cover and nature of the substratum (Fig. 5).
Thus the run varies in length and sometimes has two or even three openings.
Just before the burrow entrance proper there is usually a bulge or separate
section which could be called an ‘antechamber’ (Fig. 6B). Often food remains
are found in this area and its function appears to be a feeding area, and possibly
affords freer movement for the scorpion to turn or employ its sting. Excavation
of some fifty burrows sometimes revealed food remains in the terminal chamber
of the burrow itself. The burrow is usually a simple tunnel having a charac-
A
Canis) QrOUndlCOVeG.N v.ls 0. 2) Sia fans
_—_—_—__——~___ = +> burrow entrance
ruin entrance : : : :
terminal chamber
—
direction of slope
iS
. see eae
antechamber - - - °. \
|
Fig. 6. A. Cross-section. B. Plan of typical run and burrow of Opisthophthalmus capensis.
NOTES ON THE SCORPION FAUNA OF THE CAPE Di.
teristically oval entrance penetrating the ground at about 30° from the hori-
zontal (Fig. 6A). It often turns to right or left, if there are obstacles in the way,
and ends in a very slightly enlarged terminal chamber. Variation is due to the
nature of the substratum, size or sex of the inhabitant. Males construct shallower
burrows or none at all, living only in a run. Only one specimen occupies each
burrow. Ground cover, usually sandstone debris lying on soils of mixed origin,
often sandy loam, under which burrows were found varied from 15 to 60 cm in
diameter. If the stones lay on a slope the run entrances always faced downhill,
ensuring dry runs during rainy weather. Table 2 gives the measurements of
fifty burrows.
TABLE 2
Measurements in centimetres of fifty burrows of Opisthophthalmus capensis (36 33, 14 99)
Min. Max. Mean
Run length g ‘ : : 3 ; 2 9,0 27,0 V2,
2 ; : : ; 10,0 22,0 15,3)
Run width ¢ f ; : ; ; : 2,6 4,0 3) 2
Q : : ; : : : DS 3,4 3,1
Burrow entrance height ¢ . 4 : f 1,8 Bal 2,9
Q . ; : ; 2,0 3,6 By Il
Burrow entrance width ¢. f : ; Dell 32 2,9
Ons : : : Dep 3,0 2,8
Burrow length ¢ . p : ; ; 0,0 20,0 10,6
Oe : ; 3 : ; 10,0 21,0 19,8
Vertical depth of burrows 5. : ; 355) 6,5 S77
Orie ; : 6,0 13,5 11,1
Uroplectes lineatus occupies a wide range of shelters which may be classified
as follows:.
1. Beneath suitable stones—no evidence of any run construction, but where
concavities form a natural shelter.
Beneath stones where a run has been constructed.
Beneath debris such as logs and dried organic matter.
In rock crevices, or under stones lying on a flat rock surface.
Beneath tree bark.
A deme of U. lineatus in the south-facing subarea was found suitable for
a study of run morphology. The structure of the run is dependent on the nature
of both ground cover and substratum (Figs 7-8). The runs vary greatly in
length and width (Table 3) and may have one or two entrances.
U. lineatus in the east-facing subarea had the same shelters as U. insignis,
namely the space between dried bark and stumps of pine trees (Fig. 9), although
the two species were not found together under the same cover. Where sandstone
he eS)
TABLE 3
Measurements in centimetres of thirty runs of Uroplectes lineatus.
Min. Max. Mean
Length of run 5 : : : ; : 355 OES) 11,3
Width of run : 5 : ; 0,8 22 1,4
Ground cover size (diameter) Y , ; 9,0 55,0 31,0
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
\
|
Fig. 7. Diagrams showing variation of runs or scrapes of Uroplectes lineatus.
Fig. 8. Run of Uroplectes lineatus. Usually the runs are not as well defined as this one.
NOTES ON THE SCORPION FAUNA OF THE CAPE 239
Fig. 9. A section of bark removed from a pine stump, showing a specimen of Uroplectes
insignis, several isopods and a longicorn beetle.
debris formed the ground cover, only one specimen of U. lineatus or U. insignis
was found under each stone, whereas up to nine specimens of these species
were collected from a single stump. One specimen of U. /ineatus was found in a
shallow, wide burrow in the north-facing subarea and not covered by any
surface object. Presumably the burrow was excavated by another animal.
U. variegatus was found to occupy narrow runs, usually well defined with
a small terminal chamber. Some specimens were also found in natural depres-
sions beneath stones. Table 4 gives measurements of five runs of U. variegatus.
TABLE 4
Measurements in centimetres of five runs of Uroplectes variegatus.
Min. Max. Mean
Length . “ ; ; s : : : 4,0 13,0 9.4
Width . : d , : : 4 ; 0,75 1,4 0,92
Ground cover size (diameter) . ; : 16,0 48,0 37,0
Parabuthus capensis was not collected during the survey, but has been
collected by the author from other parts of the Cape. This species often excavates
runs which vary according to the nature of ground cover and substratum and
usually consist of a more or less round or oval space beneath the ground cover
with one or two entrances.
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
COEXISTENCE
The following pairs of species were found to coexist: Opisthophthalmus
capensis— Uroplectes lineatus, O. capensis—U. variegatus, U. lineatus—U.
insignis. In field studies demes or population aggregates were roughly delineated
by markers to determine the spatial distribution of their shelters.
Of eight demes of O. capensis, five showed a spatial overlap with demes of
U. lineatus (Fig. 10) in studies carried out in the west-facing subarea. Specimens
of U. variegatus were recorded in association with O. capensis in two of these
demes where U. lineatus was absent. In one instance a specimen of U. variegatus
had its run under the same rock as O. capensis, their respective entrances
Opening on opposite sides. |
400m _ contour
low scrub
dry proteoid
@ U.lineatus
©. O.capesis
Fig. 10. Diagram of population aggregates of Opisthophthalmus capensis and Uroplectes
lineatus in the west-facing study subarea, showing spatial overlap.
In a study of coexistence of scorpions by Williams (1970) two mechanisms
were found to permit coexistence: the choice of different-sized prey, and habitat
specialization. The first mechanism undoubtedly operates in the case of O.
capensis— U. lineatus and O. capensis—U. variegatus.
U. insignis and U. lineatus were found under the bark of pine stumps in
the same immediate vicinity, but specimens of either species did not share the
same shelter. The mechanism permitting coexistence in this case is not quite
‘clear.
The record of Parabuthus capensis from Devil’s Peak indicates the prob-
ability of the coexistence of this species with U. lineatus. P. capensis from the
Citrusdal district has been found in association with U. carinatus (Pocock)
and O. pallidipes Koch.
NOTES ON THE SCORPION FAUNA OF THE CAPE 241
SCORPION PREDATORS
Because of the nocturnal habits of scorpions, predation is not often
reported. The author has seen a troop of baboons on the northern slopes of
Table Mountain foraging beneath stones. Baboons (Papio ursinus (Kerr)) are
well-known predators of scorpions. The bokmakierie (Telophorus zeylonus (Linn.))
and the fiscal shrike (Lanius collaris Linn.) have been reported to prey on
Uroplectes lineatus (1. G. Taylor pers. comm.). Owls are undoubtedly the
commonest predators of scorpions. The spotted Eagle Owl (Bubo africanus
(Tem.)), a local species, is reported to feed on scorpions (Roberts 1940). The
Barn Owl (Tyto alba (Scopolt)) feeds on buthids and scorpionids (Vernon 1972).
SOME OBSERVATIONS ON BEHAVIOUR
The behaviour exhibited in burrow construction has been discussed by
Williams (1966) for the Neotropical species Anuroctonus phaeodactylus (Wood)
(Vejovidae), and Newlands (1972) reported on the burrowing behaviour of
Protophthalmus holmi Lawrence, Opisthophthalmus concinnus Newlands and
O. flavescens Purcell, all Ethiopian Scorpionidae.
An important adaption of Opisthophthalmus capensis is its ability to con-
struct burrows. In the laboratory specimens made burrows by loosening tightly-
packed soil with a biting action of the chelicerae. The loose sand was then
scraped out backwards by legs I and II which were tucked under the body.
Legs III and IV provided the traction while the pedipalps were extended
anteriorly and acted as supports to prevent the body tipping forward and
occasionally as levers which aided in the backward movement. The cauda
was extended straight out behind the body. The soil was transported rapidly
backward in a single movement and the soil deposited outside the entrance on a
tumulus by a slight flick of the first two pairs of legs so that the soil sprayed out.
Occasionally the scorpion moved its load of soil by moving backward for a few
centimetres, stopping, and thus pulling the soil backward with each step. With
each step there was a pause while the hind legs were moved backward again
to repeat the process. Thus two methods were employed to transport the soil:
the first was used in exceptionally dry soil where backward movement was
smooth and unimpeded; the second method was employed when the soil was
damp and therefore more difficult to move, or if the burrow was steep or filled
with small stones, twigs and similar obstacles. Several specimens did not burrow
but made runs under stones provided for this purpose. Initially the animal
would remove the soil in the usual manner as well as forming a broad chamber
beneath the stone by pushing the soil with the tail. The curved tail was laid flat
on the surface and then extended, thus moving a good volume of soil each time.
This tail scraping is reported as being a widespread phenomenon among
scorpions (Newlands 1972). It is also reported as a part of burrow maintenance
in Anuroctonus phaeodactylus (Wood) by Williams (1966).
Field observations showed that immature specimens were adept at burrow-
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
ing. Several second or third instar nymphs were found in burrows of 1-2 cm
in length under stones. These burrows rarely had the characteristic oval opening
and were often nothing more than shallow depressions.
The tarsi of O. capensis are adopted for a pelophilous habitat, that is
the claws are sharply curved and tarsal setae are not as dense as in O. wahlbergi
(Thorell) for example, a psammophilous species. Caudal segment V is heavily
sclerotized with numerous setae and well-developed keels to facilitate tail-
scraping. Collections made in the field during the winter months (June—Sep-
tember) indicate that these scorpions remained deep in their burrows on cold
days, but if the sun was hot they were often found at the entrances of their
burrows or in the runs, facing outward. This “doorkeeping’ is presumably
maintained while the scorpion waits for prey which may seek refuge under
stones (Alexander & Ewer 1958). The tail was curled around sideways, parallel
to and resting on the substratum so that the telson, which was folded over the
dorsal surface of caudal segment V, lay against the lateral side between the
sixth and seventh mesosomal segments. From this position the metasoma could
easily be brought into action sideways, as had been observed in the laboratory.
This sideways movement is obviously more suitable for stinging prey while
the scorpion is in its run. The pedipalps were arranged so that the femora were
perpendicular to the body and the tibiae and chelae lay parallel to the body.
The movable and immovable fingers were kept about 5 mm apart. If the scorpion
was just inside the burrow entrance, the chelae may have protruded and were
arranged as above. Sometimes the chelae were folded over each other with the
dorsal and outer sections facing outwards, forming a sort of shield. This is a
typical defensive stance for Scorpionidae (Newlands 1969). When a stone was
lifted while a scorpion was in its run it would lift its tail so that the telson lay
above the first caudal segment and the body was lifted off the substratum. If
further aggravated it would adopt an aggressive stance so that the last two
mesosomal segments were curved upward, the metosoma perpendicular to the
ground except for the last caudal segment which was held horizontal with the
telson above the last mesosomal segment or carapace. Another reaction was
retreat, in which case the cauda was brought down backwards and the animal
scuttled down its burrow, either backward or forward.
The function of the doorkeeping seems to be related to feeding, where
the scorpion waits for prey to come into the run. At no time was it observed
that the scorpion was an active predator. However, active foraging in the
vicinity of burrows is known for Opisthophthalmus (G. Newlands, pers.
comm.). Feeding was observed in the field in the runs or antechambers.
Feeding behaviour has been discussed for Neotropical scorpions by Baerg
(1954) and Hadley & Williams (1968) (Buthidae and Vejovidae) and for the
South African Opisthophthalmus latimanus by Alexander (1972). In the field
O. capensis was observed feeding on Dorylinus helvolus (Formicidae) and
Temnopteryx phalerata (Blattariae). In the laboratory a wide range of prey
species were accepted and eaten, while others were ignored. Prey species which
NOTES ON THE SCORPION FAUNA OF THE CAPE 243
were accepted were Periplanita americana, Temnopteri phalerata (Blatteriae),
Dorylinus helvolus (Formicidae), Holopternia valga (Hemiptera) as well as
U. lineatus, isopods, Scarabaeidae, beetle larvae, moths and moth larvae. The
following species were either killed or ignored but not eaten: Chilopoda,
Diplopoda and Anthia spp. (Carabidae).
During excavation of burrows food remains in the form of pellets were
sometimes found in the terminal chambers or in the runs. The food remains
were identified as small tenebrionid beetles and cockroaches from the indigestible
elytras and legs. Thus feeding occurred both in the burrows and the runs.
Feeding behaviour was elicited in the laboratory by placing the prey near the
scorpion which usually rests in the typical doorkeeping posture. Small prey
was firmly grasped in both chelae and the metasoma brought upright. Small
prey was not stung and the biting action of the chelicerae began immediately.
If the prey was lively but not very large the scorpion brought the telson into
play from the side and usually slowly, seeking for a soft spot in which to inject
the venom. If the prey was large and aggressive the scorpion initiated offensive
behaviour and the telson was brought swiftly over the carapace to deliver a
series of stings. The ingestion process lasted from | to 3 hours in the laboratory,
depending on the size of the prey. The formation of pellets of food remains
was not observed. Ingestion started immediately after capture and in heavily
sclerotized prey the starting point was ventrally between head and thorax. The
chelicerae moved alternately in and out while the fingers grasped at the food,
giving rise to the typical biting action.
Sponge-bathing activities were not observed after feeding, nor after burrow-
ing in wet soil when the legs and chelicerae became dirty.
In humid laboratory environments O. capensis was never seen to approach
the dish of water in the cage and it seems likely that water balance is maintained
by contact with a wet substratum as reported by Hadley & Williams (1968) in
observations at night of desert scorpions. If the cage was kept very dry and
water placed in the cage the scorpions would climb over the edge of the dish
and dip the chelicerae and pedipalps into the water. At the same time the pedi-
palps moved rhythmically up and down.
Uroplectes lineatus adopted a very characteristic resting posture while
under ground cover. The pedipalps were usually held so that they lay slightly
anterior to or lateral to and subparallel to the carapace. The tail was curved
over the abdomen dorsally so that the second caudal segment was horizontal
to the body and the terminal segments curved around sideways and the telson
lay laterally between mesosomal segments V and VI, being folded over against
caudal segment V dorsally. In the defensive posture the tail lay behind the body
with the telson curved forward so that it lay above the first caudal segment.
When agitated this scorpion can strike very rapidly to the side, forward or
backward. This species did not use the chelae to grasp an aggressor but only
struck out with the tail. In the aggressive posture there was a slight forward
stilting. Venom-dripping was associated with aggressive behaviour. In the
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
laboratory it was noticed that U. lineatus was able to excavate a run, but the
behaviour of digging was not observed. In the east-facing subarea specimens
were found in association with Isopoda, geckos, longicorn beetles, Lagria
villosa F. (Lagriidae), Pyrrhocoridae, Hemiptera, cockroaches (Temnopteryx
phalerata (Sauss)) and U. insignis. In this environment both U. lineatus and
U. insignis were observed feeding on cockroaches. Prey capture was not observed
here or in the laboratory.
Two pregnant female specimens of U. insignis and one of U. lineatus were
kept in the laboratory. Parturition was observed in both species which behaved
similarly. Parturition behaviour was as follows: the scorpion stilted on the two
last pairs of legs prior to delivery; the first two pairs were held under the body
so that the distal segments were parallel to the body. This posture was main-
tained throughout parturition. The young began passing through the genital
opening head first, one by one into the ‘birth basket’ formed by the first two
pairs of legs. The young became active after a few minutes and began ascending
the mother’s back by way of her walking legs. When all the young had reached
the back they assumed a completely random orientation, sometimes in several
layers, which continued to the first moult. This random orientation of first
instar young is characteristic of the Buthidae (Williams 1969) (Figs 11-12).
With the onset of the second instar stage the litter became distinctly more
elongate. Duration of first instar stage of U. insignis was 9-12 days with litters
of 12 and 13 specimens, and duration of first instar of U. lineatus was 11 days
with a litter of 8 specimens. Parturition took place from January to February.
KEY TO THE SCORPIONS OF TABLE MOUNTAIN
1. Chelae slender; the dorsal surface not separated by a keel (Family
Buthidae). Found in simple runs or scrapes under stones or beneath
(mee oe so a 2
— Chelae large, the dorsal curiite Renaitedl into inner anal er bee ione
by a smooth keel (Family Scorpionidae). Found in burrows, usually
opening under stones oe ie Opisthophthalmus capensis (Herbst)
2. Species large; stridulatory areas on dorsal surface of caudal segments
I and II well developed. Posterior dorsal surface of caudal segment II
raised to form a subtriangular lip Parabuthus capensis (Ehrenberg)
Species small; stridulatory areas on cauda absent .. oe me 3
3. Tergites with short lateral keels and seven black longitudinal lines
Uroplectes variegatus (Koch)
— Tergites without lateral keels, darkly pigmented Nor iReoay its 4
4. Tubercle below aculeus distinctly laterally compressed; vesicle
yellow a 5 Uroplectes lineatus (Koch)
— Tubercle below neulene cones bal blunted; vesicle black
Uroplectes insignis Pocock
NOTES ON THE SCORPION FAUNA OF THE CAPE 245
Maly
Fig. 11. First instar of Uroplectes insignis. Note the random orientation typical of the
Buthidae.
Fig. 12. Second instar of Uroplectes insignis. The young are beginning to leave the mother’s
back.
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
CONCLUSIONS
Four scorpions species were found on Table Mountain and Signal Hill. Two
species, Opisthophthalmus capensis and Uroplectes lineatus, were very abundant
whereas the other two were rare (U. insignis and U. variegatus). O. capensis
was abundant on the western north-facing subarea, the greater proportion
occurring in the dry proteoid plant community below the 400 m contour.
Fewer specimens occurred in the low scrub communities since these areas are
very steep and the substratum and ground cover unstable during the rainy
season. U. lineatus was found to be fairly abundant in all subareas studied and
has a random distribution, occurring in varied habitats. U. variegatus and
U. lineatus were found respectively to coexist with O. capensis, although these
two former species were not found to coexist. The fact that all the young pro-
duced by two females of U. insignis had the characteristics of this species
suggests that it is a true-breeding species which coexists with U. lineatus.
Apparently isolating mechanisms have been acquired for this example of
sympatric association, but what these are, is not quite clear.
The notes on the behaviour make comparison with other species possible.
In the burrowing behaviour of O. capensis two methods are employed to
transport the soil; the first is similar to the behaviour shown by Protophthalmus
holmi Lawrence (as reported by Newlands (1972)), i.e. the soil is transported
rapidly backward in a single movement, and the second is like that of O. flaves-
cens Purcell and O. concinnus Newlands, 1.e. the soil is transported backward by
a series of jerks. Observations of laboratory specimens of O. capensis revealed
that the first method was used in exceptionally dry soil where backward move-
ment was smooth and unimpeded. The second method was employed when the
soil was damp and therefore difficult to move, or if the burrow was steep or
filled with small stones, twigs or similar obstacles. Thus O. capensis, which is
pelophilous, exhibits burrowing behaviour similar to psammophilous scorpions
in certain conditions. The excavation of a run by Parabuthus capensis was
described by Eastwood (19775). Part of this process was similar to the behaviour
shown by O. capensis when obstacles had to be removed, but loose sand was
excavated in a manner characteristic to Parabuthus. The defensive behaviour
of O. capensis is the same as the generalized behaviour for Opisthophthalmus as
discussed by Newlands (1969). The resting posture of O. capensis prior to
feeding differs from that of vejovid scorpions in the position of the metasoma.
In O. capensis the metasoma is usually curved forward alongside the body
whereas the vejovid scorpions discussed by Hadley & Williams (1968) held the
metasoma straight out backwards. Feeding behaviour of O. capensis was
similar to that observed for Parabuthus capensis, except that the latter was more
reluctant to attack larger and more active prey.
Birth activities of Neotropical Buthidae were discussed by Williams (1969)
and this was found to be similar for U. insignis and U. lineatus. The first instar
young of these two species showed the random orientation on the mother’s
NOTES ON THE SCORPION FAUNA OF THE CAPE 247
back which was reported for the genus Centruroides (Williams 1969).
Because of the position of Table Mountain, the human impact on its ecology
is considerable. Frequent fires cause damage to the floral and faunal ecology.
In this study it was observed that in recently burnt areas O. capensis was abun-
dant, where U. lineatus was more abundant in adjacent unburned areas. It is
obvious that O. capensis is able to withstand a veld fire because of its ability
to burrow, whereas a local deme of Uroplectes can be exterminated. The lower
slopes of the north-facing and east-facing subareas are covered in plantations
of alien forest. In these areas there is very little insect life and consequently
few scorpions were found.
ACKNOWLEDGEMENTS
I thank Dr V. Whitehead for advice and encouragement during this project,
and the Director of the South African Museum for providing research and
transport facilities. For constructive comments and advice on this and other
projects I thank Mr B. Lamoral of the Natal Museum and Mr G. Newlands
of the South African Institute for Medical Research.
Thanks are also due to Dr V. Whitehead, Messrs A. Byron and S. Rice
for help with the photographic work, and the Cape Town City Engineer’s
Department and the State Department of Forestry for permits and permission
to collect on Table Mountain.
REFERENCES
ALEXANDER, A. J. 1972. Feeding behaviour in Scorpions. S. Afr. J. Sci. 68: 253-256.
ALEXANDER, A. J. & Ewer, D. W. 1958. Temperature adaptive behaviour in the scorpion
Opisthophthalmus latimanus Koch. J. exp. Biol. 35: 349-359.
BAERG, W. J. 1954. Regarding the biology of the common Jamaican scorpion. Ann. ent.
Soc. Am. 47: 272-276.
Eastwoop, E. B. 1977a. Notes on the scorpion fauna of the Cape. Part 1. Description of
neotypes of Opisthophthalmus capensis (Herbst) and remarks on the O. capensis and
O. granifrons Pocock species-groups (Arachnida, Scorpionida, Scorpionidae). Ann. S. Afr.
Mus. 72: 211-226.
Eastwoop, E. B. 1977b. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus
capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida,
Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214.
HADLEY, N. F. & WILLIAMS, S. C. 1968. Surface activities of some North American scorpions
in relation to feeding. Ecology 49(4): 726-734.
Hewitt, J. 1918. A survey of the scorpion fauna of South Africa. Trans. R. Soc. S. Afr. 6:
89-192.
KRAEPLIN, K. 1894. Revision der Skorpione. Jb. hamb. wiss. Anst. 12: 75-96.
KRAEPLIN, K. 1899. Scorpiones und Pedipalpi. Das Tierreich 8: 1—265.
KRAEPLIN, K. 1908. Skorpione und Solifugen. Ergebnisse e. Forsch. Siidafrika Bd 1, Lief 2.
Denksch. med.-naturw. Ges. Jena 13: 247-282.
LAWRENCE, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South
African families, genera and species. S. Afr. anim. Life 1: 152-262.
Mo Li, E. J. & CAMPBELL, B. M. 1976. Table Mountain. A conservation and management
report. Cape Town: University of Cape Town.
NEWLANDS, G. 1969. Scorpion defensive behaviour. Afr. wild Life 23: 147-153.
NEWLANDS, G. 1972. Notes on psammophilous scorpions and a description of a new species
(Arachnida: Scorpionides). Ann. Transv. Mus. 27: 241-254.
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
PeNTHER, A. 1900. Zur kenntnis der Arachnidenfauna Sitidafrikas (Scorpiones). Annln. naturh.
Mus. Wien 15: 153-163.
Pocock, R. I. 1890. A revision of the genera of scorpions of the family Buthidae, with descrip-
tions of some South African species. Proc. zool. Soc. Lond. 1890: 114-141.
Pocock, R. I. 1896. A further revision of the species of scorpions belonging to the South
African genera Uroplectes, Lepreus and Tityolepreus. Ann. Mag. nat. Hist. (6) 17: 377-393.
PURCELL, W. F. 1899. On the species of Opisthophthalmus in the collection of the South African
Museum, with descriptions of some new forms. Ann. S. Afr. Mus. 1: 131-180.
PuRCELL, W. F. 1901. On some South African Arachnida belonging to the orders Scorpiones,
Pedipalpi and Solifugae. Ann. S. Afr. Mus. 2: 137-225.
Roserts, A. 1940. Birds of South Africa. Revised by McLachlan, G. R. & Liversidge, R.
3rd ed. 1970. The Trustees of the John Voelcker Bird Book Fund.
VERNON, C. J. 1972. An analysis of owl pellets collected in southern Africa. Ostrich 43: 109-124.
WILLIAMS, S. C. 1966. Burrowing activities of the scorpion Anuroctonus phaeodactylus (Wood)
(Scorpionida: Vejovidae). Proc. Calif. Acad. Sci. 34: 419-428.
WILLIAMS, S. C. 1969. Birth activities of some North American scorpions. Proc. Calif. Acad.
Sci. 37: 1-24.
WILLIAMS, S. C. 1970. Coexistence of desert scorpions by differential habitat preference.
Pan-Pacif. Ent. 46: 254-267.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon. ~
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
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In describing new species, One specimen must be designated as the holotype; other speci-
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must be recorded, e.g.:
Helciype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; ‘.. . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. 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 be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
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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.
E. B. EASTWOOD
NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 3
SOME OBSERVATIONS ON THE DISTRIBUTION
AND BIOLOGY OF SCORPIONS ON
TABLE MOUNTAIN
: Oy es
VOLUME 74 PART 11 JANUARY 1978 ISSN 0303-2515
: MUS. COMP. ZOOL
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APR 28 1978
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634. \
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of:Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 +#&Band
January 1978 Januarie
Part 11 Deel
Yia,, S
AU /0Ui0. NONI spas
NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 4
THE BURROWING ACTIVITIES OF SOME
SCORPIONIDS AND BUTHIDS
(ARACHNIDA, SCORPIONIDA)
By
E. B. EASTWOOD
Cape Town Kaapstad
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NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 4
THE BURROWING ACTIVITIES OF SOME
SCORPIONIDS AND BUTHIDS (ARACHNIDA, SCORPIONIDA)
By
E. B. EAsTwoop
South African Museum, Cape Town
(With | figure and 2 tables)
[MS. accepted 23 August 1977]
ABSTRACT
Burrowing activities of the scorpionid Opisthophthalmus macer Thorell were studied in
the field and laboratory, and comparisons made with O. capensis (Herbst). O. karrooensis
Purcell, O. pallidipes Koch and O. peringueyi Purcell were found to burrow only in their
immature stages. The morphological adaptions of the adults are discussed in relation to their
scrape-dwelling habit. The specialized digging behaviour of the buthid Parabuthus planicauda
(Pocock) is described. Pregnant females were often found to construct proper burrows prior
to parturition.
CONTENTS
PAGE
IiEKOGUCHIONS 9 oe ee we a bat ie Pte ae, Pale DAD
DISCUSSION eee eee css sae A OS oh el OO
GOnclusiGns) ten) we oe eee ah agar a 25S
ACknOWwledSementS es gu.) sacs.) te a el 254
IRNGLGKENCES Sr eae oe et ee a ee 4
INTRODUCTION
Opisthophthalmus macer Thorell is a true burrowing scorpionid, distributed
from False Bay to Port Elizabeth along the south coast. In this paper the burrow
morphology and burrowing behaviour of this scorpion are reported and com-
pared to the corresponding activities of O. capensis (Herbst).
O. karrooensis Purcell, O. pallidipes Koch and O. peringueyi Purcell all
occupy the drier semi-desert regions of the Cape, namely the Karoo and
Namaqualand southwards to Piketberg. The young of these species are true
burrowers, whereas the adults have in common a scrape-dwelling habit with
special morphological adaptions for this particular way of life.
Digging behaviour has been reported in the Buthidae and shows certain
similarities to Parabuthus planicauda (Pocock) and P. capensis (Ehrenberg).
This was illustrated by Pavlovsky (1924) for the transcaspian species Liobuthus
kessleri Birula, where the first three pairs of walking legs were used for excava-
tion. Lankester (1882) reported on the digging behaviour of the Egyptian
species Buthus australis citrina (Hemprich & Ehrenberg). In this case the
249
Ann. S. Afr. Mus. 74 (11), 1978: 249-255, 1 fig., 2 tables.
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
pedipalps aided the first three pairs of legs in digging operations.
P. planicauda is a widely distributed buthid of the southern Cape, extending
from Worcester and Tulbagh in the west to Albany and Cathcart in the east
(Lawrence 1955). Under certain conditions this species may construct true
burrows. The specialized digging behaviour of this species is discussed in
relation to the activities of P. capensis (Ehrenberg).
DISCUSSION
Opisthophthalmus macer Thorell
Two demes of this species were studied. The first was found on the southern
slopes of the Kleinriviersberge in a habitat consisting of macchia, namely
Proteaceae, Restionaceae, Ericaceae, Geraniaceae and other families. Available
ground cover was sandstone and quartzite debris and the substratum was deep,
sandy loam rich in organic matter.
O. macer was found to burrow only beneath suitable ground cover.
Typically a run was constructed with a single entrance, leading to the burrow
opening. Often an antechamber was present, the probable function of this
being a feeding area. The burrows of females were invariably longer than those
of males and often males were found to construct only a run. Table | gives the
measurements of twenty burrows and runs.
The second deme was located next to the sea in the southern part of
Walker Bay. This habitat consisted of rocky limestone slopes with very little
soil and low coastal scrub. In this habitat the burrows were generally shorter
and none were found with an antechamber in the run. Specimens were also more
darkly coloured. Table 1 gives the measurements of ten burrows and runs of
the second deme.
Burrowing behaviour of O. macer was observed as follows: compacted
soil was loosened by the action of the chelicerae and scraped into a heap by
the first two pairs of walking legs. The heap was then moved backward by legs
I and IJ, which were tucked beneath the body. Legs III and IV provided the
traction while the tail was extended, the pedipalps being used as supports.
The tarsal claws of O. macer are more sharply curved than those of O. capensis.
According to Newlands (1972) this is related to a pelophilous habit. This view
could not be substantiated in the present study, since the Kleinriviersberge deme
was located in sandy loam and the deme from the coastal habitat was found
to burrow in compacted sand, whereas O. capensis from the Cape Peninsula
burrowed in sandy loam only. Tail scraping was not observed although the
heavily sclerotized fifth caudal segment of O. macer is adapted for this function.
The scrape-dwellers, Opisthophthalmus karrooensis Purcell, O. pallidipes Koch
and O. peringueyi Purcell
Observations of a deme of O. karrooensis showed that although the adults
were found to occupy scrapes or runs, or simply depressions beneath stones,
NOTES ON THE SCORPION FAUNA OF THE CAPE 251
TABLE |
Measurements in centimetres of twenty burrows and runs of Opisthophthalmus macer in
a habitat of Cape macchia (Deme 1), and ten burrows in a limestone and coastal scrub
habitat (Deme 2).
Deme | Deme 2
Min. Max. Mean Min. Max. Mean
Run length 8,0 19,0 16,5 6,0 10,0 7,0
3,0 14,0 10,2 Ds) 7,0 5,0
Run width DS) 3) 3,0 Dp 3,0 2,6
Burrow entrance height .
Burrow entrance width .
Burrow length
Vertical depth of burrow
+O Os +O Oy +O Os =} OOS tO MON +0 Oy
ww
i)
ww
ey
we
N
_
the immature stages constructed burrows characteristic of other groups in
this genus. These burrows opened beneath ground cover and the length and
breadth varied with the stage of development. The burrows were often simple,
straight tunnels varying from 2 to 9 cm in second and third instars to 6 to 17
cm in subadults.
The scrape-dwelling habit of adults and burrowing by the young was also
evident in populations of O. pallidipes and O. peringueyi from the Clanwilliam
district. Table 2 shows the relationship between the mode of shelter and certain
morphological characters for ten species of Opisthophthalmus.
The pedipalps of these three species are powerful and heavily sclerotized.
The chelae of the males in particular are very long and afford frontal and
lateral shielding of the opisthosoma (Fig. 1). The mesosoma is broad and
distinctly dorsoventrally compressed, and the metasoma slightly more laterally
compressed than those species which burrow in all stages. The tail is adapted
for tail-scraping operations by being heavily sclerotized.
Parabuthus planicauda (Pocock)
While collecting this species in a variety of habitats over a very large area
it was noticed that pregnant females were usually found in deep, narrow
ANNALS OF THE SOUTH AFRICAN MUSEUM
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“I9}JOYS JO spou pue sIa}ORILYO [BOISOTOYdIoUL UIe}ID Ud9MjOq AIYsUONeIOI 9Y} SuNeoIpur sniupoysydoyssid—E Jo sa1oadg
Z AIGVL
NOTES ON THE SCORPION FAUNA OF THE CAPE 253
Fig. 1. Pedipalp and prosoma of male specimens of A. Opisthophthalmus karrooensis and
B. O. macer.
depressions or properly constructed burrows, opening beneath stones. The
entrances were oval or round and the burrows varied from 4 to 11 cm in length.
Males, non-pregnant females, and young were found in scrapes of varying
shapes and lengths. Often specimens occupied natural depressions under stones,
where there was no evidence of excavation.
Laboratory specimens of pregnant females were observed scraping sand as
follows: the tail was curved either sideways or directly over the mesosoma so
that the telson lay above the carapace, and the first two pairs of walking legs
were used in alternative scraping motions to loosen the sand. The body was then
raised by stilting on the hind legs while the first two pairs of legs and sometimes
the third pair scraped the sand very vigorously so that it sprayed out behind
the scorpion; the pedipalps and fourth pair of legs were used as supports during
this operation. Transport of soil from a burrow was not observed.
The defensive armature of Parabuthus consists of the well-developed meta-
soma and highly toxic venom. It was observed that the terminal chambers of
the burrows were wide enough to enable the sting to be used effectively, and
the deep narrow depressions were constructed so as to allow the sting to be
employed without hindrance.
CONCLUSIONS
When the burrowing behaviour of O. macer was compared to O. capensis
only one difference was noted, namely the scraping of soil into a heap by the
first two pairs of legs prior to removal from the burrow in the former species.
Eastwood (1978) noted that O. capensis modified its behaviour under different
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
substratum conditions. For example, when the soil was dry the scorpion
transported it backwards out of the burrow in a single rapid movement, whereas
if it was damp or there were stones or other obstacles in the soil, it was removed
in a series of backward jerks. Thus it appears that the pattern of behaviour is
flexible to accommodate various ecological conditions. It may be that the
scraping of soil into a heap prior to transportation is used by O. capensis, but
was not observed in this species because the conditions of the soil did not require
it. However, this observation illustrates that more detailed information is
required to determine significant specific behaviour patterns. The measurements
of burrows and runs for the demes occupying two different habitats indicate
that burrow and run length are a function of the nature of the habitat, i.e. the
Kleinriviersberge specimens had deeper burrows since the substratum consisted
of deep, sandy loam, whereas in the coastal habitat of the second deme,
specimens constructed shallower burrows in pockets of compacted sand on a
limestone outcrop. Generally O. macer and O. capensis show similar trends
in burrow construction and behaviour, and more in-depth studies would be
necessary to determine significant differences.
The scrape-dwelling species of Opisthophthalmus are adapted to their
habitat by the development of a flattening of the body and the increased effective-
ness of defensive and aggressive armature. That the young of these forms are
burrowers indicates that they may have evolved from the true-burrowers. The
flattening of the opisthosoma and metasoma and elongation of the chelae are
also characteristic of the ischnurinine genus Hadogenes, a lithophilous form,
and is a good example of parallel evolution.
The burrowing habit of P. planicauda appears to occur only in females
prior to parturition. The sand-scraping behaviour of this species is essentially
the same as that reported for P. capensis by Eastwood (1977). Removal of
soil in large quantities by P. capensis was not observed in P. planicauda. However,
this process is obviously necessary for burrow excavation, since the confined
space at the entrance would not allow for the posture required for sand-scraping
operations. No doubt many other Parabuthus species will be found to burrow
and a great deal needs to be studied with respect to functional morphology,
burrowing behaviour and the relationship to the type of habitat.
ACKNOWLEDGEMENTS
I thank Dr V. Whitehead for his advice and help, and the Director of
the South African Museum for providing research and transport facilities.
REFERENCES
EAstwoop, E. B. 1977. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus
capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida,
Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214.
NOTES ON THE SCORPION FAUNA OF THE CAPE 255
Eastwoop, E. B. 1978. Notes on the scorpion fauna of the Cape. Part 3. Some observations
on the distribution and biology of scorpions on Table Mountain. Ann. S. Afr. Mus.
74: 229-248.
LANKESTER, E. R. 1882. Observations on scorpions. Proc. R. Soc. 33: 95-104.
LAWRENCE, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South
African families, genera and species. S. Afr. anim. Life 1: 152-262.
NEWLANDS, G. 1972. Ecological adaptions of Kruger Park scorpionids (Arachnida:
Scorpionides). Koedoe 15: 37-48.
PAVLOVSKY, E. 1924. Zur Ausseren Morphologie der Scorpione. Ezheg. zool. Mus. 25: 125-141.
6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).
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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:
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In describing new species, one specimen must be designated as the holotype; other speci-
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not regarded as paratypes should be listed separately. The complete data (registration number,
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Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
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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
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Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
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Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
E. B. EASTWOOD
NOTES ON THE SCORPION FAUNA OF THE CAPE
PART 4
THE BURROWING ACTIVITIES OF SOME
SCORPIONIDS AND BUTHIDS
(ARACHNIDA SCORPIONIDA)
a ASod
VOLUME 74 PART 12 FEBRUARY 1978 , | ISSN 0303-2515
MUS. COMP. ZOOL:
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Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 Band
February 1978 Februarie
Part 12 Deel
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
THE AMMONITE FAMILY LYTOCERATIDAE
NEUMAYR, 1875
By
WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER
Cape Town Kaapstad
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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
THE AMMONITE FAMILY LYTOCERATIDAE NEUMAYR, 1875
By
WILLIAM JAMES KENNEDY
Geological Collections, University Museum, Oxford
&
HERBERT CHRISTIAN KLINGER
South African Museum, Cape Town
(With 53 figures)
[MS. accepted I September 1977]
ABSTRACT
Ammonites referred to the family Lytoceratidae Neumayr, 1875, occur in rocks of
Barremian, Aptian and Albian age in northern Zululand, South Africa, and represent the
most southerly Cretaceous records of all the species present. The following are described:
Lytoceras vogdti Karakasch, which is common in the Aptian, Lytoceras aff. sauclum
(Drushchitza), represented by a single specimen from the Aptian, and Lytoceras hennigi
Zwierzycki, which is frequent in the Upper Barremian and was previously known only from
Tanzania. Eulytoceras is represented in the Upper Barremian by the widely occurring
E. phestum (Matheron); A. (Ammonoceratites) mahadeva (Stoliczka) occurs in the Middle and
Upper Albian, and A. (Ammonoceratites) crenulatum (Crick), A. (A.) ezoense (Yabe) and
A. (A.) crenocostatum (Whiteaves) are shown to be synonyms of Stoliczka’s species. A.
(Argonauticeras) depereti (Kilian), of which A. (A.) argonautarum (Anderson) and A. (A.)
belliseptatum (Anthula) are considered synonyms, occurs in the Upper Aptian; a form referred
to as A. (A.) aff. depereti is present in the Albian, accompanying A. (A.) besairiei Collignon.
Protetragonites is represented in the Upper Albian by the widely occurring P. aeolus aeolus
(d’Orbigny).
The genus Pictetia is regarded as only a doubtful lytoceratid, perhaps better classed with
the Ancyloceratina; it is represented by a specimen referred to as Pictetia aff. depressa (Pictet
& Campiche). A number of important specimens, including the types of A. (Ammonoceratites)
ezoense, A. (Am.) crenulatum and A. (Argonauticeras) belliseptatum, are figured photographi-
cally for the first time.
CONTENTS
PAGE
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INTRODUCTION
The Lytoceratidae are represented by nine species in the South African
Cretaceous, all referrable to the subfamily Lytoceratinae Neumayr, 1875, of
AST
Ann. S. Afr. Mus. 74 (12), 1978: 257-333, 53 figs.
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
which the Hemilytoceratinae Spath, 1927, and Protetragonitidae Spath, 1927,
are regarded as inseparable parts.
The Lytoceratinae are the rootstock of the Lytoceratina, ranging from
Triassic to mid-Cretaceous. They are a morphologically conservative group,
with long and widely ranging genera and species. During the late Jurassic and
Cretaceous the group were commonest in the Mesogean Realm, although known
from as far north as Greenland and as far south as the sub-Antarctic Islands,
and typically rare in the Boreal Realm of western Europe and the United
States Western Interior.
The following species are described below:
Lytoceras vogdti Karakasch, 1907
Lytoceras aff. sauclum (Drushchitza, 1956)
Lytoceras hennigi Zwierzycki, 1914
Eulytoceras phestum (Matheron, 1878)
Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865)
Ammonoceratites (Argonauticeras) depereti (Kilian, 1892)
Ammonoceratites (Argonauticeras) aft. depereti (Kilian, 1892)
Ammonoceratites (Argonauticeras) besairiei Collignon, 1949
Protetragonites aeolus aeolus d’Orbigny, 1850
Pictetia aff. depressa (Pictet & Campiche, 1861)
LOCATION OF SPECIMENS
The following abbreviations are used to indicate the repositories of the
materials studied:
BMNH British Museum (Natural History), London
IGS Geological Museum, London
MNHP Muséum d’Histoire Naturelle, Paris
SAM South African Museum, Cape Town
SAS South African Geological Survey, Pretoria
GSC Geological Survey of Canada, Ottawa
UPG University of Pretoria, Geology department
FIELD LOCALITIES
Details of field localities referred to in the paper are given in Kennedy &
Klinger (1975).
DIMENSIONS OF SPECIMENS
Dimensions of specimens are given in millimetres; abbreviations are as
follows:
D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical
diameter.
Figures in parentheses are dimensions as a percentage of total diameter.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 259
SUTURE TERMINOLOGY
The suture terminology of Wedekind (1916; see Kullman & Wiedmann
1970 for a recent review) is followed in the present work:
I; = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral
lobe, E = External lobe.
SYSTEMATIC PALAEONTOLOGY
Phylum MOLLUSCA
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order LYTOCERATIDA Hyatt, 1899
Superfamily LYTOCERATACEAE Neumayr, 1875
Family LYTOCERATIDAE Neumayr, 1875
Subfamily Lytoceratinae Neumayr, 1875
Genus Lytoceras Suess, 1865
Types species
Ammonites fimbriatus J. Sowerby, 1817 (1.C.Z.N. Opinion 130).
Diagnosis
Evolute, serpenticone, whorls slowly expanding, with a rounded to quadrate
whorl section. The surface of the shell is ornamented by fine crenulate ribs
or growth lines, and there are periodic flares, each associated with a constriction
on the internal mould. The shell surface may be finely striate. Suture formula
ELU, (Uyy : Uya) Is; highly subdivided, with a short external lobe (E) and a
triangular, highly subdivided lateral (L) lobe. There is a large septal lobe.
Discussion
Lytoceras is a conservative genus, ranging through the Jurassic and Lower
Cretaceous with very little change. There have been repeated attempts to sub-
divide the genus, but the authors follow Arkell & Wright (1957), Wiedmann
& Dieni (1968) and others in regarding Ophiceras Suess, 1865, Fimbrilytoceras
Buckman, 1918, Thysanoceras Hyatt, 1867, Thysanolytoceras Buckman, 1905,
Kallilytoceras Buckman, 1921, Crenilytoceras Buckman, 1926, Orcholytoceras
Buckman, 1926, Biasaloceras Drushchitza, 1953, Valentolytoceras Beznosov,
1958, and Dinolytoceras Beznosov, 1958, as synonyms.
Hemilytoceras Spath, 1927 (type species Ammonites immanis Oppel, 1865),
has rounded and smooth inner whorls, and on the outer whorl develops high,
concave, closely spaced lamellar flares. Metalytoceras Spath, 1927 (type species
Lytoceras triboleti Hohenegger in Uhlig 1883), is a compressed, finely ribbed
genus, with very weak constrictions, and is readily separable from Lytoceras
by virtue of the consistent splitting of the ribs into fine riblets over the venter.
Pterolytoceras Spath, 1927 (type species Ammonites exoticus Oppel, 1865), is
very loosely coiled, with slowly expanding subcircular whorls; the ornament is
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
much finer than in Lytoceras, and rather irregular.
Eulytoceras Spath, 1927 (type species Ammonites inaequalicostatus
d’Orbigny, 1840), differs from Lytoceras in having regular, distant ribs and
periodic flares, with fine striae or riblets between. In the type species, this
ornament survives in adults, as it does in E. phestum (Matheron). There are,
however, species such as ‘Biasaloceras’ sauclum Drushchitza which have
Eulytoceras-like inner whorls, but develop a Lytoceras-like adult ornament of
fine, crinkled ribs and flares with associated constrictions. These forms are both
classed with Lytoceras sensu stricto here, but they point to the homogeneity of
the Lytoceratinae and the problem of placing many ‘intermediate’ species
within the group.
The remaining Cretaceous Lytoceratinae are readily distinguished from
Lytoceras; in Pictetia Uhlig, 1883 (type species Crioceras astierianum d’Orbigny,
1842), the whorls are not in contact, whilst there are no flares or constrictions.
These are also generally absent in the rapidly expanding, finely ribbed Ammono-
ceratites (Ammonoceratites) and A. (Argonauticeras), although, as is noted
later, the dividing line between many lytoceratid genera discussed here is
sometimes exceedingly thin.
Occurrence
Lytoceras ranges from the Lower Jurassic (Pliensbachian) to the
Cenomanian, with a world-wide distribution extending as far north as Japan,
Greenland and Alaska, and as far south as Zululand and the sub-Antarctic
Islands. In the Cretaceous, the group is unknown in the Boreal region of
northern Europe and the Western Interior of North America.
Lytoceras vogdti Karakasch, 1907
Figs 1-8, 9A, 12A, F-G
Lytoceras vogdti Karakasch, 1907: 51, pl. 5 (fig. 1), pl. 24 (fig. 31), pl 26 (fig. 7). Forster, 1975:
143, pl. 1 (fig. 3).
Pictetia vogdti Drushchitza, 1956: 82, pl. 5 (fig. 19), text-fig. 36a-d. Drushchitza &
Kudryavtseva, 1960: 257, pl. 6 (fig. la—b), text-fig. 65.
‘Pictetia’ vogdti Schindewolf, 1961: 679.
Type
The larger of the two specimens figured by Karakasch (1907, pl. 5 (fig. 1))
is herein designated lectotype of this species; it is from the Barremian of the
Crimea.
Material
Large lytoceratids referred to this species are common in the Upper Aptian
of Zululand, especially along Mlambongwenya Spruit and in the Mkuze Game
Reserve; SAS L/Li, Z1113-4, Z805, 805a and A1158, are all from the Makatini
Formation, Aptian IV, at locality 151 in the Mkuze Game Reserve. SAS Z7
(Haughton Collection) is from the Makatini Formation at Haughton’s (1936)
CRETACEOUS FAUNAS FROM SOUTH AFRICA 261
Fig. 1. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38.
locality Z7. UPG B391-393 and BMNH C78716 are from the Makatini
Formation at locality 171; BMNH C78712, C78715, and C78717 are from the
same formation, Aptian IV, at locality 172, Mlambongwenya Spruit. SAS
LJE71, BMNH C78702, and possibly C78710 are from the Makatini Formation,
Aptian III-IV, at locality 168, Mfongozi Spruit; and BMNH C78703 is from
the same horizon at locality 34 on a tributary of the Mzinene River near
Hluhluwe, Zululand.
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 263
Fig. 3. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. x 0,38.
264 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. x 0,38.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 265
Fig. 5. Lytoceras vogdti Karakasch, 1907. A. SAS Z1731, showing septal lobe. xX 0,45. B. SAS
2805, showing relationship between septal lobe and cruciform internal lobe. x 1,3.
266 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. External suture of Lytoceras vogdti Karakasch, 1907. x 4,5.
267
CRETACEOUS FAUNAS FROM SOUTH AFRICA
T x “€0L8L0 HNWG
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ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Lytoceras vogdti Karakasch, 1907. Copy of the original figures of the lectotype. x 1.
Dimensions
Lectotype (after
Karakasch)
SAS L/Li
SAS Z805
SAS Z805a
UPG B393
LJE 71
SAS Z113
UPG B391
SAS Z1114
SAS A1158
Wb
34,0(40)
67,5(44)
82,0(—)
16,0(—)
95,0(45)
130,0(46)
149,0(45)
165,0(47)
198,0(49)
20)
Wh
32,0(38)
67,0(43)
TOS
16,0(—)
94,0(45)
135,0(48)
142,0(43)
161,0(46)
170,0(42)
200,0(—)
Wb: Wh
1,06
1,00
1,01
1,00
1,00
0,96
1,05
1,02
1,16
1,05
U
30,0(35)
55,0(35)
=)
= (G3)
80,0(28)
GS)
115,0(33)
120,0(30)
—O
CRETACEOUS FAUNAS FROM SOUTH AFRICA 269
Description
This is a very large lytoceratid; the largest fragment is still septate at a
whorl breadth of 211 mm, corresponding to an estimated total diameter of
close on 450 mm for the phragmocone alone. Almost, all specimens have very
poorly preserved inner whorls, commonly encrusted with oysters, and it has
proved impossible to dissect out the nuclei of the giant specimens to confirm
that they indeed correspond to the few associated juveniles, the best preserved
of which is illustrated as Figures 7A-C, 12A. This individual is partially
exfoliated and does not preserve the outer shell surface; it shows, however, very
evolute coiling with a shallow dorsal impressed zone, a wide umbilicus and
rapidly expanding depressed whorls (Fig. 7A). The exfoliated ornament (Fig.
7B-C) consists of prominent, narrow, fairly distant ribs separated by from three
to four fine growth striae. Both ribs and growth striae arise at the umbilical
seam; they are concave on the umbilical wall, sweep forwards across the shoulder
and are markedly prorsiradiate on the flank, crossing the venter with a broad
shallow convexity (Fig. 7B). The specimen bears a single broad constriction,
parallel to ribs and striae.
BMNH C78710 (Fig. 12F—G), a fragment tentatively referred to this species,
shows perfectly preserved ornament at a whorl height of 42,5 mm. The ribs
and striae are markedly prorsiradiate and pass across the venter with but a
shallow convexity. Ribs are distant, and minutely crenulated, whilst between
them are from three to five growth striae of variable strength and development.
In addition, there is a delicate spiral ornament connecting the crinkles on
successive ribs.
Beyond: 150 mm diameter, it is convenient to describe the specimens in
terms of two morphological extremes, with passage forms between. In variety «
the dorsum is flattened and the venter quite narrowly rounded. The ribs flex
slightly backwards over the umbilical wall and are radial to feebly prorsiradiate
on the flanks, crossing the venter with only slight curvature. There are sparse,
broad, shallow constrictions bounded by a prominent flared adoral rib. As
diameter increases, the ribs become widely spaced and in most specimens
growth striae are not preserved although a few specimens (e.g. Fig. 9A) show
that both they and the delicate spiral ornament survive. On moulds the ribs
appear as low, rounded undulations. In variety 8, the venter and dorsum are
equally rounded, and the greatest breadth is at mid-flank—a square cross-
section in which the corners are rounded. Ribs are distinctly prorsiradiate and
there are occasional flared ribs, not, apparently, accompanied by constrictions.
The suture line of this species is deeply and intricately subdivided (Fig. 6);
L is large and asymmetrically bifid, U, larger than Uj, and I cruciform (Fig. 5B).
The septal lobe is massive (Fig. 5A).
Discussion
The lectotype of Lytoceras vogdti is a juvenile, but shows an expansion rate
and differentiation of ornament into ribs and striae like that of some of the
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
present similarly sized fragments, sufficient to suggest them to be conspecific.
Some of the specimens resemble Lytoceras mikadyense Krenkel (1910: 233,
pl. 22 (fig. 5)), but this species has consistently wider spaced ribs with striae
between, indicating it to be closer to Eulytoceras. Of other species described by
Krenkel, the unique holotype of ‘Crioceras’ schlosseri Krenkel (1910: 227,
pl. 22(3) (fig. 15)), although more compressed, has ornament like that of some
of the present specimens.
Of species described from approximately contemporaneous strata in
Madagascar, Lytoceras aff. vicinum Douvillé of Collignon (19625: 5, pl. 217
(fig. 952)) has a compressed whorl section with delicate concave, prorsiradiate
flank ribs, whilst Lytoceras belliseptatiforme Collignon (19626: 6, pl. 217
(fig. 953), pl. 218 (fig. 953)) is densely and evenly ornamented by fine ribs, has
a lower expansion rate and circular whorl section. Lytoceras mahafalense
Collignon (1963: 14, pl. 246 (fig. 1503)) has a distinctly compressed oval whorl
section and is of Albian age. The most similar species appears to be Lytoceras
(Thysanolytoceras) strambergense mut. albensis Collignon (1963: 9, pl. 244
(fig. 1047)), a Lower Albian form with rather finer ribbing and a distinctively
compressed whorl section and lower expansion rate. The species may be a
descendant of the present form.
Occurrence
Lytoceras vogdti was first described from the Barremian of the Crimea;
it has been subsequently recorded from the Upper Aptian of Mozambique,
and now Zululand.
Lytoceras aff. sauclum (Drushchitza, 1956)
Figs 9B, 1OA—D, 11B
Compare:
Biasaloceras sauclum Drushchitza, 1956: 71, pl. 4 (fig. 13); Drushchitza &
Kudryavtseva 1960: 256, pl. 5 (fig. 1).
Lytoceras sauclum Wiedmann and Dieni, 1968: 29.
Type
The holotype is Drushchitza’s (1956, pl. 4 (fig. 13)) original specimen from
the Hauterivian of the Crimea.
Material
BMNH C78704 from the Makatini Formation, Albian IV, locality 171,
Mlambongwenya Spruit, Zululand.
Dimensions
D Wb Wh Wb: Wh U
BMNH C78704. ._—‘122,0 52,5(43) 48,0(39) 1,1 49,0(40)
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CRETACEOUS FAUNAS FROM SOUTH AFRICA
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Die ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
The coiling is very evolute, serpenticone, with a narrow, shallow, impressed
dorsal area. The whorl section is depressed (whorl breadth to height ratio may
be up to 1,1), expanding quite rapidly, with the greatest breadth some way
below mid-flank. The umbilical seam is deeply incised, the umbilicus of moderate
width (about 40% of diameter) and depth. The umbilical wall is high and
rounded, the umbilical shoulder broadly rounded, the flanks flattened, ventro-
lateral shoulders broadly rounded, and the venter somewhat flattened.
The shell is ornamented by growth striae, and widely spaced crinkled ribs.
Ribs and striae arise at the umbilical seam, curve backwards across the umbilical
wall and forwards across the umbilical shoulder and are straight and slightly
prorsiradiate across the flanks. The inner whorls have a distinctive Eulytoceras-
like appearance, with between seven and ten striae between each pair of ribs,
the ribs themselves being distinctly—if minutely—crenulate, with a gently
inclined apical slope and an abrupt apertural face which give the surface of the
shell a scale-like appearance (Fig. 9B). As size increases (Fig. 10) the ribs become
more closely spaced, striae lose their prominence, crenulations become more
distinct, and on some specimens, fine strigations appear on some areas of shell.
All specimens bear occasional broad, deep constrictions on both shell and
mould, preceded by a high, flared rib. Both flares and constrictions become
increasingly prominent as diameter increases.
The suture line (Fig. 11B) includes a deeply incised elongate E with a
long, narrow lanceolate median saddle, a large, triangular, asymmetrically
bifid L, and a similar smaller U, lobe, separated by a large, splayed bifid L/U,
saddle.
Discussion
Drushchitza figured the type specimen of Biasaloceras sauclum in side view
only, but the illustration shows clearly the differentiation of ornament on the
Eulytoceras-like early whorls into ribs with groups of striae between, an inter-
mediate stage with rather irregular ornament, and middle to late growth
stages with straight, variably spaced crenulate ribs and occasional flares. The
whorl section is slightly depressed, with the greatest breadth some way below
mid-flank. Although of Hauterivian age, the striking similarity between the
illustration and the present material suggests that they belong to the same
group, in spite of age difference.
Biasaloceras striatum Drushchitza (1956: 78, pl. 4 (figs 14a—b); Drushchitza
& Kudryavtseva 1960, pl. 4 (figs 5a—b)) has a similar juvenile ornament, but a
rather different whorl section; based on a juvenile, the authors are inclined to
regard it as a possible variant of L. sauclum. The markedly different early and
late ornament of Lytoceras sauclum helps to differentiate it from most Cretaceous
Lytoceras species. Thus, of the long-ranging Stramberg species, Lytoceras
liebigi Oppel (Zittel, 1868: 74, pl. 9 (figs Sa—b, 7a—b), pl. 10) has a depressed
oval whorl with concave crenulate ribs. Lytoceras strambergense Zittel (1868:
ATE:
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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274 ANNALS OF THE SOUTH AFRICAN MUSEUM
74, pl. 11 (figs 1-3)) has flexuous crenulate ribs, and Lytoceras sutile Oppel
(1865: 551; Zittel 1868: 76, pl. 12 (figs 1-5)) has a circular whorl section and
flexuous ribbing throughout ontogeny. Lytoceras aff. strambergense of Collignon
(1949a: 71, pl. 12 (fig. 1), text-fig. 5) from the Hauterivian of Madagascar has
distinctly concave ribs and flares. Lytoceras mahafalense Collignon (1963: 14,
pl. 246 (fig. 1053)) from the Albian of Madagascar has a distinctly compressed
whorl section and flattened flanks, whilst Lytoceras vicinum Douvillé (1916:
93, pl. 11 (fig. 6)) is also a compressed species with an elliptical to oval whorl
section. A final species to mention, Lytoceras hennigi Zwierzycki, described
below, has a depressed oval whorl section, and a very even, regular ornament.
Occurrence
Lytoceras sauclum was originally described from the Hauterivian of the
Crimea and is also recorded from the Valanginian of Sardinia.
Lytoceras hennigi Zwierzycki, 1914
Figs 11A, 12B—E, 13A-E
Lytoceras sp. Krenkel, 1910: 224, pl. 22 (fig. 10).
Lytoceras hennigi Zwierzycki, 1914: 40, pl. 4 (figs 6-7). Spath, 1939: 7.
Type
Lectotype herein designated is Zwierzycki’s (1914, pl. 4 (figs 6-7)) original
figured specimen from the Trigonia schwartzi Bed (Barremian—Aptian) of
Mikadi, Tanzania.
Material
Fragments of five individuals and one fairly complete specimen, BMNH
C78705-9, C78711, from the Makatini Formation, Barremian 1, locality 171,
Mlambongwenya Spruit, northern Zululand.
Dimensions
D Wb Wh Wb:Wh U
Lectotype (from
IE MATOS) 5 5 ‘ISS 58,0(37) 57,0(37) 1,02 61,0(39)
BMNH C78708 — 37,0(—) 31,0(—) 1,19 —
BMNH C78705___.. — 35,0(_ ) SUSE) 1,10 --
Description
The coiling is very. evolute, with a narrow, shallow impressed area, the
whorls expanding quite rapidly. The whorl section is a depressed oval, the
greatest breadth being at mid-flank. Ornament consists of fine ribs, growth
striae, and occasional flares. These arise at the umbilical seam, sweep sharply
backwards over the inner umbilical wall where they are concave, then forwards
CRETACEOUS FAUNAS FROM SOUTH AFRICA 275
Fig. 11. A. Partial suture of Lytoceras hennigi Zwierzycki, 1914. BMNH C78705. x 5.
B. Partial suture of Lytoceras aff. sauclum (Drushchitza, 1956). BMNH C78704. x 4,5.
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
to pass straight across the umbilical shoulder and flanks where they are markedly
prorsiradiate, to cross the venter with a faint forwards projection. The best
preserved specimens show the ribs to have been even, sharp-topped and
minutely crinkled, and separated by wider interspaces ornamented by irregular
fine striae. Delicate spiral ridges connecting crinkles are present over the
whole of the shell surface when well preserved. There are periodic high, sharp
flares (Fig. 12D-E), although these are only rarely preserved. When shells
are partially exfoliated, growth striae, crinkles and strigations disappear,
leaving only evenly spaced, rounded ribs and occasional constrictions (corres-
ponding to the site of flares). Moulds are smooth save for constrictions
(Fig. 12D).
The sutures are poorly exposed, but include a large E/L and smaller L/U,,
both deeply incised, asymmetric and triangular, separated by a deep splayed
L (Fig. 11A). A distinct siphonal band is present in one specimen (Fig. 12D).
Discussion
The present specimens are much smaller than Zwierzycki’s lectotype, but a
comparison with topotype specimens in the British Museum (Natural History)
suggests that they are indeed referrable to Lytoceras hennigi. The whorl section
of topotype specimens is depressed at small diameters, whilst the ornament
consists of even, crinkled ribs with smooth, somewhat wider interspaces between,
and occasional strong periodic flares, marked by broad constrictions on the
internal mould.
The regular, fine ribs and periodic flares of L. hennigi readily distinguish
it from Eulytoceras species discussed below, whilst Lytoceras liebigi, L. stramber-
gense and L. sutile differ in proportions and ornament, as noted above. Lytoceras
subsequens Karakash (1907: 49, pl. 5 (fig. 9, 9a), pl. 24 (fig. 32)) lacks prominent
constrictions and flares; L. aff. sauclum has completely different, Eulytoceras-
like juvenile ornament as discussed above, whilst Lytoceras puezanus Haug
(1889: 197, pl. 8 (fig. 1) pl. 10 (fig. 1)) has less marked flares and very strong
strigations. Lytoceras densefimbriatum Uhlig (1883: 191, pl. 6 (figs la—c, 2)) is
a very finely ornamented, compressed form with weaker flares and stronger
strigations, Lytoceras mahafalense Collignon is compressed, whilst Lytoceras
subfimbriatum d’Orbigny (1840: 121, pl. 35) has a compressed whorl section,
very fine fimbriate ornament, and distant flares, four to five per whorl.
Occurrence
Lytoceras hennigi is known from the Lower Cretaceous of Tanzania and
the Upper Barremian of Zululand only.
Genus Eulytoceras Spath, 1927
Type species
Ammonites inaequalicostatus d’Orbigny 1840.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 277
MMM, yy
to)
y Ye TY,
Yih:
G
Fig. 12, A, F, G. Lytoceras vogdti Karakasch, 1907. A. BMNH C78703. F-G. BMNH C78710.
x 1. B-E, Lytoceras hennigi Zwierzycki, 1914. B—D. BMNH C78708. E. BMNH C78705.
ae
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 13. Lytoceras hennigi Zwierzycki, 1914. A-B. BMNH C78705. C-E. BMNH C78706.
Phe Os
CRETACEOUS FAUNAS FROM SOUTH AFRICA 279
Diagnosis
Compressed to round-whorled serpenticone lytoceratids with slowly
expanding whorls ornamented by fine, regular, distant rectiradiate to prorsi-
radiate ribs and periodic flares, all of which extend across the venter without
interruption.
Discussion
The regular, fine, distant ribs and occasional simple flares of typical
Eulytoceras clearly separate them from Lytoceras itself (with typically crinkled
ribs and flares and periodic constrictions), Hemilytoceras (with smooth inner
whorls and high, closely spaced concave flares), Pterolytoceras (with minutely
crinkled growth lines and irregular fine ribs), Ammonoceratites (Ammonoceratites)
and A. (Argonauticeras), both of which have fine, dense, subdued ornament.
As the authors have noted, however, some species such as Lytoceras
sauclum, described above, have Eulytoceras-like inner whorls, whilst ‘Lytoceras’
mikadyense Krenkel has Eulytoceras inner whorls and develops very closely
spaced ornament when adult, being transitional to Lytoceras sensu stricto.
Occurrence
Eulytoceras ranges from the Hauterivian to Lower Aptian, and species
are known from southern Europe, the Crimea, Caucasus, Bulgaria, east Africa
(Tanzania), South Africa (Zululand), Madagascar, and California.
Eulytoceras phestum (Matheron, 1878)
Figs 14A-E, 15
Ammonites phestus Matheron, 1878, pl. c—20 (fig. 5).
Lytoceras phestus Uhlig, 1883: 187, pl. 5 (figs 1-4, 20). Trautschold, 1886: 137. Haug, 1889:
196, pl. 8 (fig. 2). Simionescu, 1898: 59, pl. 2 (figs 4, 10), pl. 20 (fig. 17). Sarasin &
Schondelmayer, 1901: 19. Karakasch, 1907: 46, pl. 4 (fig. 10), pl. 20 (figs 5—6). Pervinquieére,
1907: 64. Kilian & Reboul, 1915: 21, pl. 1 (figs 1-2). Petkovic, 1921: 48. Kulzhinskaya-
Voronets, 1933: 5, fig. 6. Rouchadzé, 1933: 174. Ksiazkiewicz, 1938: 230, pl. 1 ee 1).
Eristavi, 1955: 53; 1957: 59.
Lytoceras gresslyi Somogyi (non Hantken), 1914, pl. 11 (fig. 12).
? Lytoceras lorentheyi Somogyi, 1914: 300, pl. 11 (figs 10-11).
? Protetragonites phestus Nagy, 1967: 62, pl. 2 (fig. 1).
Eulytoceras phestum Spath, 1927: 64. Drushchitza, 1956: 87, pl. 5 (fig. 18); Drushchitza &
Kudryavtseva, 1960: 258, pl. 6 (fig. 3). Collignon, 1962a: 95, pl. 214 (fig. 935). Dimitrova,
1967: 27, pl. 9 (fig. 3). Kotetichvili, 1970: 60, pl. 4 (fig. 3). Vasicek, 1972: 37, pl. 2 (fig. 6).
Murphy, 1975: 18, pl. 2 (fig. 3).
Type
The ‘holotype’ is Matheron’s original specimen, in the collections of the
Muséum d’Histoire Naturelle de Marseille according to Murphy (1975: 18),
but Matheron in fact gives three figures, and it is not clear if these represent
individual specimens or composite drawings. Lectotype designation (if necessary)
must await fuller redescription of the type material.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 14. Eulytoceras phestum (Matheron, 1878). A-C. BMNH C78698. D-E SAS L7(5).
; A=Di xX ke Bx xe
CRETACEOUS FAUNAS FROM SOUTH AFRICA 281
Fig. 15. Partial suture of Eulytoceras phestum (Matheron, 1878) based on SAS L7(5). x 8.
Material
Two specimens, SAS. L7(5) and BMNH C78698, both from the Makatini
Formation at locality 170, Mlambongwenya Spruit, Zululand (Barremian 1).
Description
The material consists of a wholly septate juvenile 27,5 mm in diameter,
and a body chamber fragment of a larger individual.
The coiling is very evolute, the whorls only just in contact, with, as a
consequence, a very narrow impressed area. The coiling is evolute, the umbilicus
being approximately 30 per cent of the diameter, and shallow with a rounded
wall. The whorl section is as broad as high, rounded trapezoidal, with a rounded
ventrolateral shoulder, convergent, somewhat flattened sides, the greatest
breadth being some way beyond mid-flank, and a broadly rounded venter.
Inner whorls up to a diameter of 15 mm appear to be devoid of all ornament.
Beyond this ornament consists of fine, narrow ribs separated by wide, smooth
interspaces. The ribs arise at the umbilical seam, pass forwards across the
umbilical wall and shoulder and are prorsiradiate and weakly flexed or convex
on the flanks. Interspaces are ornamented by fine growth striae.
The external suture is partially exposed, as shown in Figure 15.
Discussion
The present specimens match closely with Matheron’s original figures in
terms of general ornament and proportions, his illustrations being based upon
crushed specimens. Eulytoceras phestum is readily distinguished from Eulytoceras
inaequalicostatus (d’Orbigny) (1840: 18, pl. 29 (figs 3-4)), for that species
develops strong flares, nine per whorl in D’Orbigny’s figure. Eulytoceras
raricinctum (Uhlig) (1883: 88, pl. 5 (figs 5—7)) is a distantly ribbed form with only
fifteen ribs per whorl rather than the forty to forty-two typical of E. phestum.
Eulytoceras rotundatum Drushchitza (1956: 86, pl. 5 (fig. 15)) has a depressed
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
whorl section with fewer, more convex ribs. Eulytoceras intemperans (Coquand
in Matheron) (1878, pl. c—20 (fig. 4-4c)) has only twenty-one ribs per whorl,
with distinctive striae between, whilst Eulytoceras electra (Coquand in Matheron)
(1878, pl. c-20 (fig 2a—2b)) has fifteen flexuous ribs per whorl, also with striae
between.
The east African “Lytoceras’ mikadyense (Krenkel) (1910: 223, pl. 22 (fig. 5))
is a depressed species, has periodic flares, rather irregular ribs, and prominent
striae between. Eulytoceras belchasifakaense Collignon (1949a: 70, pl. 11
(fig. 2-2a), text-fig. 4), from the Hauterivian of Madagascar, has straight,
quite distantly spaced ribs, rather than the slightly flexuous ornament of
E. phestum, but may subsequently prove to be a synonym. E.? komihevitraense
Collignon (1963: 15, pl. 247 (fig. 1055)), from the Middle Albian of Madagascar,
is said to possess constrictions, and has dense, distinctly concave ribs with a
ventral sinus. Finally, Eulytoceras lepidum (d’Orbigny) (1840: 149, pl. 48
(figs 3—4)) bears strong flares, and is more distantly ribbed.
Occurrence
Eulytoceras phestum is a typically Barremian species, known from southern
France, the Tyrol, Czechoslovakia, Hungary, Bulgaria, the Crimea, Georgia
and the Carpathians, Tunisia, Madagascar, and South Africa. Kilian & Reboul
(1915: 21) also cite the species from the Lower Aptian of southern France,
whilst Murphy (1975: 18) records it from the Upper Barremian of California.
Genus Ammonoceratites Bowdich, 1822
Subgenus Ammonoceratites Bowdich, 1822
Type species
Ammonoceratites lamarcki Bowdich, 1822.
Diagnosis
Large, evolute, serpenticone lytoceratids with a circular whorl section,
ornamented by dense, crenulate fine ribs and growth striae, sometimes combined
with faint spiral striae. Occasional broad, shallow constrictions may appear.
Suture highly subdivided, with a massive cruciform septal lobe (Is).
Discussion
A general lack of flares and associated constrictions, fine ornament and
rounded whorls separates Ammonoceratites (Ammonoceratites) from Lytoceras
Suess, 1865 sensu stricto, whilst lack of regular, distant flares differentiates it
from Eulytoceras Spath, 1927, Hemilytoceras Spath, 1927, and Pterolytoceras
Spath, 1927. Ammonoceratites (Argonauticeras) is readily separated by virtue
of its rapidly expanding whorls and an ornament of very fine, even, generally
non-crenulate ribs.
Considerable confusion surrounds the introduction of this genus, as much
CRETACEOUS FAUNAS FROM SOUTH AFRICA 283
as that surrounding the type species, as discussed below. Following Casey
(1960: 2), Ammonoceratites Rafinesque, 1815, is regarded as a nomen nudum
and Ammonoceras Lamarck, 1822, a nomen dubium (as introduced) over which
Ammonoceratites Bowdich, 1822 (both of which have type species based on the
same specimen), has priority.
The following species have been referred to the restricted subgenus:
A. (Ammonoceratites) lamarcki Bowdich (1822: 21, pl. 3 (fig. 14)).
A. (Ammonoceratites) glossoidea (Lamarck) (1822: 144 in Chenu 1859: 90,
text-figs 391-392). |
A. (Ammonoceratites) mahadeva (Stoliczka) (1865: 16, pl. 80 (fig. 1-1b)),
Middle Albian of southern India.
A. (Ammonoceratites) crenulatum (Crick) (1907: 236), Upper Albian of
Zululand.
A. (Ammonoceratites) betiokyense Collignon (19625: 8, pl. 218 (figs 954-955)),
Upper Aptian of Madagascar.
A. (Ammonoceratites) crenocostatum (Whiteaves) (1884: 45, pl. 9 (fig. 2)),
Albian of British Columbia.
. (Ammonoceratites) ezoense (Yabe) (1903: 9, pl. | (fig. 1), pl. 5 (fig. 1)), Upper
Albian of Japan.
aN
Occurrence
Ammonoceratites ranges from Upper Aptian to Upper Albian. There are
records from British Columbia, southern France, southern India, Japan,
Madagascar, and South Africa.
Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865)
Figs 16-33
? Ammonoceratites lamarcki Bowdich, 1822: 21, pl. 3 (fig. 14).
? Ammonoceras glossoidea Lamarck, 1822: 644.
? Ammonoceratites glossoidea Chenu, 1859: 90, text-figs 391-392.
Ammonites mahadeva Stoliczka, 1865: 165, pl. 80 (fig. 1).
Ammonites crenocostatus Whiteaves, 1876: 45, pl. 9 (fig. 2).
Lytoceras batesi Whiteaves (non Trask), 1884: 202, pl. 27 (fig. 1); 1900: 270.
Lytoceras mahadeva Kossmat, 1895: 17.
? Lytoceras ezoense Yabe, 1903: 9, pl. 1 (fig. 1), pl. 5 (fig. 1).
? Lytoceras crenulatum Crick, 1907: 236.
? Lytoceras sp. aff. mahadeva: Jeannet, 1908: 105-119, pls 3-6.
Lytoceras (Ammonoceratites) crenulatum: Breistroffer, 1936: 169, text-fig. 10h.
Ammonoceratites cf. mahadeva Collignon, 19495: 45, text-fig. 6.
Ammonoceratites mahadeva: Collignon, 1963: 12, pl. 245 (fig. 1051).
Ammonoceratites crenocostatus McLearn, 1972: 22, pl. 1 (fig. 5), pl. 2 (figs 1-3).
Type
The holotype is Stoliczka’s (1865, pl. 80 (fig. 1)) original specimen from
the Lower Utatur Group of the environs of Moraviatoor, southern India, and
presumably of Albian age. The original illustrations are reproduced here as
Figures 27-28.
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
Eight specimens: BMNH C18265, the holotype of Lytoceras crenulatum
Crick, from the Albian of the Mzinene River area; BMNH C78890, from Bed
2 of the Mzinene Formation at locality 35 on the Mzinene River (Albian III);
BMNH C78700-1, from the Mzinene Formation at locality 54 on the Mzinene
River (Albian V), SAS A1195 and 2004 come from the same horizon at locality
53. SAS Z418 and Z428 in the Van Hoepen Collection are from Impala, Ndumu,
Mzinene Formation (Albian II or III).
Dimensions
D Wb Wh Wb:Wh U
Holotype (after
Stoliczka . 218,0 82,0(38) 78,0(36) 1,05 95,0(44)
A. ezoense (after
Yabe) . 200,0 75,0(37) 80,0(40) 0,94 70,0(35)
A. crenocostatum
holotype, GSC
4987 (after
McLearn) 44,0 14,7(33,5) 15,4(35,0) 0,95 20,7(47)
A. crenocostatum,
GSC 4976 = 47,0(—) 40,0(—) 1,08 — (—)
BMNH C78700 ae 44,8(—) 41,5(—) 1,08 — (—)
— 29,0(—) 28,0(—) 1,04 — (—)
SAS A1195, at . 115,0 45,0(39) 44,0(38) 1,03 48,0(42)
SAS Z428, at 137,0 49,0(36) 48,0(36) 1,02 58,0(42)
ate" 194,0 75,09) 74,0(38) 1,03 83,5(43)
BMNH C78890 220,0 93,0(42) 88,0(40) 1,05 85,0(38)
SAS Z418 — 78,0(—) 76,0(—) 1,03 — (—)
SAS A2008 . . . 253,0 98,0(39) 96,0(38) 1,01 101,0(40)
Description
Very large (the Zululand specimens are still septate at diameters of over
300 mm), very evolute, the whorls only just in contact with a narrow, shallow
dorsal impressed zone. The whorl section is almost circular, typically just
slightly broader than high, the ratio varying from 1,0 to 1,05. The whorls expand
quite rapidly and the umbilicus is broad, varying from 38 to 42 per cent of the
diameter and increasing slightly during ontogeny, with a deeply incised umbilical
seam.
Ornament changes markedly throughout growth. At the smallest diameters
visible (Figs 16A, 31C) the shell surface appears virtually smooth, bearing
only fine prorsiradiate growth striae, distant, minutely crenulated ribs, and
occasional flares which correspond to distinct constrictions on internal moulds.
Ribs, flares and striae arise at the umbilical seam, are concave on the inner
part of the umbilical wall, then sweep forwards and are prorsiradiate and feebly
285
CRETACEOUS FAUNAS FROM SOUTH AFRICA
~ we
oe .
ee SRE PR.
B
we
x
a
RAT
“aS
SS
©
ASSES
ERS ens
. SSS
Fig. 16. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). A. SAS A2004.
B. BMNH C78890. x 1.
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 17. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45.
convex on inner and mid-flank (Figs 17, 20). Beyond this, the crenulated ribs
become increasingly frequent, and there are sparse prominent flares and
associated constrictions. In middle and later growth ribs are strong and closely
spaced (Fig. 20). During these later growth stages, ribs and striae arise at
the umbilical seam, sweep strongly backwards across the inner part of the
umbilical wall and forwards over the umbilical shoulder. They are distinctly
prorsiradiate and variably, although commonly, weakly concave across the
CRETACEOUS FAUNAS FROM SOUTH AFRICA 287
We Has
highs y
ws
oe (a
Fig. 18. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45.
flanks and sweep forwards across the ventrolateral shoulder to produce a broad
ventral peak of variable depth. Growth striae are scarcely visible to the naked
eye, but the ribs are very distinct and strongly crenulate with a vertical apertural
face and a gently inclined apical slope. On the ventral and ventrolateral region
of some of the Zululand specimens there are distinct, curved longitudinal striae
between and connecting the crinkled ribs, arising from individual crinkles
(Fig. 20). .
Partially exfoliated shells show low, broad, irregular, fold-like constricted
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 19. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS 2418. x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 289
Fig. 20. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78890.
x 0,6.
areas during middle and later growth (Figs 22—23), and in some places, transverse
rows of large pits corresponding to the site of crinkled ribs. Weathered shell
surfaces are also covered in minute pits. Internal moulds are essentially smooth,
save for broad, shallow, fold-like constrictions (Fig. 19).
One specimen shows a dorsal callus, through which traces of-the under-
lying ventral ornament are still visible.
ANNALS OF THE SOUTH AFRICAN MUSEUM
290
BMNH C78890.
text-fig. 391.
°
9
1865)
ka,
icz
Copy of Chenu 1859
h.
i
Ammonoceratites (Ammonoceratites) mahadeva (Stol
A.
°
Dal
x 0,6. B—D. Ammonoceratites lamarcki Bowd
1g
F
CRETACEOUS FAUNAS FROM SOUTH AFRICA 291
*
< : - ,
4 - i, i ‘
dic Ae oe Gee ‘
Gy 7. ae
Fig. 22. The holotype of Ammonoceratites (Ammonoceratites) crenulatum (Crick, 1907).
BMNH C18265. Slightly reduced.
The suture line is highly subdivided with a long spatulate ventral saddle,
a large bifid asymmetric E/L saddle and a smaller also asymmetrically bifid
L/U, saddle, separated by a very large lateral lobe (L), deeper than the external
lobe (E) and bifid, with a large subdivided median element. U, is highly sub-
divided, dorsal internal lobe (1) cruciform, septal lobe large (Fig. 18).
Discussion
The Zululand specimens show some variation in strength and direction of
ornament and relative proportions when compared with Stoliczka’s magnificent
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 23. The holotype of Ammonoceratites (Ammonoceratites) crenulatum (Crick, 1907).
BMNH C18265S. Slightly reduced.
specimen. In particular, the ventral peak is more prominent than shown in
his figure and the ribs are prorsiradiate rather than radial. The material varies
in these respects, and rather than trying to separate it into Ammonoceratites
mahadeva with straight ribs and a shallow ventral peak and Ammonoceratites
crenulatum, with prorsiradiate ribs and a deeper peak, the authors regard their
material as one variable species. It is, however, far from clear whether
Ammonoceratites mahadeva is the earliest name used for this species.
Lamarck (1822, vol. 7: 644) introduced the name Ammonoceras glossoidea,
CRETACEOUS FAUNAS FROM SOUTH AFRICA 293
A B C
Fig. 24. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78700. x 1.
without figuring the material. His description is brief: ‘Cette coquille rompue
en trois morceaux, qui s’appartiennent successivement, et dont l’un offre
Pextrémité supérieure de cette méme coquille, est d’une assez grande taille,
fort épaisse en sa partie inférieure, arcuée presque en demi-cercle, et se termine
supérieurement en forme de langue. Ses loges sont remplies de matiére pierreuse,
et leurs cloisons ne se distinguent que dans les parois ot! leurs concours forment
des sutures lobées, laciniées, rameuses, tout a fait analogues a celles des ammo-
nites. Mais la coquille dont il s’agit en est trés-distincte par sa forme générale;
car malgré son arcuation, elle n’efit point forme de tours contigus, si la nature
Petit agrandie davantage. Sa longueur est de cinquante centimetres.’
In the same year, Bowdich (1822: 21, pl. 3 (fig. 14)) introduced the name
Ammonoceratites lamarcki, based upon the same fragments. His description is
ANNALS OF THE SOUTH AFRICAN MUSEUM
294
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CRETACEOUS FAUNAS FROM SOUTH AFRICA 295
: | a
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Fig. 26. External suture of Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865).
BMNH C78890. x 1,6.
also brief: -“Septa numerous, undulated at the margins, pl. 3, fig. 14. Siphon
marginal, interior. Rolled on itself in the same plane? The locality is unknown,
M. Lamarck purchased it by accident: he kindly allowed me to take it home
in order that the figure, the first that has been made, might be as accurate as
possible.’
Subsequent authors have varied in their views of the nature of Ammonoceras
glossoidea— Ammonoceratites lamarcki (the latter has priority; fide Casey 1960),
regarding it as of both Jurassic and Cretaceous age and as a species of Hamites,
Toxoceras, and Pictetia. The age is, however, in all probability Cretaceous,
for as Chenu (1859: 90) notes, the specimen was not purchased by Lamarck,
for he (Chenu) quotes Valenciennes as telling him that ‘Lamarck lui a souvent
répété que ce fossile, dont il faisait grand cas, avait été rapporté par le secrétaire
de la colonie de Pondichéry, et donné au fils de Buffon, qui lui en avait luiméme
fait cadeau. M. Valenciennes ajoute que parmi les ammonites rapportées de
l’Inde par Jacquemont, il se trouve aussi un fragment de I’A. fimbriatus.’
Chenu’s figure is much better than that of Bowdich, and is reproduced
here as Figure 21B—D. The specimen was said to be 500 mm in diameter, and
corresponds to the generally accepted interpretation of Ammonoceratites; its
occurrence in southern India indicates that it may well be a specimen of what
has subsequently been termed Ammonoceratites mahadeva. The figures are,
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 27. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka’s
figures of the holotype, reduced here by approximately 0,75.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 297
Fig. 28. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka’s
figure of the holotype, reduced here by approximately 0,75.
however, so poor that, until the specimen is redescribed and refigured, the
authors would prefer to use Stoliczka’s name, having sought without success
for the Lamarck/Bowdich type in the Paris museums.
Of other species, Ammonoceratites (Am.) betiokyense, from the Aptian of
Madagascar, is based on two small fragments only, but appears to have
distinctly flattened sides and a markedly asymmetric or even subtrifid U, lobe
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 29. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Lectotype. x 0,75.
according to Collignon (19625: 8).
Ammonoceratites (Am.) ezoense (Yabe), lectotype, herein designated,
Yabe’s (1903: 9, pl. 1 (fig. 1), pl. 5 (fig. 1)) original specimen is also an Albian
species (see range charts in Matsumoto 1954), the original material being
preserved in the collections of the Geology Department of Tokyo University,
and refigured here as Figures 29-30, 31A—B. Yabe separated it from Ammo-
CRETACEOUS FAUNAS FROM SOUTH AFRICA 299
Fig. 30. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Lectotype. x 0,75.
noceratites (Am.) mahadeva on the basis of more rapidly expanding whorls and
a smaller umbilicus (Yabe’s measurements are reproduced above), but as can
be seen from the present photographs, the preservation is poor and the specimen
probably within an acceptable range of variation for A. (Am.) mahadeva as
shown by the Zululand specimens (e.g. compare Figs 29-30, 31A—B with Figs
17-19); the authors therefore place it as a questionable synonym.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
» iy
oe se
if
RY
re
,
Fig. 31 A-B. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Part of lectotype,
showing suture. C. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1861); inner
whorls of SAS A2004. x 1,5.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 301
Ammonoceratites (Am.) crenocostatus (Whiteaves) is a further Middle? to
Upper Albian species known only from the Queen Charlotte Islands off the
coast of British Columbia. The authors have been able to examine casts of the
holotype and other specimens from the area, illustrated here as Figures 32-33.
The holotype is only 43 mm in diameter, and it retains only part of the
shell. The inner whorls are strongly constricted on the mould up to a diameter
of 20 mm, with traces of delicate striae where shell is preserved; from 26 to
40 mm there are distant fine ribs with delicate growth striae between, both
ribs and striae being prorsiradiate and feebly convex on the venter. Over the
last preserved part of the specimen the ribs crowd markedly. GSC 4976 is a
larger specimen, with an original diameter of approximately 120-130 mm. The
outer whorl shows distinctly concave prorsiradiate flank ribs, with a marked
convex ventral peak, rib density, form and strength matching closely to the
similarly sized Zululand specimen BMNH C78700 (Figs 24-25). So far as can
be judged there are no criteria that can be used to separate A.(Am.) crenocostatum
from the Zululand material, and it is therefore suggested that it is a further
synonym of Ammonoceratites (Am.) mahadeva.
Occurrence
As defined above, Ammonoceratites (Am.) mahadeva ranges from Middle
to Upper Albian, and is known from southern India, the Queen Charlotte
Islands, Madagascar, Japan, Zululand, and perhaps the Jura (Jeannet 1908:
105-119, pls 3-6).
Subgenus Argonauticeras Anderson, 1938
Type species
Lytoceras argonautarum Anderson, 1902.
Diagnosis
A subgenus of Ammonoceratites with a high expansion rate, subrectangular
to trapezoidal whorl section, fine, equal, dense, straight to gently flexed ribs
or striae without obvious crinkles. Weak, broad constrictions may be present
on early whorls.
Discussion
Argonauticeras can be separated from other Cretaceous lytoceratid genera
by its fine, even, typically non-crenulate ornament and high expansion rate.
The trapezoidal whorl section, emphasized by Wright (in Arkell & Wright
1957), is a less distinctive feature, as discussed below. Differences from
Ammonoceratites (Ammonoceratites) are outlined above.
Four species have been referred to the subgenus:
Argonauticeras argonautarum (Anderson) (1902: 85, pl. 7 (figs 154-155);
1938: 149, pl. 17 (fig. 3), pl. 19 (figs 1-2)). Upper Aptian of California.
ANNALS OF THE SOUTH AFRICAN MUSEUM
302
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CRETACEOUS FAUNAS FROM SOUTH AFRICA
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304 ANNALS OF THE SOUTH AFRICAN MUSEUM
Argonauticeras depereti (Kilian) (1892: 8, pl. 1 (fig. 2). Thomel, 1968: 684-687,
pl. 35 (figs 1-4)). Upper Aptian of southern France.
Argonauticeras besairiei Collignon (19495: 46, pl. 8 (fig. 1)). Upper Aptian to
Middle Albian of Madagascar.
Argonauticeras belliseptatum (Anthula) (1899: 97, pl. 6 (fig. 1), pl. 7 (fig. 1)).
Aptian of the Caucasus and Madagascar.
Of these species, Dimitrova (1967: 28, pl. 10 (fig. 6)) figured a fragment
referred to as Pictetia belliseptata (Anthula) which appears actually to uncoil.
The authors therefore take the opportunity to figure photographically the
holotype of A. (Ar.) belliseptatum (Figs 40-43) which shows it to be an
Argonauticeras.
Pseudotetragonites Drushchitza, 1956 (type species P. kudrjavzevi
Drushchitza), is a subjective synonym of Argonauticeras, based on juveniles.
Occurrence
Argonauticeras is known from the Upper Aptian of southern France, the
Caucasus, California, the Lower and Middle Albian of Madagascar, and the
Middle Albian of Zululand.
Ammonoceratites (Argonauticeras) depereti (Kilian, 1892)
Figs 34-37, 38B, 39-43
Lytoceras depereti Kilian, 1892: 8, pl. 1 (fig. 2a—b). Simionescu, 1900: 656.
Lytoceras belliseptatum Anthula, 1899: 97, pl. 6 (fig. 1), pl. 7 (fig. 1). Collignon, 1963: 12, pl.
245 (fig. 1052).
Lytoceras argonautarum Anderson, 1902: 85, pl. 7 (figs 154-155).
Tetragonites depereti Jacob, 1907: 12. Non Fallot, 1920: 243, pl. 2 (fig. 5), text-fig. 13.
Lytoceras (Argonauticeras) argonautarum: Anderson, 1938: 140, pl. 17 (fig. 3), pl. 19 (figs 1-2).
Pseudotetragonites kudrjavzevi Drushchitza, 1956: 80, pl. 7 (fig. 25), text-fig. 35. Orlov,
1958: 58, pl. 20 (fig. 4), Drushchitza & Kudryavtseva, 1960: 257, pl. 5 (fig. 2a—b),
text-fig. 64.
Ammonoceratites (Argonauticeras) depereti Collignon, 1956: 100. Thomel, 1968: 684, pl. 35
(figs 1-4), text-figs.
? Pictetia belliseptata Dimitrova, 1967: 28, pl. 10 (fig. 6).
Argonauticeras belliseptatum Forster, 1975: 144, pl. 1 (fig. 6).
Type
The lectotype, herein designated, is Kilian’s original specimen from the
Upper Aptian of the environs of Barréme (Basses Alpes), south-eastern France,
in the collections of Muséum d’Histoire Naturelle, Lyon.
Material
Two specimens, UPG B4 and UPG B5 from the Makatini Formation,
Upper Aptian, Manyola Drift on the Usutu River, at 26°49’15” S 30°13’30" E.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 305
Fig. 34. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). UPG B4. x 0,78.
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 35. External suture of Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). x 0,8.
Dimensions
Californian specimen
(after Anderson,
1933) ere ree
Holotype of A. (Ar.)
belliseptatum (after
Anthula, 1899)
Lectotype of A. (Ar.)
depereti (after
Kilian, 1892) .
French specimens
(after Thomel,
1968)
LSI.
i527.
158)
UPG B4.
UPG B5.
Description
D
250,0
32550
53,0
38,3
55,0
102,0
179,0
Wb
125,0(50)
153,0(47)
25,0(47)
19,3(50)
27,0(49)
43,0(42)
70,0(39)
40,0(—)
Wh
113,0(45)
130,0(40)
27,0(39)
16,0(41)
23,0(41)
44,5(43)
67,0(27)
7.0)
Wb: Wh
iol
1,20
7
0,96
1,04
1,08
U
87,0(35)
123,0(38)
21,0(39)
14,4(37)
19,2(34)
38,0(36)
73,0(40)
aa ee)
The largest specimen in the collection consists of one and a half whorls,
parts of which are damaged. Coiling is very evolute, and whorls increase rapidly
in height and width. The whorl section is slightly wider than high with rounded
umbilical and ventrolateral edges; the dorsal impressed zone is small.
Ornament consists of narrow, dense unbranched ribs which pass forwards
over the flanks with a slight curvature and straight across the venter. Broad,
CRETACEOUS FAUNAS FROM SOUTH AFRICA 307
A
B
Fig. 36. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). UPG BS. x 1.
low constrictions are also present, paralleling striae; the number of constrictions
per whorl cannot be determined, but on the smaller specimen, UPG BS, there
are four equidistant constrictions in one-third of a whorl. The suture is only
partially exposed; E is narrow and bifid, L is very wide and deeper than both
E and U;. The saddles E/L and I/U, are bifid and have slender stems.
Discussion
Thomel recently (1968) reviewed this species and pointed to the changes
in whorl section, relative dimensions and density of ornamentation during
ontogeny. The Zululand specimens fall within the range of relative proportions
given by Thomel (1968: 685) and are virtually identical to the figured specimens
as far as the whorl section, fine ornamentation and constrictions are concerned.
The largest known French specimen of A. (Ar.) depereti is only 102 mm in
308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 37. Ammonoceratites (Ammonoceratites) depereti (Kilian, 1892). A-—B. Copies of Kilian’s
figures of the holotype. C. Copy of Anderson’s (1938: pl. 17 (fig. 3)) figure of a juvenile
Lytoceras (Argonauticeras) argonautarum (Anderson, 1902). All figures x 1.
309
CRETACEOUS FAUNAS FROM SOUTH AFRICA
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CRETACEOUS FAUNAS FROM SOUTH AFRICA 311
Fig. 40. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45.
(Photograph supplied by R. A. Reyment.)
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
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Fig. 41. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45.
(Photograph supplied by R. A. Reyment.)
CRETACEOUS FAUNAS FROM SOUTH AFRICA 313
Fig. 42. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45.
(Photograph supplied by R. A. Reyment.)
314 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 43. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45.
(Photograph supplied by R. A. Reyment.)
CRETACEOUS FAUNAS FROM SOUTH AFRICA B15
diameter, and thus difficult to compare with the holotype of Argonauticeras
argonautarum (Anderson), the original figures of which are reproduced here
as Figure 39. A smaller specimen figured subsequently by Anderson (1938:
pl. 17 (fig. 3)) is reproduced here as Figure 37C; it appears identical to Kilian’s
lectotype (reproduced here as Fig. 37A—B), and in consequence the authors
regard A. (Ar.) argonautarum as a junior subjective synonym of A. (Ar.) depereti.
A. (Ar.) belliseptatum Anthula is based on a giant disc 325 mm in diameter.
When compared with the present material, the overall proportions, expansion
rate, style and direction of ornament are so similar that the authors believe this
to be a synonym of A. (Ar.) depereti. The remaining species referred to the
subgenus, A. (Ar.) besairiei Collignon (19495: 46, pl. 8 (fig. 1)), is also very
similar, and, indeed, Forster (1975: 144) regarded it as a synonym of A. (Ar.)
belliseptatum. The juvenile holotype has, however, a distinctly trapezoidal
whorl section with a broad, flattened venter, flattened flanks and, according to
Collignon, has straight and rather coarse ribs for the genus. A larger Mada-
gascan specimen figured subsequently (Collignon 19626: 10, pls 219-220 (fig.
956)) has a very high expansion rate, the proportions (after Collignon) being:
D Wb Wh Wb:Wh U
270,0 160,0(58) 136,0(48). 1,18 82,0(30)
Given a larger sample it might prove possible to demonstrate that A. (Ar.)
besairiei is also within the range of variation of A. (Ar.) depereti, but the authors
maintain the species separate at this time whilst acknowledging the fine division
between them.
Occurrence
As defined above, Ammonoceratites (Argonauticeras) depereti is known
from the Upper Aptian of the Caucasus, Bulgaria, southern France, California,
Mozambique and Zululand, and the Albian of Madagascar.
Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892)
Figs 38A, 44-46
Material
One specimen only, UPG B465, from the Mzinene Formation, Albian
III-IV, Mlambongwenya Spruit, northern Zululand.
Dimensions
D Wb Wh Wb:Wh U
UPRG B465*.5 052% 47,5 23,5(49) 21,0(44) 12 — (—)
84,0 40,0(48) 38,5(46) 1,04 — (—)
149,0 68,5(46) 67,5(45) 1,02 — (—)
248,0 99,5(40) 98,5(40) 1,01 — (—)
352,0 127,5(36) 134,0(38) 0,97 ~—«135,0(38)
316 ANNALS OF THE SOUTH AFRICAN MUSEUM
OL IOT
yy
Yl
Fig. 44. Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG B465.
x OLS:
CRETACEOUS FAUNAS FROM SOUTH AFRICA B17
Yj
YY
Fig. 45. Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. x 0,5.
Description
The specimen is beautifully preserved, retaining part of the original shell.
All but the last quarter whorl are septate.
Coiling is very evolute with the whorls increasing gradually in height and
width. The dorsal area of impression is very shallow. The whorl section varies
with increasing diameter as shown in the table of dimensions. It is initially rect-
angular, but with increasing diameter the venter becomes narrower and the
whorl height becomes greater than the whorl breadth. Maximum whorl breadth
is at the dorsal third of the flanks. The surface is ornamented by numerous thin,
318 ANNALS OF THE SOUTH AFRICAN MUSEUM
(NEage
\
Fig. 46. Suture of Ammonoceratites (Argonauticeras) aff. depereti (Kilian, 1892). UPG
B465. x 3,6.
fine ribs. The ribs pass radially over the umbilical wall and then forwards
across the flanks with a slight curvature, and straight over the venter. On
the venter of one of the inner whorls, where the specimen had been broken, the
ribs are minutely crinkled and are connected longitudinally by faint striae.
At a whorl height of 63 mm there are 45 ribs in a distance equal to the whorl
height; at 85 mm the figure is 42. On the inner whorls the flanks bear low folds,
so that the ribs appear to bunch. On the outer whorls this effect is less noticeable,
but two wide and shallow, barely noticeable constrictions occur. The external
suture (Fig. 46) is highly divided with narrow-stemmed saddles. The lobes are
asymmetrically bifid.
Discussion
As can be seen from the dimensions, the ratio of Wb/Wh decreases with
increase in diameter, a situation somewhat comparable to that encountered in
A. (Ar.) depereti, from which it differs in the lack of constrictions on inner
whorls (Fig. 38A), as a consequence of which the authors refer to it as A. (Ar.)
aff. depereti. It is readily separated from A. (Ar.) besairiei which is markedly
depressed at large diameters.
Occurrence
Middle Albian of northern Zululand.
Ammonoceratites (Argonauticeras) besairiei Collignon, 1949
Figs 47-48A
Argonauticeras besairiei Collignon, 1949a: 46, pl. 8 (fig. 1); 1950: 36, pl. 5 (fig. 6); 19625:
10, pl. 219-220 (fig. 956).
Holotype
The original of Collignon (1949a: pl. 8 (fig. 1)), a juvenile specimen from
the Lower/Middle Albian transition beds of Ambarimaninga, Madagascar.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 319
Material
A single fragment, BMNH C78697, from the Mzinene Formation, Albian
III, of locality 171, Mlambongwenya Spruit, northern Zululand.
Dimensions
D Wb Wh Wb:Wh U
Holotype (after
Collignon, 1949a) 43,0 21,0(49) 18,0(42) LLY 16,0(37)
Specimen 1 of
Collignon, 1950 . 70,0 35,0(50) 33,0(47) 1,06 26,0(35)
Specimen 2 of
Collignon, 1950 . 130,0 51,0(50) 48,0(47) 1,06 28,0(27)
Original of Collig-
non, 19625: pls
219-220 (fig. 956) 200,0 106,0(53) 94,0(47) 1,13 63,0(32)
BMNH C78697 __.. — 121,0(—) 107,0(—) (133 — (—)
Description
The specimen is a wholly septate fragment of a large, massive ammonite
whose original diameter must have been well over 200 mm. The coiling is very
evolute, with a narrow, shallow impressed area. The umbilicus is of moderate
size, and deep. The whorls expand rapidly and are depressed, rounded-
trapezoidal, the greatest breadth being well below mid-flank. The umbilical
wall is high, rounded, merging with broadly rounded umbilical shoulders and
lower flanks. The upper flanks are flattened and converge to a broadly rounded
venter.
Ornament consists of fine, non-crenulate, narrow, flattened ribs separated
by wider interspaces. These arise at the umbilical seam, sweep abruptly back
across the lower part of the umbilical wall and curve forwards over the shoulder
to pass across the flanks in a rectiradiate or slightly prorsiradiate direction,
with a faint convexity. They pass straight across the venter.
The external suture is not visible, but the fractured sections indicate the
presence of large L and U, lobes, together with a large septal lobe. The dorsum
shows the impression of the venter of the penultimate whorl, which was
distinctly flattened.
Discussion
At the diameter represented by the Zululand specimen, the trapezoidal
whorl section of Argonauticeras is lost, but when compared with the series of
specimens of A. (Argonauticeras) besairiei figured by Collignon (1949a—19625),
the present specimen is clearly an adult of that species.
Occurrence
Upper Aptian to Middle Albian of Madagascar, Middle Albian of
Zululand.
320 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 47. Ammonoceratites (Argonauticeras) besairiei Collignon, 1949. BMNH C78697. x 0,7.
CRETACEOUS FAUNAS FROM SOUTH AFRICA SAI
Genus Protetragonites Hyatt, 1900
Type species
Ammonites quadrisulcatus d’Orbigny, 1841.
Diagnosis
| Very evolute lytoceratids with circular to oval whorl sections, sparse to
frequent radial or curved constrictions and a shell ornamented by growth
striae only.
Discussion
Repeated attempts have been made to justify a separation of a family
Protetragonitidae Spath, 1927, from the Lytoceratidae. This has been based on
the tendency for species to develop one or more auxiliary elements in the
suture line. Schindewolf (1960: 681) and Wiedmann (1962: 17) have both
demonstrated that Protetragonites follow the same sutural development as the
Lytoceratidae, and are derived from Lytoceras sensu stricto. The authors
further follow Schindewolf and Wiedmann in regarding Hemitetragonites
Spath, 1927, and Leptotetragonites Spath, 1927, as subjective synonyms.
Occurrence
Protetragonites is a typically Mesogean genus, known chiefly from the
Mediterranean region, but also known from central Europe (Austria), the
Crimea, Caucasus, north Africa, Madagascar, and now Zululand. Its strati-
graphic range is from Tithonian to Upper Albian.
Protetragonites aeolus aeolus (d’Orbigny, 1850)
Fig. 48B—F
Ammonites aeolus @’Orbigny, 1850: 125.
Lytoceras (Gaudryceras) aeolus Jacob, 1908: 14 (pars), pl. 1 (fig. 17), non pl. 1 (figs 14-16).
Gaudryceras aeolus: Fallot, 1920: 12. Collignon, 19495: 50.
Hemitetragonites aeolus Breistroffer, 1936: 175. Breistroffer & Villoutreys, 1953: 71 (Pars):
Protetragonites aeolus aeolus Wiedmann, 1962: 24, pl. 10 (fig. 3), text-fig. 6.
Protetragonites aeolus: Wiedmann & Dieni, 1968: 31.
Type
The lectotype, designated by Wiedmann (1962: 25), is MNHP 5770 in the
D’Orbigny Collection, Muséum d’Histoire Naturelle, Paris, and is from the
Middle? Albian of Escragnolles, Var, France.
Material
Two specimens, SAS PJR/1—2, from the Mzinene Formation, Albian V,
at locality 65 along the Munywana Creek, east of Hluhluwe, Zululand, are
definitely referred to the subspecies, whilst a third specimen, BMNH C78699,
from the same horizon at locality 56 on the Mzinene River, may also belong
here.
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 48. A. Ammonoceratites( Argonauticeras) besairiei Collignon, 1949. BMNH C78697. x 0,7.
B-F. Protetragonites aeolus aeolus (d’Orbigny). B—C. PJR/1. D-F. BMNH C78699. B-C, x 1;
Ds <2: 5E-F alk
CRETACEOUS FAUNAS FROM SOUTH AFRICA 323
Dimensions
D Wb Wh Wb:Wh U
Lectotype, MNHP
SHOW : . % 1 31,0 12,5(40) 10,5(34) 1,19 14,5(47)
RMB ee 30,7 12,5(41) 8,7(28) 1,43 14,3(47)
BMNH C78699
(approximate) . 30,1 — (—) 9,3(31) — 15,0(50)
Description .
The coiling is very evolute, serpenticone, with a very wide umbilicus
(47-50% of diameter). The whorl section is depressed, reniform, with a gently
rounded venter and swollen flanks (the greatest breadth is below mid-flank).
The dorsal impressed zone is narrow and shallow.
All specimens are corroded and no external shell surface survives except
in a few areas which retain a suggestion of growth striae. There are four straight,
prorsiradiate constrictions per whorl, bounded aperturally by a thickened rib.
The sutures are undecipherable.
Discussion
Relative proportions, expansion rate, position and form of constrictions
indicate these specimens to be Protetragonites, most closely resembling the
well-known species Protetragonites aeolus (d’Orbigny). Wiedmann (1962: 23)
recognized three subspecies, P. aeolus aeolus, P. aeolus neptuni Wiedmann
(1962: 29, pl. 1 (fig. 2), text-fig. 7b), and P. aeolus aeoliformis (P. Fallot)
(Wiedmann 1962: 26, pl. 1 (fig. 1), pl. 2 (fig. 4), text-fig. 4a). P. aeolus aeoliformis
has a more or less rounded whorl section rather than reniform as in the nominate
subspecies, whilst P. aeolus neptuni has strongly ornamented inner whorls and
much more prominent constrictions.
Of other species, Protetragonites crebrisulcatus (Uhlig) (Wiedmann 1962:
19, pl. 1 (fig. 3), pl. 3 (figs 2, 4)) has rather similar overall proportions, but
bears more constrictions which efface at maturity. P. obliquestrangulatus
(Kilian) and its subspecies (Wiedmann 1926: 21 et seg.) bear many more prorsi-
radiate constrictions, whilst Protetragonites laevis Wiedmann (1962: 31, pl. 10
(fig. 4) text-figs 9-10) has a subquadrate depressed whorl section and is smooth.
Occurrence
Protetragonites aeolus and its subspecies are best known from the western
Mediterranean, i.e. the Balearics, Sardinia and south-eastern France, but there
are also records from Madagascar, and now Zululand. The species is restricted
to the Albian.
SYSTEMATIC POSITION UNCERTAIN
Genus Pictetia Uhlig, 1883
Type species
Crioceras astierianus d’Orbigny, 1842.
324 ANNALS OF THE SOUTH AFRICAN MUSEUM
Diagnosis
Loosely coiled with whorls separated throughout; body chamber may
straighten. Whorl section rounded, compressed to depressed; ornamented by
weak, irregular, feebly crinkled ribs.
Discussion
Pictetia is an enigmatic genus, known chiefly from septate fragments only
which are either curved or straight. Few good illustrations of the type species
are available and in consequence the authors reillustrated (Fig. 49) the original
of D’Orbigny (1842: pl. 115 (figs 3-5)) which is herein designated lectotype of
P. astieriana (the specimen has been commonly referred to as the holotype, but
D’Orbigny clearly states that he was aware of specimens in the Astier, Duval
and Puzos collections, although that which he figured was ‘a lévé tous mes doutes
a été découverte par M. Astier’. The authors also take the opportunity of
illustrating two of the straight fragments, doubtful paralectotypes, also from
Escragnolles, Var (Figs 50-51).
In his generic diagnosis of Pictetia, Casey (1960: 3) indicated that it had a
‘suture line with simple lobe formula IU LE, but highly complex’. Such a
formula is not typical of the Lytoceratina, which are characterized by a quin-
quelobate suture. Unfortunately there is no detailed work on the sutural
ontogeny of Pictetia; Schindewolf (1961: 678) examined specimens of the type
species, and concluded that the small incision in saddle I/U should be regarded
as an incision, and not a lobe. If this be true, then Pictetia is either to be trans-
ferred to the Ancyloceratina (some members of which have fimbriate ornament),
as did Schindewolf and Hyatt (1900: 588) before him, or retained within the
Lytoceratida as an uncoiled Lytoceras (Casey 1960: 3) in which the quadrilobate
suture is secondarily derived from quinquelobate ancestors. In Spiroceras
bifurcati (Quenstedt) Schindewolf (1961: 762, text-fig. 45) demonstrated that
two of the original five lobes became so reduced that a pseudo-trilobate condition
prevailed in adults; might not reduction in Pictetia be a similar reflection of
acquisition of the heteromorph habit? Unfortunately the Zululand material does
not help in the resolution of this interesting problem.
Occurrence
Pictetia is known from the Lower and Middle Albian of western Europe,
Transcaspia, India, Madagascar and Zululand; Collignon (19626: 13) has
recorded two species from the Upper Aptian of Madagascar.
Pictetia aff. depressa (Pictet & Campiche, 1861)
| Figs 52A-C, 53
Compare:
Crioceras depressus Pictet & Campiche, 1861: 28, pl. 45 (figs 3-4).
Pictetia depressa Casey, 1960: 5, pl. 1 (fig. la—c), text-fig. la—d (with synonomy).
Collignon, 19626: 13, pl. 221 (fig. 457).
35)
CRETACEOUS FAUNAS FROM SOUTH AFRICA
"TX °€669p HNING ‘2d410399T “(78 T “AUSIQIO.P
peste
ye
) DUDLAAIISD DIAN “6p “314
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ge
B
Fig. 50. Pictetia astieriana (d’Orbigny, 1842). BMNH C5440, wholly septate ancyloceratid-
like shaft. x 1.
CRETACEOUS FAUNAS FROM SOUTH AFRICA a0)
A B
Fig. 51. Pictetia astieriana (d’Orbigny, 1842). BMNH C5439, septate straight shaft. x 1.
328 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 52. A-C. Pictetia aff. depressa (Pictet & Campiche, 1861). SAS LJE 173. x 1.
D-F. Pictetia depressa (Pictet & Campiche, 1861). Copies of Pictet & Campiche’s original
figures of the species.
CRETACEOUS FAUNAS FROM SOUTH AFRICA 329
Fig. 53. Pictetia aff. depressa (Pictet & Campiche, 1861). Suture of SAS LJE 173. x 3,6.
Material
One specimen only, SAS LJE 173 from the Mzinene Formation, Albian
II-III, locality 171, Mlambongwenya Spruit, northern Zululand.
Description
The fragment is wholly septate, consisting of three-quarters of a whorl
only of a juvenile specimen. Coiling is very open, the whorl section depressed,
reniform, with a very small dorsal impressed zone. Ornament is subdued, and
consists of single, low prorsiradiate folds on the shell surface. The suture line
is partially exposed, showing a long E/L saddle with a narrow stem, a large L
and low L/U saddle. I is trifid (Fig. 53).
Discussion
Depressed whorl section, together with the presence of a narrow dorsal
impressed area place this specimen closest to Pictet & Campiche’s species,
the original illustrations of which are reproduced here as Figure 52D-—F; the
whorl section is more depressed in that species, whilst internal moulds are
smooth, in consequence of which the authors refer to their specimen as P. aff.
depressa. In P. astieriana (Fig. 49) the whorl section is more rounded; in
P. crassecostata Collignon (1963: 8, pl. 243 (fig. 1046)) the whorl section is
oval with broad, well-differentiated convex ribs; in P. ovalis Collignon (1963:
8, pl. 243 (fig. 1045)) the whorl section is also oval, with dense, relatively
prominent flexuous ribs. P. arcuata Collignon (19626: 13, pl. 221 (fig. 958)) is
an Aptian species based on gently curved fragments with a circular whorl
section, smooth surface and folds corresponding to the site of individual
sutures.
330 ANNALS OF THE SOUTH AFRICAN MUSEUM
Occurrence
Pictetia depressa is best known from the Lower Albian of western Europe
and Transcaspia, although also recorded from the Upper Aptian of Madagascar.
ACKNOWLEDGEMENTS
We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the
British Museum (Natural History), Dr R. Casey and Mr C. J. Wood (Institute
of Geological Sciences), Dr J. Sornay (Muséum d’Histoire Naturelle, Paris),
Professor T. Matsumoto (Kyushu, Japan), Dr I. Hayami (Tokyo), Professor
R. A. Reyment (Uppsala) and Professor D. Visser (Pretoria) for allowing us to
study material in their care and for assistance in many other ways.
We also thank General M. Collignon (Moirans), Professor Dr J. Wiedmann
(Tubingen), Mr C. W. Wright and Dr M. R. Cooper (Oxford) for their advice
and useful discussions. Dr J. A. Jeletzky of the Canadian Geological Survey
kindly supplied casts of Ammonoceratites crenocostatus and Professor
R. A. Reyment (Uppsala) provided photographs of the holotype of
Argonauticeras belliseptatum for this publication. The help of the staff of the
Geological Collections, Oxford University Museum; Geological Survey of South
Africa, Pretoria; and South African Museum, Cape Town, is gratefully
acknowledged, as is the financial support of the Trustees of the Sir Henry
Strakosch Bequest, the Royal Society, and the Natural Environment Research
Council.
Thanks are due to the Director, Geological Survey of South Africa,
Pretoria, for the loan of part of the material described.
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’
“bts
bie
I P ‘ 1
he eh a ane ) end 4
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WILLIAM JAMES KENNEDY
| | &
HERBERT CHRISTIAN KLINGER
CRETACEOUS FAUNAS FROM ZULULAND
AND NATAL, SOUTH AFRICA
THE AMMONITE FAMILY LYTOCERATIDAE
NEUMAYR, 1875
B= 77; -
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on cenns (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320
Konn, A. J. 19605. Spawning behaviour, ee ta and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4):
THIELE, J. 1910. Mollusca: B. aha Re Ee Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 74 ~ Band
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Part 13 #Deel
RE-EXAMINATION OF ARCHAEICHNIUM,
A FOSSIL FROM THE NAMA GROUP
By
MARTIN F. GLAESSNER
Cape Town Kaapstad
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RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM
THE NAMA GROUP
By
MARTIN F. GLAESSNER
University of Adelaide, South Australia
(With 3 figures) -
[MS. accepted 6 September 1977]
ABSTRACT
The fossils described by Haughton in 1960 as Archaeocyatha from the Nama Group were
later, after the study of plaster casts, considered as worm tubes or worm burrows by the
present author and named Archaeichnium haughtoni. The type specimens have now been
re-examined. They consist of agglutinated sand grains and are similar to tubes made actively
by living polychaete worms. They are not closely similar to the partly calcareous tubes of
Cloudina or to the distinctively structured Volborthella. In the absence of any taxonomically
significant characters of tube construction the systematic position of the originator of these
fossils remains uncertain.
CONTENTS
PAGE
Introduction=." 2" cas see 335
New observations . © 2... 338
INntenpretaliOniass ewan tee ee SS
Conclusions = «bh ees 341
Acknowledgements. . . . . 342
References: 2) Ge) haere 342
INTRODUCTION
Haughton (1960) described what he considered “An archaeocyathid from
the Nama System’, from two specimens (housed in the South African Museum)
of a quartzitic sandstone on Griindoorn near the Ham River, ‘a typical member
of the Kuibis beds’. Germs later re-examined the stratigraphy of the Nama
Group. From his brief description of the eastern area it appears that Haughton’s
fossils came from the Upper Clastic Member of the Nababis Formation, probably
corresponding to the Upper Clastic Member of the Kuibis Formation in the west
(Germs 1972a: 19, figs 14, 22). The interpretation of these fossils hinges on their
mode of fossilization. It is therefore necessary to summarize Haughton’s views
on this matter. He considered them as moulds on bedding planes and noted
correctly that some of the supposed moulds are ‘marked off’ by a bordering
depression and that some are lying parallel to each other, as if deposited by
current action. They are described as elongated tubular cones with circular cross-
sections. “The more complete individuals have a pointed lower end.’ In two
335
Ann. S. Afr. Mus. 74 (13), 1977: 335-342, 3 figs.
336 ANNALS OF THE SOUTH AFRICAN MUSEUM
instances, ‘two individuals appear to have been joined at this pointed end’.
Haughton noted also the presence of a number of elongate (longitudinal)
‘pustulate ridges’ and of an individual ‘mould’ appearing as an approximately
circular ridge which ‘can be interpreted as an infilling of the intervallum or space
between the internal and external walls of a cup, or as a replacement of a single
thick exoskeleton’. The second alternative was abandoned without further com-
ment and the first one adopted. Thus the longitudinal section (as seen below the
centre of Fig. 1) was interpreted as showing an infilling of a central cavity sur-
rounded by concentric inner and outer walls which are now indicated by
depressions. The solid specimens being considered as internal moulds, the present
longitudinal ridges on their outer surfaces were thought to separate what would
have been broad, rounded ridges on the inner surfaces of the vanished outer
walls. If pustules are now present, they would have originated as rows ‘of evenly
spaced pores’ in longitudinal channels. This reconstruction of the fossils from
their supposed ‘moulds’ led Haughton without much difficulty to his interpreta-
tion as Archaeocyatha.
The obvious significance of Haughton’s classification of his fossils for the
dating of the Nama Group led to their re-examination (Glaessner 1963). Those
studies were based on plaster casts of the originals. It appeared that the fossils
represented infillings of cemented tubes made, probably, by worm-like animals
which incompletely agglutinated the sediment surrounding their burrows. The
tubes were considered as generally cylindrical but they appeared to be conical
where they entered the rock matrix. The specimen (below the centre in Fig. 1,
marked M in Glaessner 1963, pl. 3 (fig. 1)) which Haughton saw as two indivi-
duals joined at their pointed ends, represents one tube dipping from the bedding
plane into the rock matrix and re-emerging with a 90° change in direction. The
other specimen, marked with an arrow on Haughton’s Plate 4 (L in Glaessner
1963, pl. 3 (fig. 1)), shows a similar change of direction. The number of longitu-
dinal ridges was correctly described by Haughton as ten to twelve on each
individual. Five to six ridges can be seen on one side but the supposed pustules
are sand grains. The ridges are also visible on several fragmentary specimens but
appear only faintly at the ends of the largest specimen. The longitudinal ridges
on the outer surfaces of the agglutinated tubes and the transverse layering of the
1 mm thick walls appear to be diagnostic for these fossils. None of the distinctive
characters of the Archaeocyatha can be seen in them. There are no pointed cones,
no traces of a former calcareous double wall, no septa or other intervallum
structures. There is no evidence of an intervallum.
The description of the partly agglutinated tubes of Cloudina by Germs
(1972a, 1972b) and recent studies of this genus and of the fully agglutinated
Volborthella (Glaessner 1976) suggested that a re-examination of Archaeichnium
would be desirable, not because they could still be considered as Archaeocyatha
or as identical with Cloudina, as subsequently suggested by Kaever & Richter
(1976: 30), but because more is now known about early tube-building worms.
The observations of the morphology of these fossils made previously on plaster
RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 331
Fig. 1. Archaeichnium haughtoni Glaessner, 1963, on a slab of quartzite from the Nama Group
(Late Precambrian). Slab B, SAM-K4812. The holotype is the specimen below the centre of
the figure. Natural size.
338 ANNALS OF THE SOUTH AFRICAN MUSEUM
casts have been supplemented by examination of the original specimens. One of
them is marked Slab A, SAM-K4813, referred to correctly as Slab A by Haugh-
ton (1960) on pages 57 and 58, but as Slab B on plate 3 (true magnification x 0,9,
not ‘above natural size’). The other is marked Slab B, SAM—K4812, which is
correctly referred to by Haughton on pages 57 and 58 and on plate 5, but as
Slab A on plate 4. The true magnification of this plate is about x 1,5, not x 5/8.
Part of a plaster cast of Slab A was previously figured (Glaessner 1963) as plate 3,
figure 2, and part of Slab B as plate 3, figure 1 and they are now shown as
Figures | (Slab B) and 2 (Slab A).
NEW OBSERVATIONS
The fossils consist of firmly cemented sand grains, mostly quartz. Although
the slabs could not be cut and without sections no measurements could be made,
the grains forming the tubes appear to be slightly coarser and more uniform than
the matrix. They must have been agglutinated prior to fossilization as they are
mostly sharply delimited from the matrix and do not grade into it on the bedding
planes. There is no indication of any previous presence of thin ‘walls’ between
the matrix and the fossils. The rock slabs consist of a slightly glassy quartzite.
Diagenetic alteration of a sandy layer has fused the tubes with the sediment
represented by the slabs, leaving them projecting in sharp semirelief from one
bedding plane. On the analogy of similarly preserved Australian fossils collected
in situ, this bedding plane was probably underlain by a silt or clay layer. The
preservation is likely to be in hyporelief. On Slab A (Fig. 2) and possibly also
on Slab B (Fig. 1) where numbers are smaller, the tubes show (as Haughton had
recognized) a preferred orientation, indicating some current sorting. This is
demonstrated by Figure 3. Sizes of the tubular fossils range from | mm to almost
5 mm in diameter. There is some flattening and some apparent twisting of tubes.
Some are cylindrical, some appear to be barrel-shaped, some were probably
conical and a few, including one near the bottom of Figure | (L in pl. 3 (fig. 1)
in Glaessner 1963) thin abruptly at one end to form a long, thin, twisted tail.
Irregular forms are more common than geometrically regular ones. Some ends
are sharp, others fade out in the matrix. Few if any clean breaks which would
indicate brittleness are visible and it is concluded that the tubes were Cea
sible and flexible, particularly where their walls were thin.
INTERPRETATION
In 1963 it was demonstrated that the fossils could not be considered as
Archaeocyatha. What are these fossils? Specimen M (Fig. 1, below centre) was
selected as the holotype of Archaeichnium haughtoni because it shows the most
distinctive characters of the new monotypic genus which are seen less clearly
among the other fossils on the two slabs. The holotype specimen can be followed
in appropriate lighting conditions over a length of 70 mm, almost to the edge of
RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 339
Fig. 2. Archaeichnium haughtoni Glaessner, 1963. Slab A, SAM-—K4813. (The lobe projecting
from the lower part of the bedding plane may be a bioturbated load cast.) Natural size.
340 ANNALS OF THE SOUTH AFRICAN MUSEUM
fo) ° °
Clo 0. 0 20° or 20 AN 60 B80 oO"
Fig. 3. Histogram of positions of 73 individual fossils on Slab A showing deviation of long
axis from modal direction.
the slab. Its maximum width is almost 5 mm. Its wall is about 1 mm thick and
shows pronounced transverse layering of sand grains. The centre of the tube is
filled with sand grains which show a much fainter and less regular transverse
fabric. The outer surface of the tube is marked with fine longitudinal ribs, but
they are visible in this specimen only faintly near the ends of the tube, which is
either smoothed by weathering or split open for most of its length. At the end
nearer the edge of the slab the ribs are preserved as impressions in the matrix,
hence they are an external sculpture, not part of an internal mould.-In the other
large specimen (bottom of Fig. 1, L in pl. 3 (fig. 1) in Glaessner 1963) the
external ribs are well preserved and straight but its internal structure is not
revealed. This tube is 35 mm long and up to 4,5 mm wide. At one end it is
narrowed and appears twisted. Several other specimens show some of these
characters such as external longitudinal ribbing or transverse wall fabric. Their
size ranges fit those of the larger specimens and the indications of original
flexibility are similar. On these grounds the author sees no reason to distinguis]
any specimens taxonomically from the typical ones.
It is difficult to decide whether these fossils are burrows in the strict sensi
of the word. There is no doubt that they were built by a worm-shaped animal
from sand grains selected and agglutinated by some organic cement which
allowed them some flexibility. The clear separation between the tube walls and
the matrix brought out by weathering of the bedding plane indicates that the
composition of the walls was different from that of the matrix. The wall material
appears to have been actively selected by the animal. There is no evidence for
calcareous walls having been dissolved during fossilization. The selection of
grains and their arrangement so as to form external and possibly strengthening
ridges shows active building from transported grains, rather than passive cemen-
RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 341
tation of a burrow wall from grains of matrix in situ. Faint transverse texture
of the infilling of the tube in the holotype specimen may indicate active filling,
but there is no confirmation from other specimens. It is likely that the tubes were
made by organisms burrowing in or living on the sediment and constructing
agglutinated walls from sand grains. It is also probable that subsequent current
action as indicated particularly by the orientation of burrow fragments on
Slab A washed the weakly cemented burrows or tubes out of a layer of sediment
and distributed them over a bedding plane. If this were the case, it raises the
semantic point whether we are dealing with trace fossils in the strict sense of the
word or whether worm tubes agglutinated from sand grains with organic cement
are properly considered as body fossils, but this is not a matter of substantial
importance. The Slabs A and B are so similar that they may well represent not
very distant parts of the same bedding plane, yet one (A) indicates current action
while the other (B) may be thought to have at least the two curved, long tubes
preserved in situ. These questions cannot be solved without further field studies
and discovery of material in situ.
CONCLUSIONS
Archaeichnium haughtoni is a name given to tubular fossils with walls
agglutinated from small detrital quartz grains. The best preserved of these tubes
show external, longitudinal, straight, fine ribs and a transverse layering of the
grains forming the wall. They may have been actively infilled. Most of the tubes
seem to be fragments washed out of the original sediment within or from which
they were built. They appear to have been redeposited on bedding planes where
they were subsequently buried by layers of sand and diagenetically fused with it.
There is no significant resemblance between this fossil and others from the Nama
Group or indeed from anywhere else. After this study had commenced, Kaever
& Richter (1976) described Buschmannia roeringi from the Buschmannsklippe
Formation of the Nama Group as Archaeocyatha. The author was able to
re-examine this fossil. Its resemblance to Praesolenopora Tsao & Zhao, 1972,
from the Late Precambrian (Upper Sinian) of China (see Tsao & Liang 1974)
was noted and it was concluded that it is probably a calcareous alga.
Archaeichnium differs from Cloudina Germs from the Nama Group (Germs
1972a, 19726). This genus builds partly calcareous, secreted tubes with only
minor, external agglutination of sand grains and transverse sculpture. ‘The tube
walls seem to have contained organic material. Their distortions suggest some
flexibility ... reminiscent of some agglutinating foraminifera with walls of
quartz or carbonate grains and much organic cement’ (Glaessner 1976: 266).
Cloudina is generally placed in the Order Cribricyathida, most of which are of
early Cambrian age. Their morphological characters make probable their
‘possible relation to the serpulids’ (Germs 1972b: 753) and suggest that they are
‘best classified as an extinct Order of the Class Polychaeta’ (Glaessner 1976: 268).
Archaeichnium differs from Volborthella which is built from invaginated cones
342 ANNALS OF THE SOUTH AFRICAN MUSEUM
of agglutinated sandy material round a central tube in distinctly sloping layers,
and from the similar but predominantly calcareous Sal/terella. This mode of tube
construction is similar in some detail to that used by living Sabellariidae.
Volborthella can be considered as the product of polychaete annelid worms.
While the tubes and the inferred tube-building activities of Archaeichnium are
comparable and compatible with those of polychaete worms of the Order Tere-
bellida (and probably others), there is as yet nothing sufficiently specific in the
known morphological characters of this fossil to support its definite systematic
assignment to the Annelida Polychaeta. It can only be tentatively attached to
them and must remain for the present incertae ‘sedis. No occurrences of this fossil
other than that at its type locality, in strata considered to be of Late Precambrian
age, are known.
ACKNOWLEDGEMENTS
I wish to thank the authorities of the South African Museum for the loan
of the type material for this study; the support of this loan by Mr P. J. Rossouw,
South African Geological Survey, Pretoria, is gratefully acknowledged. Through
the kind co-operation of Prof. M. Kaever I was able to re-examine Buschmannia
roeringi. Mr Richard Barrett, Department of Geology, University of Adelaide,
took the photographs of the rock specimens.
REFERENCES
GERMS, G. J. B. 1972a. The stratigraphy and paleontology of the lower Nama Group, South
West Africa. Bull. Chamber Mines Precambr. Res. Unit 12.
GERMS, G. J. B. 1972b. New shelly fossils from the Nama Group, South West Africa. Am. J.
Sci. 272: 752-761.
GLAESSNER, M. F. 1963. Zur Kenntnis der Nama-Fossilien Stidwest-Afrikas. Annln naturh.
Mus. Wien 66: 133-120.
GLAESSNER, M. F. 1976. Early Phanerozoic annelid worms and their geological and biological
significance. J. geol. Soc. Lond. 132: 259-275.
HAUGHTON, S. H. 1960. An archaeocyathid from the Nama System. Trans. R. Soc. S. Afr. 36:
57-59.
KAEVER, M. & RICHTER, P. 1976. Buschmannia roeringi n. gen. n. sp. (Archaeocyatha) aus dei
Nama-Gruppe Siidwestafrikas. Paldont. Z. 50: 27-33.
Tsao, RuI-CuHi & LIANG, YU-ZHOU. 1974. On the classification and correlation of the Sinian
System in China, based on a study of algae and stromatolites. Mem. Nanking Inst. Geol.
Pal. 5: 1-26.
—1
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7
6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in-chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: he pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861:
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new Species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Relaivea:
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus ...’; °. . . in C. namacolus (Fig. 10) .
(b) The prenines of prefixed surnames in all dames: hen used in the text, if not precicd
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when LONE part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
MARTIN F. GLAESSNER
RE-EXAMINATION OF ARCHAEICHNIUM,
A FOSSIL FROM THE NAMA GROUP
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