iy,

vine

Anil h

Portico a

inate Weel Wh

Ad Reng ABA!

feta en Mee

nytt ede Uf

fie i tele shel!

ILO aie

Aah ae a

SRE Marthe gt tae Cats

acide eit

ied (he

yriked letter

pony

torial!

neh

Cadets!

jaan eget «dart

i nd

Axe! Gs

AYE hestondhon at!

ithe eget

peated tine ®

iwi§h ote

well ere

% f

seybactnate ye?

gant ta eds

Cad at tet e Atrefe

Mid Din Wed pret Ae

warn atte

Bee Cee

Apres Pel

mistentie!

arses

a bogs

eat

Hotes Needing Hah

sO Athy Beds ws

a Mgt

Fin Maser H

Road

the

Mdivalertae ot

oie esfiers ana ken Oe A

saftey

NdeI AT WEY Lie AEE

the Bad et in

Ginette dogo ted

Ge he Docvttn FortcahelF

pet

ani yee Ge Gin sari

AE AE DN AE

hays

oer

AW OMe et lce

Pe aensatin die tents

ea ad me SY

arab Wieretal ebeteatinaeawamearesalar nnn

ity

YOM

ene

Cite

“

Dineeas

aN OS

ir,

Soph nie

Td ey pe

Ga ann

Tilt Ue ag x

eed SEN

Ss aneit

ora

Pere

11s

HYMAN a

Wy GEM weil

wat

Wy anens “

“ jets

Say fh

needs lest 44

erect

Hdonden sw iy

Wenn

war

wave EN

Py tly Mleth Phd #

LANL OT ae

ate CMa Petal

priven eon a

Pee bigt

abet yee Lay Me dae

LIL FN

[Ne bets Du Ma Nei)

Lipa ut

é 12%

nies AN Ee AAES EY OMe det dN Oe bow,

Pa Pea

Depiectert pi ele

gag fs reuss

SOS) 1

we hee Bee ty

Apridy Fy gy

1% dren = aes

tasty fe

“

Pde toy

errr a

PAPEL Oud OO

fai nay

Bin

Heke Ve

oP setsy

herve V4y

ene ants ea

Hye

yaya ae OY, Esyon phat

Sana ves Beet

Uy aidan

ep iN fa

me Tenia

Var enemy ¢

Shae

yh Oeste Def

ret)

Myitwase he

pate SK vey yet obi

ry AL ci eM Ay egegy fee

t vga ath

Hb a AWN

Bn ASAP NO tie $

sp ts “ jn an yoda

sg My At]

Ae ya

geese he

Pier a Den na bier

: Rien

Pere Sea

te DS

say ere

Yaak om sols

pees

TAS Deh

rh}

ote Gh

Dei iG

Be Fh cn oa Hen pS

fate

dete Wyte ye

Ted ANNES CaO

FN aM ny ike <j capt y th

Sands emake dey

eM Ne | aN cH sPM gh §

tyes

SNe ae

geeky

SAME ye

f maoeaT

mer fw iw

anny nine hee es

Aika Dead Wwe ale SiN

fins Ly dM YG MEN Ie Wi Tegel Sg a EMERY

van ¢

vay os ge fest

easy oy

Rede

PaO Hipety

Ade oeey

Leh 8 te

rere

ewer Sega

Serre

Vig tsb aL RetNat S

Feed te TN Ag wag pnts PS SPER LMA

At yt eet EN

AY UA Oe te

reap itie

cate

Pree

Horm 9.

ore)

“es

why

¥ ¥

“ Laie

Wei Gy 0d et tee We ae

te hasty

? fod

ents tow

“a

Peed A ee SAN

Wed EN Dye Mir ON ENR Ede PAV La

pees

“iy

eke

See

eer ck

ee DMN

PSA NAA ad ee

INAS ANNU Neger ee

Lar aw ey Yo

Vp seaewns

Cre ake ath eee es

a Se

ea eed

poy eer eae

PHAN tye E

pees ma

DERN ON eters ae

eke

Sek rene S

Pd SAORI

ee SINE AN

2h Oral

hgh AWS ON

ree

” me

Prdede ay “sha

aye

vad AO NY eirewreteawaw ey aW ewe ha wminuediamenm

monn Ratti e

Manian ene

: 0 yeh d toa baie

nity

ee dS eT

ena cee nt Bb HY

SONY try

Sede deep ND

vou

aye

eas

Ay t,

yin

ener

mi yore dare Ab at

ytey a

PARANA OP ators Ys Ns

yhaterde Pel heheh th

HE AWN AN,

as ott a

Fah aly da Ws

eres tae

arr

or eae CD

ad ve MIN cy Ye

AASV

bytes

aly

DAW paige 0 2%

Hr Nateatns

Ms)

MG Se Me

Wa Th Qs

Vegeevee

soe

He Nath

peebelns

He spade INES UE

aera

Bh Sadi chidee 4) St

wetyih

ENeegtia!

Yerba Verne

Shei Nya E

Cer Paar ve

Sale Suter

CI YE ie

Dike seathdh emotes

Ai\ag

j Le eMten enh

ste

Ryo g 7 ’

Pita

Aha Re Nudthg Aly

ey td bof 2 FWW NY (RY tana STA

they

29.

se EW

iy Sinn oth om

oyna

edition

MOIR ADT IN aT

nana’ sieerten Ni

aves

Saath fo

Meare

capi hes yoik

Pye ek IN MOMS G

i teat et Zh oe

wiih hie ot

he Nene Neath

aot th le AY 1%

: wie

Se gee the vv be

Baeth had aah atte a

eee es

wae

RoeNeds othe) oh oe

Rar “ath th pith ate

Dew edthnny

toners

Nate

pre

sHazi

AN ASM a ait 0

zr ayn aay hs aor bet

ay REA ReS 2s Oa AP aC

Aver ie et eM ses 8 hag te

eertnaty ys

Se EK teh ote eb SLT AB ken

pe th FS Mita tot

isthe Wo hat

y 9 at

pint eck anh el HT ANY fr! <p

Bert rruts Bay SAY eben foie 8

ANE te SIGE,

Vat 71's

Oot unis) AE hm Fhe ee ty Be

% a yy Matha athe A ala Mh

Ae pep thg Md ofigihon yh

Ny mat

2B mia aithorth

AT beakers Daren nthe he Aaa

hel sooe

EYL anade

Ueshatha tts “ale Pe

AE os HE te 9

A eee wE he Ne

y Kier

reba AY aera:

Stee cep eent woe we

CoS eas

Fue Wy gar he kash wot Sth

SUNN ate Ps ete te ie a

SR athe tet

ry ten nad tat Fal oS Ay Ty Ths,

apache ov aD Winsett

mth (GaSe 9%

Dee,

ats + TIA

DV Aaylye ash

Nate oun)

hey at

fae ee ee

naw

Gow taeee

PIM MN Peg Ae he ey eA

Weed ONE

tes

see a

le Seated ee eee

“mee

cf RRM eat bea fh al

ye ORES,

FLAW y os Tie Preheat

mont atl pase

eed SPA D say Ue AD DER Sate a = ae

et Tene Serre

hs =~

ALS Na iret SoBe he

ede St! oy td ee

Sa AL Bike ent oh

Sreahenth adie at

ens Ma

We oh Rely Za Detar le

Wath ald anh tg

Mas Tani eden Mant ys

pl eat Pake hy

Enews

whats bean

oD nun th hore

patedt nae gTnn a Mw baal mma ibe

Say aE tom

pe emeticeie

Wha ete bee ew

aban

pata 8 Na Rhee Ting

Cee Be ala 9 vette!

wee Stee ene eee Peet Cee ae

Te athe Rach os Ga

Ape ha aap Nee Sete athe te

“a an pean e tn at

4 ue

+ Nis

ma tn ule

a ee ee

gigi 2)

et eet

. tin oy

ee oe

wo eBiet %

aceite Rear tun teaaem ar eM A

Fe citar aN aD

Wa tier «Vet ct 6h =e? = <8

hag hen Ee AD nts

a atten ©

did

th eee

RA eRE Ska

ea er

me Ne

scone ape +:

a vain

ANNALS OF THE

SOUTH AFRICAN MUSEUM

VOLUME 101

ANNALE VAN DIE

SUID-AFRIKAANSE MUSEUM

BAND 101

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 101 BAND

TREN

2S B:8B:8e

Jt g

@ S

Soin noi N°

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1991-1992

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA

BY THE RUSTICA PRESS (PTY) LTD, NDABENI, CAPE

D2109

LIST OF CONTENTS

Page

DE Decker, A. H. B., KACZMARUK, B. Z. & Marska, G.

A new species of Calanus (Copepoda, Calanoida) from South African waters.

(RublishedeAucustHOoi sia sete. Cree ease ace as Ua Re cli ae err ee eee 7)

GriFFITHS, C. L. see STEWART, B. A.

KACZMARUK, B. Z. see DE DECKER, A. H. B. et al.

KENNEDY, W. J. see KLINGER, H. C.

KLINGER, H. C. & KENNEDY, W. J.

Cretaceous faunas from Zululand and Natal, South Africa. Barremian represen-

tatives of the ammonite family Ancyloceratidae Gill, 1871. (Published March

LOD SS St Near eee PIA rt Get OPI Str Sree eth PM aed ae MC Sees 7

Marska, G. see DE Decker, A. H. B. et al.

POORE, G. C. B.

Neoarcturus Barnard, 1914: diagnoses of the genus and its type species and substi-

tution for the nomen nudum ‘Microarcturus’ Nordenstam, 1933 (Isopoda,

Arctumoac).(Eublished: Marcht99i))i oc. een ee ei one ae ae ee 1

RAYNER, N. A.

Revision of the freshwater diaptomid genus Lovenula (Crustacea, Copepoda) in

PatniCae(EUOLIShed Septemberl992 ew nce ce cine ars oe ae ence eae 297

SHONE, R. W.

Trace fossils of the ?Early Ordovician Sardinia Bay Formation, Table Mountain

Groups (Rublisneddume 199M eas ck Sa Sse Aieeare «tc a eee es ae ee Oe 9

STEWART, B. A. & GriFFITHS, C. L.

Four new species of the genus Paramelita (Amphipoda, Crangonyctoidea) from

SouchyAiricay (Published MarchWlO92 yi: Sear are es oe ae no eee 139

THANDAR, A. S.

The South African Museum’s Meiring Naude cruises. Part 18. Holothuroidea.

RBublishea une M992) ene oad eee cies Pee rts wane crap aces Bie tea 159

WILLIAMS, G. C.

The Alcyonacea of southern Africa. Gorgonian octocorals (Coelenterata, Antho-

ZO) EUOnNShe Galva 992s) ors tes os 2 enn at tes ee ere ole tesla ey a ert ae 181

WITTMER, W.

Die in Afrika vorkommenden Colotes Erichson mit einfarbig metallischen Fligel-

decken (Coleoptera, Malachiidae). (Published September 1991.).............. 45

Volume 101 is complete in 9 parts.

t : ots

f =, Pay el

au }

; 4

+ A 4 ;

: 1

j rs

: = !

= |

i Jes

(

{

3 q

‘

f ) <

i {

i a a 1

“

. i

{ .

}

VOLUME 101 PART 1 MARCH 1991 j ISSN 0303-2515

JM SON a

JUN 9 6 1991

Lirneparits

I BRARIE

ANNALS

“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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes. . .’

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

‘As described (Smith 1969a, 19696; Jones 1971)’

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 19692).

For books give title in italics, edition, volume number, place of publication, publisher

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 #£Band

March 1991 Maart

Part 1 Deel

QO) SS

Stouin now 3%

NEOARCTURUS BARNARD, 1914: DIAGNOSES

OF THE GENUS AND ITS TYPE SPECIES AND

SUBSTITUTION FOR THE NOMEN NUDUM

‘MICROARCTURUS’ NORDENSTAM, 1933

(ISOPODA, ARCTURIDAE)

GARY C. B. POORE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

i, DOES; 5-8), S02, 425, 6, tat), SMEs, 5, 7S),

Ge ask) TOO), By (GED), NOGES) TIGER, 5. Wo...)

(=D), ISAS), DAO, 5), Mil, SC), 2G), 33,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 121 0

Printed in South Africa by In Suid-Afrika gedruk deur

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

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

D213

NEOARCTURUS BARNARD, 1914: DIAGNOSES OF THE GENUS

AND ITS TYPE SPECIES AND SUBSTITUTION FOR THE NOMEN

NUDUM ‘MICROARCTURUS’ NORDENSTAM, 1933

(ISOPODA, ARCTURIDAE)

Gary C. B. POORE

Department of Crustacea, Museum of Victoria, Swanston Street,

Melbourne, Victoria 3000, Australia

(With 3 figures)

[MS accepted 27 June 1990|

ABSTRACT

The arcturid isopod genus Neoarcturus Barnard and its type species N. oudops are

rediagnosed largely as follows: body not geniculate between pereonites 4 and 5; pereopod 1 a

gnathopod; propodus ovoid, with mesial and posterior setae; dactylus stout, with mesial setae

and well-developed unguis; pereopods 2—4 with double row of long setae, dactylus with

elongate unguis; male pleopod 1 exopod thickened, with lateral setal row, apex bilobed, inner

lobe with long plumose setae, outer lobe carrying distal end of open duct on posterior surface;

endopod shorter, membranous; male pleopod 2 with short appendix masculina; uropod with

two rami.

Twenty-eight species previously assigned to the nomen nudum ‘Microarcturus’ Nordenstam

are reassigned, most to Neoarcturus.

CONTENTS

PAGE

METERO CU CLIO Mee ee er ee ae aa ict Re Be Sve der ay Uta 1

ON See Wrap OE ey ee een pet ty ty al ace te tate oh sa ctl ahah Susy Shas 2

Neoarciurus Barnard: 1904) 5228 oe eee er ee oe eV ak Pe 2

Neoareiurus oudops Barnard, 1914 2. 05 bse os ee ee eee 4

EVEMBO WCU PEMENtS it. Sas ce Peta caine ae Pee ee as fe Pema oe ahs 7

1 EN SHETRET TOSS SS Pea One oh tary Res Pe alae ae Ney SRL A REE ee 7

INTRODUCTION

In 1914 Barnard erected the genus Neoarcturus for a single species,

Neoarcturus oudops. The generic name has not been used for any other species.

Nordenstam (1933) proposed the name ‘Microarcturus’ for a genus to

accommodate twelve (plus possibly two) species of Arcturus Latreille and

Antarcturus zur Strassen. A type species was not selected by Nordenstam nor by

any subsequent worker. The name is therefore a nomen nudum (ICZN

Article 13 (b)). In spite of this the name has been widely used and, in addition

to the species listed by Nordenstam, 13 species have been assigned to

Microarcturus or described as species of it (Hale 1946; Kensley 1975, 1977,

1978a, 1978b, 1984; Kussakin 1967, 1982).

Ann. S. Afr. Mus. 101 (1), 1991: 1-8, 3 figs.

D ANNALS OF THE SOUTH AFRICAN MUSEUM

As well as being a nomen nudum and therefore unavailable, Nordenstam’s

name poses additional problems, not the least being that the generic concept is

poorly defined. He diagnosed the genus as follows: ‘Antennae shorter than the

body, with a short flagellum consisting of three joints (occasionally two or four).

Abdomen short, never longer than the last four pereion segments together,

(pleotelson posteriorly pointed or cleft). Small forms.’

These characters are weak and not clear cut. At least two of the three

species examined by Nordenstam (‘M.’ rugosus and ‘M.’ stebbingi) and some of

the others listed by him can be readily assigned to the older genus Neoarcturus

Barnard, 1914. The remaining species are a heterogeneous assemblage, some of

which have already been placed in other genera (Schultz 1981).

I suggest that the name, already a nomen nudum, be allowed to lapse. Since

Neoarcturus—a suitable substitute name for many species of “Microarcturus’—

is poorly known, its type species is here redescribed and a new generic diagnosis

presented. The generic positions of all species previously allocated to

“Microarcturus’ are discussed on the basis of the literature.

The material of Neoarcturus oudops examined in the course of this study

was lent by the South African Museum (SAM) and Nordenstam’s material of

‘Microarcturus’ rugosus and ‘M.’ digitatus by the Swedish Museum of Natural

History. Redescription of Nordenstam’s species is beyond the scope of this short

contribution.

DESCRIPTION

Neoarcturus Barnard, 1914

Neoarcturus Barnard, 1914: 213-214, 216; 1940: 508. Nordenstam, 1933: 115.

‘Microarcturus’ Nordenstam, 1933: 128. Hale, 1946: 199-200. Kensley, 1975: 47, 54; 1984:

234-238.

Antarcturus (Microarcturus) Kussakin, 1982: 310.

Type species. Neoarcturus oudops Barnard, 1914 (by monotypy).

Diagnosis

Body not geniculate between pereonites 4 and 5. Head coalesced with

pereonite 1. Pereonites 2, 3 and 4 subequal in length, broader in female than

male. Pleonal segments completely fused to each other and to pleotelson.

Antenna 1 flagellum with short first article, second long article with distal

aesthetascs, and third minute article with aesthetascs. Antenna 2 with flagellum

of three articles, the last an elongate claw. Pereopod 1 a gnathopod; propodus

ovoid, with mesial and posterior setae; dactylus stout, with mesial setae and

well-developed unguis. Pereopods 2—4 with double row of long setae; dactylus

with elongate unguis. Pereopods 5-7 ambulatory. Male pleopod 1 exopod

thickened, with lateral setal row, apex bilobed, inner lobe with long plumose

setae, outer lobe carrying distal end of open duct on posterior surface; endopod

DIAGNOSES OF THE ISOPOD GENUS NEOARCTURUS 8}

shorter, membranous. Male pleopod 2 with short appendix masculina. Uropod

with two rami.

Remarks

The similarity between the species that should now be placed in

Neoarcturus was discussed by Kensley (1984) under the name ‘Microarcturus’.

He noted the problems with Nordenstam’s definition and suggested that they

were held together by general habitus and male pleopods rather than the

characters used by Nordenstam. He figured 12 South African species differing

mostly in ornamentation and ranging from granulate to obviously tuberculate.

Tuberculation was used by Schultz (1981) to differentiate some of these species

as members of his new genus Rectarcturus. Given the wide range of forms seen

in the suite of South African species, separation of Rectarcturus from

Neoarcturus is hard to justify.

Most genera of the Arcturidae have been loosely defined. It is my belief

that characters such as the form and setation of pereopod 1, the nature of the

male pleopods, and the uropodal rami will shed light on affinities but, until the

family is revised, relationships cannot be fully explored.

The species previously assigned to “Microarcturus’ may be discussed in

groups as follows.*

1. Species that are almost certainly, in my view, placed in Neoarcturus: Ant-

arcturus hirticornis Monod, 1926; Arcturus stebbingi Beddard, 1886; Micro-

arcturus barnardi Kensley, 1984; M. biserialis Kensley, 1978; M. dayi

Kensley, 1977; M. halei Kensley, 1984; M. longispinus Kensley, 1984; M. maw-

soni Hale, 1946; M. nordenstami Kensley, 1984; M. quadriconus Kensley, 1975;

M. rugosus Nordenstam, 1933; M. tannerensis Schultz, 1966; M. youngi

Kensley, 1978.

2. Species assigned to or placed very close to Rectarcturus by Schultz (1981):

Arcturus kophameli Ohlin, 1901 (type species); A. patagonicus Ohlin, 1901;

Antarcturus similis Barnard, 1925; M. laevis Kensley, 1975; M. ornatus Kensley,

1975. These could also be species of Neoarcturus.

3. A species placed in Pseudarcturella Tattersall by Hale (1946): Arcturus ocula-

tus Beddard, 1886.

4. A species placed in Pseudidothea Ohlin, 1901 (Pseudidotheidae) by Sheppard

(1957): Microarcturus scutatus Stephensen, 1947.

5. Species, in Schultz’s judgement, perhaps related to Acantharcturus Schultz,

1981: Antarcturus acanthurus Monod, 1925; M. digitatus Nordenstam, 1933.

These differ from the previous groups in having a uniramous uropod.

6. Another species with uniramous uropod very different from all others but

included in ‘Microarcturus’ by Hale (1946): Arcturus serrulatus Whitelegge,

1904.

* See note added in press.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

7. Species too poorly known to be assigned to genus: Arcturus alcicornis White-

legge, 1904; A. dentatus Whitelegge, 1904; A. nodosus Whitelegge, 1904;

A. simplicissimus Whitelegge, 1904; M. abnormis Kussakin, 1967; Antarcturus

kilpoae Kussakin, 1971.

Neoarcturus oudops Barnard, 1914

Figs 1-3

Neoarcturus oudops Barnard, 1914: 214-216, pls 18C, 19B; 1920: 397-398; 1940: 493.

Microarcturus oudops Kensley, 1978a: 130-131 (not fig. 3 which is male pleopod 1 of another

species of Neoarcturus); 1978b: 28 (part: fig. 13A is female of an undescribed species of

Neoarcturus); 1984: 252 (part: figs 19B, C and probably D, E and F are of a male of

N. oudops; fig. 19A is female of an undescribed species of Neoarcturus).

Material

Holotype, male, off Cape Point, South Africa (SAM-—A69, with 2 slides).

1 male, eastern South Africa (SAM-—A15472, only pleopod 1 on slide seen).

1 female with slide, eastern South Africa (SAM-—A39991 taken from SAM-—

A17819; remaining 2 males, 3 females, 4 juveniles are another species).

Diagnosis

Integument granulate. Pleotelson with lateral subapical wings, apex broadly

acute (see Barnard 1914, pl. 19B; Kensley 1984, fig. 19B-C). Eyes absent.

Pereopod 1 propodus with c. 16 setulate mesial setae loosely arranged in pairs,

posterior margin with c. 16 setulate setae in two rows; dactylus with oblique

mesial row of seven setae, one separate on anterior margin, four on posterior

margin, unguis one-third total length. Pereopod 2 unguis two-thirds total length

of dactylus, shorter in pereopods3 and4. Male pleopod1 exopod with

12 lateral simple setae plus four plumose setae, outer lobe with 16 teeth in a

curved row on anterior face, inner lobe of apex with six setae; endopod with

four apical plumose setae. Male pleopod 2 appendix masculina with club-shaped

apex.

Remarks

There has been some confusion over the identity of the South African speci-

mens referable to Neoarcturus oudops. The holotype and some of the material

identified by Kensley were lent to me by the South African Museum. Ten speci-

mens in one lot from the east coast of South Africa (SAM-—A17819) belong to

two species differing most obviously in the shape of the pleotelson and setation

of the male pleopod 1. Both species have been illustrated by Kensley under the

name Microarcturus oudops (1984—see synonymy above). The female habitus

and male pleopod 1 of other material figured by Kensley (1978a, 1978b) were

also not of N. oudops.

The specific diagnosis given here therefore is intended to differentiate the

species primarily from others from South Africa. More detailed information is

contained in the figures of the type material and in the original description.

DIAGNOSES OF THE ISOPOD GENUS NEOARCTURUS 5

—>

SSS

ESS

SSS

Fig. 1. Neoarcturus oudops Barnard, male holotype (from Barnard’s slides which are not

clear). A. Last two articles of antennal. B-C. Left and right mandibles. D-E. Maxil-

lae 1,2. F. Left maxilliped. G. Left maxillipedal endite. H. Right pleopod 1, posterior

view. I-J. Apex of endopod of pleopod 1, posterior and anterior views. K. Pleopod 2 (no

setae shown).

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Wy Wye

SS Y/ Wye

—

ROSS

EES

sla

Dos

LOL

Ess} Z

> 2

ee,

—————

Fig. 2. Neoarcturus oudops Barnard, female, SAM—A39991 (left limbs). A-B. Antenna 1 and

flagellum. C—D. Antenna 2 and last article of flagellum. E. Pereopod 1. F—G. Pereopod 1

propodus, mesial and lateral views. H. Pereopod 1 dactylus, mesial view. I. Pereopod 2.

J. Pereopod 2 dactylus. K. Uropodal rami.

DIAGNOSES OF THE ISOPOD GENUS NEOARCTURUS 7/

Fig. 3. Neoarcturus oudops Barnard, female, SAM—A39991 (left limbs). A-—D. Pereopods 3,

4,5, 7. (Fig. 3 and Fig. 2E, I to same scale.)

ACKNOWLEDGEMENTS

This paper was made possible with the support of a grant from the Austra-

lian Biological Resources Study which is acknowledged. The manuscript

benefited from discussions with Helen Lew Ton and Brian Kensley, and I thank

them for their comments.

REFERENCES

BARNARD, K. H. 1914. Contributions to the crustacean fauna of South Africa. 1. Additions to

the marine Isopoda. Annals of the South African Museum 10 (7): 197-230.

BARNARD, K. H. 1940. Contribution to the crustacean fauna of South Africa. XII. Further

additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identifi-

cation of hitherto recorded marine and freshwater species. Annals of the South African

Museum 32 (5): 381-543.

Hate, H. M. 1946. Isopoda—Valvifera. Reports of the British, Australian and New Zealand

Antarctic Research Expedition (Series B, Zoology, Botany) 5 (3): 161-212.

KENSLEY, B. 1975. Marine Isopoda from the continental shelf of South Africa. Annals of the

South African Museum 67 (4): 35-89.

KENSLEY, B. 1977. New records of marine Crustacea Isopoda from South Africa. Annals of

the South African Museum 72 (5): 239-265.

KENSLEY, B. 1978a. The South African Museum’s Meiring Naude cruises. Part 7. Marine

Isopoda. Annals of the South African Museum 74 (5): 125-158.

KENSLEY, B. 1978b. Guide to the marine isopods of southern Africa. Cape Town: Trustees of

the South African Museum.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

KENSLEY, B. 1984. The South African Museum’s Meiring Naude cruises. Part 15. Marine

Isopoda of the 1977, 1978, 1979 cruises. Annals of the South African Museum 93 (4):

213-301.

KussAKIN, O. G. 1967. Fauna of Isopoda and Tanaidacea in the coastal zones of the Antarctic

and Subantarctic water. [Translation by the Israel Program for Scientific Translations,

Jerusalem, 1968.] Biological Reports of the Soviet Antarctic Expedition (1955-1958) 3:

220-380.

KuSSAKIN, O. G. 1982. Marine and brackish-water Crustacea (Isopoda) of cold and temperate

waters of the Northern Hemisphere. Suborders Anthuridea, Microcereberidea, Valvifera,

Tyloidea. Opredeliteli po Faune SSR 131: 1-460. [In Russian. ]

NorRDENSTAM, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudido-

theidae, Arcturidae, Parasellidae and Stenetriidae mainly from the South Atlantic.

Further Zoological Results of the Swedish Antarctic Expedition, 1901-1903 3: 1-284, pls

1-2, errata.

ScHULTZ, G. A. 1981. Arcturidae from the Antarctic and Southern seas (Isopoda, Valvifera).

Part I. In: Biology of the Antarctic Seas X. Antarctic Research Series 32: 63-94.

SHEPPARD, E. M. 1957. Isopod Crustacea. Part II: The suborder Valvifera. Families: Idothei-

dae, Pseudiotheidae and Xenarcturidae fam. n. With a supplement to isopod Crustacea,

Part I, the family Serolidae. ‘Discovery’ Reports 29: 141-198.

NOTE ADDED IN PRESS

Since the completion of this paper Brandt’s (1990) paper on Antarctic valvi-

ferans has been published. She too recognized the priority of Neoarcturus over

Microarcturus. She erected five new species: Neoarcturus elongatus, N. minutus,

N. robustus, N. sclerosus and N. stebbingnordenstami. The first four of these

were figured and described and the last is based on the 146 specimens (from

Shag Rock Bank and South Georgia) that Nordenstam (1933) called Microarctu-

rus stebbingi Beddard, 1886. No holotype or type locality was selected for this

species.

Brandt placed Antarcturus hirticornis Monod, 1926, which I suggest is a

species of Neoarcturus, in a new genus, Fissarcturus Brandt.

These species and N. mawsoni Hale, all from Antarctica, fit the generic

diagnosis given here but differ from N. oudops in structure of the male pleopod

1 exopod. They lack teeth on the distal anterior surface and have different

setation. The appendix masculina is acute rather than blunt.

BranpT, A. 1990. Antarctic valviferans (Crustacea, Isopoda, Valvifera). New genera, new

species and redescriptions. Leiden: E. J. Brill.

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

ete.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

eam . 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 ae ee by

initials or full names

e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

GARY C. B. POORE

NEOARCTURUS BARNARD, 1914: DIAGNOSES OF

THE GENUS AND ITS TYPE SPECIES AND

SUBSTITUTION FOR THE NOMEN NUDUM

‘MICROARCTURUS’ NORDENSTAM, 1933

| (ISOPODA, ARCTURIDAE)

1x POLUME 101 PART 2 °&#&JUNE 1991 | ISSN 0303-2515

OF THE SOUTH AF RICAN”

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes... .”

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 °#£4Band

June 1991 Junie

Part 24 Deel

MRACE FOSSILS OF THE 7EARLY

ORDOVICIAN SARDINIA BAY FORMATION,

TABLE MOUNTAIN GROUP

RUSSELL W. SHONE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

i, WOES, SO), OleWs FES. 3, haat), SOLS, 5. 7D),

GG. tos), CED), 8, CCE, 7), MOOS), INGED, 5.7, Bos),

14(1-2), 15(4-5), 24(2, 5), 27, 31(1-3), 32(5), 33,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 122 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

D340

—————

TRACE FOSSILS OF THE 7EARLY ORDOVICIAN

SARDINIA BAY FORMATION, TABLE MOUNTAIN GROUP

RUSSELL W. SHONE

Department of Geology, University of Port Elizabeth, Port Elizabeth

(With 12 figures)

[Paper presented at the Palaeontological Society of southern Africa Symposium, Cape Town,

September 1986]

ABSTRACT

Several sandstones in the 180 m thick Sardinia Bay Formation of the lower portion of the

Table Mountain Group, contain trace fossils. Endichnial burrows are most common, including

traces that resemble the genera Ophiomorpha, Thalassinoides, Diplocraterion, Skolithos, and

Fascifodina. Hypichnial traces on the sole of one sandstone are tentatively identified as Cru-

ziana. A number of enigmatic traces occur on bedding surfaces. Three trace fossil associations

are recognized in the Sardinia Bay Formation. Two of these correspond broadly with the Cru-

ziana and Skolithos ichnofacies models of Seilacher (1967) and Frey & Pemberton (1984), but

differ significantly in detail. The differences are ascribed to reduced circulation resulting in

anoxic conditions in parts of an ancient epeiric sea. Barren sandstone units in the Sardinia Bay

strata represent former high-energy depositional environments and shifting substrates hostile to

sediment-dwelling organisms. Pyritic trace-free mudrocks may have been deposited in biologi-

cally sterile parts of the epeiric sea.

CONTENTS

PAGE

PS PeEClOMICEIO HME M Ur San ae re gs ninth eas sae oie Gatiohe 10

Mecationrand peneraliseOlogy i.” ois ei cee wee cole cle es thete oops toe 10

Vertical facies changes and depositional model................. it

Age and correlation of the Sardinia Bay Formation............. 12

BICSGRIPEIONSFOM LACE LOSSIIS. 658 ok ne ie see Se Sian iE eee ee ee 12

2 DCU OTELDN TTL OT a seesunn Ae a ae ae ee en EE 12

1 COLES SEDI LIGS SS Oeste OE 1 CPR Oe RC a a eS er 14

DETHLOCEECHION ec caine ene Skee a ee gsi hs Oe a Se pas, eas 15

SESUTT DDS oe AIS ee Ak Led iN a GSR Se EE RPE PoE ee 15

hmivermitorm,sand-filled tubes... 22. .2.5-52: 420.0026 sees os 16

(CEUTA OR ai Ee CECH Oe Soe ee Ee 18

Horizontal traces on upper bedding surfaces................... 18

Ira ICERPOULEL CS Fete eS. rt ee Fede eh Scheer See Chin wt ghee ieee vnalin cin rey ote 19

Ichnofacies models and depositional environments ................. 20

INNCHCMNOFACICS CONCEDE. Care a6 tee eo eee eke Hee sles colon 20

SCOUTEOSDICIMOLACIESE <.4.447 Seas ek a i cic HNSS Maas Aes 22)

GAA ANCHMOLACIES tt specie ak eye eninge, Seta dey ape Bras sae ogtaee 22

Trace fossil associations in the Sardinia Bay Formation.......... ip

PAEMMOMICCOCINENUS Cie. cima cer eae Suna scl Oa a cea es Sake ee et: 24

[2 EN ETSTTG SS, Arete tens ear e ik oer ole o Me OU N o oR eae a 24

Ann. S. Afr. Mus. 101 (2), 1991: 9-25, 12 figs.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

LOCATION AND GENERAL GEOLOGY

The Sardinia Bay Formation crops out near Port Elizabeth in the vicinity of

Sardinia Bay (Fig. 1). These strata represent part of the lowermost Table Moun-

tain Group in the Port Elizabeth area. The probable base of the Table Mountain

Group is represented by an outcrop of granite overlain by conglomerates con-

taining boulders of granite and quartzite, which lies about 2 km west of Sardinia

Bay. The thickness of strata separating these conglomerates and the base of the

Sardinia Bay Formation is not known.

Rocks of the Sardinia Bay Formation occur on the northern limb of a major

asymmetrical anticlinal structure in the Table Mountain Group formed in response

to compressional forces directed from the south. There is evidence of a subsequent

east-west compression in the Sardinia Bay area (Shone 1983: 65-71). Rocks of

the Sardinia Bay Formation appear to have been subjected to low-grade load

metamorphism prior to the folding episodes (De Swardt & Rowsell 1974; Shone

1983: 125).

Characteristic rock types are metaquartzites and phyllites. The emphasis on

ichnology and sedimentology in this paper favours reference to them as sand-

stones and mudrocks.

Table Mountain ELIZABETH

Group

;

Sardinia Bay

O km 10

_———)

+-34°10'

25°30:

Fig. 1. Map showing location of Sardinia Bay.

TRACE FOSSILS OF THE SARDINIA BAY FORMATION i

SCALE INMETRES

LEGEND

25+] Conglomerate

ee] Sandstone

GE Mudrock

Z| Planar

ee cross-beds

— Low-angle

-——] sand wave

erosion planes

(ar| Trough cross-

beds

Horizontal

lamination

Hummock

cross- y

stratification

GRAIN SIZE

180

GRAVEL

COARSE

MEDIUM

Skolithos

Fascifodina

Cruzianag

Ophiomorpha

Thalassinoides

SILT/MUD

FACIES INTERPRETATIONS

SHALLOW WATER TURBIDITES

deposited bya density current

generated via wave loading

liquefaction and ebbing storm surge

INNER SHELF MUD AND SAND

deposited under the influence of

tidal currents local wind-generated

currents andwaves. Restricted biota

-Shallow marine sediment-dwellers

INNER SHELF MEGARIPPLE FACIES

INNER SHELF MUD AND SAND

SHELF SAND WAVE FACIES

deposited by tidal currents with a degree

of time - velocity symmetry

INNER SHELF MUD AND SAND

AND INNERMOST SHELF MUDS

anoxic hypersaline conditions in oreas

shorewards of the tidal reach.

Elswhere, tidal currents, wind-

generated currents and waves

increased circulation sufficiently for

colonization by sediment- dwellers

INNERMOST SHELF MUD FACIES

dark, pyritic silt- streaked muds deposited

in a low-energy, anoxic, hypersaline

environment beyond tidal reach

INNER SHELF MUD AND SAND

small tidal megripple fields colonized

by sediment -dwelling organisms.

Muddy anoxic where tide-damped

SHOREFACE DEPOSITS associated

with line of wave break, including beach,

beach runnel andlower shoreface lag

deposits

INNER SHELF_MUD AND SAND

INNER SHELF MEGARIPPLE FACIES

deposited by weak fidal currents

assisted by storm- generated flow

Fig. 2. Simplified stratigraphic section of the Sardinia Bay Formation showing facies

interpretations and units containing trace fossils.

VERTICAL FACIES CHANGES AND DEPOSITIONAL MODEL

The 180 m thick Sardinia Bay Formation (sensu Shone 1979, 1983) can be

subdivided into a number of facies on the basis of geometry, lithology, sedimen-

tary structures, trace fossils and palaeoflow data (Shone 1983, 1987). Figure 2

shows a simplified stratigraphic section and the corresponding facies interpret-

ations. Bioturbated units, indicated by the generic names of the more important

trace fossils (Fig. 2), are limited to planar cross-bedded sandstones attributed to

the migration of small megaripples in an inner-shelf depositional environment.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Elsewhere, the sandstones are fossil free but contain sedimentary structures that

denote high-energy hydraulic conditions normally associated with beaches, tidal

sand-waves (cf. Allen 1980), storm-affected shelf floors, and ebbing storm-surge

turbidity flows (Shone 1983, 1987).

AGE AND CORRELATION OF THE SARDINIA BAY FORMATION

The absolute age of the Sardinia Bay Formation is not known. On the basis

of extremely tenuous correlation with the Graafwater Formation of the Western

Cape (Rust in Tankard et al. 1982: 339), an Early Ordovician age is possible.

Even if the Graafwater—Sardinia Bay correlation is untenable, the Sardinia Bay

rocks are likely to be older than the Ashgillian age proposed for rocks of the

Upper Table Mountain Group (cf. Cocks et al. 1970). The granite outcrops west

of Sardinia Bay have not yet been dated but, if sufficiently fresh material can be

obtained, a lower age limit for the Sardinia Bay Formation could be deduced.

The absence of body fossils other than enigmatic stromatoporoid fragments

(Shone 1983: 257-259) aggravates the dating problem still further.

DESCRIPTIONS OF TRACE FOSSILS

Classification of trace fossils cannot be accomplished in the same manner as

zoological classification (Hantzschel 1975: W16), and trace fossil names are not

recognized in the International Code of Zoological Nomenclature (Ekdale et al.

1984: 17). In the case of the Sardinia Bay Formation, the identities of the organ-

isms responsible for producing the traces are not known; trace morphology is all

that can be determined. Even so, individual traces can seldom be named with

confidence and some of the trace fossils could represent hitherto undescribed

forms. In certain instances, trace fossils in the Sardinia Bay Formation exceed

the time ranges known for comparable ichnogenera from other strata. The

descriptions that follow are set out in accordance with the problems outlined

above.

?O0PHIOMORPHA

Description

Large sub-cylindrical sand-filled vertical burrows with dark pyritic linings

occur as endichnial traces in several sandstone horizons. Differential weathering

of the lining temporarily exposes the burrow-fill in bas-relief, but eventually

both burrow lining and fill are excavated leaving the bedding surface crowded

with perpendicular holes (Fig. 3). These weathered-out burrows seldom exceed

4 cm in diameter and penetrate little more than 12 cm into the substrate. In a

few examples relict mammilation of the linings can be discerned (Shone 1983:

241-242). No clearly defined branching of the tubes has been observed, but

strata adjacent to the vertically burrowed horizons contain numerous branching

sub-horizontal burrows of similar diameter.

TRACE FOSSILS OF THE SARDINIA BAY FORMATION 18)

Fig. 3. Large-diameter vertical burrows. Differential weathering of the less resistant burrow-

fill and lining results in a bedding surface crowded with holes perpendicular to bedding.

Scale = 5 cm.

Interpretation

These vertical burrows are broadly similar to Ophiomorpha but lack the

typical dichotomous branching of that ichnogenus. If the openings were more

distinctly funnel-shaped and the plugged tubes curved in vertical section, Mono-

craterion might be a more plausible identification. The latter has a stratigraphic

Fig. 4. Horizontal burrow with a dark pyrite-rich lining. Differential weathering of lining and

fill results in excavation of the burrow (left).

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

time range from Cambrian to Jurassic (Hantzschel 1975: W84), whereas Ophio-

morpha (Lower Permian—Recent) is typically associated with strata younger

than the Sardinia Bay Formation (Hantzschel 1975: W85). Ophiomorpha-like

structures in modern sediments are produced by callianassid decapods (Weimer

& Hoyt 1964).

THALASSINOIDES SP.

Description

Smooth-walled sand-filled sub-horizontal to inclined burrows, with a lining

of dark pyritic sand and circular to flattened-oval in cross-section (Fig. 4), occur in

close stratigraphic proximity to beds permeated by large vertical Ophiomorpha-

type burrows. These burrows often show repeated branching (Fig. 5). Burrow

diameters range from 2 to 10 cm and individual shaft sections are up to 50 cm

long. Inclined shafts may run parallel to foreset surfaces in the cross-bedded

sandstone.

Interpretation

The sub-horizontal branching burrow systems described above are closely

similar to Thalassinoides (Triassic-Recent), which are generally considered to

be the dwelling burrows of callianassid decapods (Hantzschel 1975: W117) or

crabs (Radwanski 1977: 233-235).

Fig. 5. Partly weathered burrows showing initial erosion of burrow lining and eventual excava-

tion of fill and lining. Basal surface of a sandstone. Note the branching burrow (arrowed)

which resembles Thalassinoides.

TRACE FOSSILS OF THE SARDINIA BAY FORMATION ILS

DIPLOCRATERION

Description

Spreiten demarcated by pyrite-rich laminae, indicating the former position

of a U-tube, were found in a loose boulder of quartzite (Fig. 6). ‘The exact strati-

graphic position of this fragment has not been fixed but is probably close to the

interval 30-60 m (see Fig. 2). Irregular U-tubes up to 2 cm in diameter, with the

limbs of the tube up to 7 cm apart, occur 48 m from the base of the Sardinia Bay

Formation. Spreiten can be discerned between the limbs of one U-tube (Fig. 7).

Fig. 6. Spreiten demarcating the former positions of U-tubes indicating upward migration of

U-shaped burrows, probably Diplocraterion. Tracing from a loose boulder.

Interpretation

Figure 6 shows a typical example of Diplocraterion in which upward

migration of the U-shaped burrow has occurred in response to sedimentation.

The U-tubes themselves are not represented, possibly because of erosional

truncation following sedimentation (cf. Goldring 1964, in Hantzschel 1975:

W30). The irregular U-tubes shown in Figure 7 are less convincing examples of

the ichnogenus. Diplocraterion is thought to be the dwelling trace of a

suspension-feeding organism living in a high wave energy environment, and is

known from strata of Cambrian to Cretaceous age (Hantszchel 1975: W62).

SKOLITHOS

Description

Millimetre-thin straight or gently-meandering sand-filled tubes, perpendicu-

lar to the bedding, occur in sandstones of the Upper Sardinia Bay Formation

(see Fig. 2). Tube cross-sections are oval to roughly circular. The tubes are gen-

erally crowded to the extent that the internal lamination of the host bed is

virtually destroyed (Fig. 8). Individual tubes appear to penetrate whole beds of

up to 60 cm in thickness.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Photograph (top) and tracing (bottom) showing an irregular U-tube with spreiten in a

bioturbated sandstone.

Interpretation

These are probably Skolithos tubes (Upper Precambrian—Lower Creta-

ceous), usually attributed to the activities of marine worms but of uncertain origin

(Hantzschel 1975: W108). Skolithos tubes are widespread in other Table Moun-

tain Group outcrops (Rust 1967, 1973, 1977; Cocks et al. 1970: 603).

THIN VERMIFORM SAND-FILLED TUBES

Description

Vermiform sand-filled tubes, up to 1 cm in diameter and oval to circular in

cross-section, occur in spaghetti-like masses towards the bases of thin sandstone

beds (Fig. 9) in the Upper Sardinia Bay Formation. Individual tubes tend to

straighten and become sub-vertical upwards.

Interpretation

The vermiform traces described above resemble lower parts of the ichno-

genus Fascifodina, the upper parts of which, including the master-shaft, have

been removed by erosion (cf. Hantzschel 1975: W63, fig. 3). Fascifodina is,

TRACE FOSSILS OF THE SARDINIA BAY FORMATION

Fig. 8. Crowded Skolithos tubes in a quartzitic sandstone that has acquired massive texture

a result of the bioturbation.

Fig. 9. Vermiform sand-filled tubes resembling Fascifodina on the basal surface of a sand-

stone, Upper Sardinia Bay Formation.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

however, more usually identified by the presence of horseshoe-shaped groups of

vermiform epireliefs (Hantzschel 1975). The vermiform burrows from the

Sardinia Bay Formation could also be the result of interfering Chondrites

burrows, but apparently lack the repeated branching characteristic of

Chondrites. Fascifodina has been interpreted variously as the grasping traces of

nautiloid tentacles (Flower 1955) or as a feeding burrow (Osgood 1970). The

association of Skolithos-like traces and these enigmatic vermiform burrows

suggests a similar trace-maker for both.

?CRUZIANA

Description

Figure 10 represents a sandstone sole marking that consists of a series of

raised elongate ridges in a V-shaped pattern. It is clear that the ridges on the

sandstone sole are casts of a series of V-shaped grooves cut in an underlying

muddy horizon, now removed by erosion. The V-angle formed by the ridges is

approximately 90°. The trace as a whole is not markedly bilobate.

Interpretation

The V-shaped pattern of scratch marks deduced from the sole marking

(Fig. 10) is broadly suggestive of the trace fossil Cruziana, but lacks the typical

bilobed morphology of the ichnogenus. It is nevertheless possible that the

scratch marks represent shallow furrowing of a trilobite-like arthropod. Cru-

ziana traces are known from strata of Upper Precambrian to Triassic age

(Hantzschel 1975: W55) and younger (Shone 1978).

HORIZONTAL TRACES ON UPPER BEDDING SURFACES

Description

Rare upper bedding surface exposures 45-50 m above the base of the Sar-

dinia Bay Formation are covered by horizontal sand-filled burrows preserved in

convex epirelief. These traces, shown in Figure 11, are up to 7 mm in diameter

and 10 cm long, branch repeatedly, and cross one another. Raised stellate forms

with burrows radiating from a central hub can also be recognized. Small,

rounded, sand-filled tubes appear to intersect the bedding surface but their verti-

cal extent has not yet been established.

Interpretation

These enigmatic trace fossils resemble Chondrites, their surface appearance

corresponding to the three-dimensional tunnel patterns illustrated by Simpson

(1956: 484, fig. 2) and Hantzschel (1975: WS1, fig. 32 (1c)). The tendency of

horizontal tunnels to cross one another is more typical of Fucusopsis (Upper

Ordovician—Tertiary— Hantzschel 1975: W64) but the dichotomous branching

and vertical tunnel sections weigh heavily in favour of Chondrites as the prob-

able ichnogenus. Chondrites is known to occur in strata from Cambrian to Ter-

TRACE FOSSILS OF THE SARDINIA BAY FORMATION 19

Fig. 10. Photograph (top) and tracing (bottom) of a fibreglass cast of a sandstone sole showing

a series of ridges and grooves arranged in a V-shaped pattern. The general appearance is that

of Cruziana but the trace is not markedly bilobed.

tiary in age, and is generally considered to be the feeding burrow of a sediment-

eating animal (Richter 1927; Seilacher 1955; Osgood 1970).

PLANOLITES

Description

Short, gently-curved, subcylindrical burrows with rounded ends. The burrow

fill is almost indistinguishable from the surrounding muddy substrate, so that the

walls of individual traces are not clearly defined (Fig. 12). These mud-filled barely

discernible tubes occur in the interval 60-73 m (see Fig. 2) in a dark pyritic mud-

rock unit, which weathers to a very light grey colour.

Interpretation

These traces resemble Planolites (Precambrian—Recent), which are believed

to be burrows filled with the alimentary casts of sediment-ingesting animals,

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Photograph (left) and tracing (right) showing thin horizontal sand-filled burrows and

rounded cross-sections of vertical sand-filled tubes on a sandstone bedding surface. These

traces are similar to Chondrites.

probably worms (Nicholson 1873; Hantzschel 1975: W95—W97). The indistinct

walls suggest the host sediment and burrow-fill were identical in composition

and that the burrowing activities may have taken place in a soupy mud (cf.

Ekdale et al. 1984: 219).

ICHNOFACIES MODELS AND DEPOSITIONAL ENVIRONMENT

THE ICHNOFACIES CONCEPT

Trace fossil assemblages are often associated with particular depositional

environments. Seilacher (1967) attempted to correlate trace fossil assemblages

with water depth, but it is now clear that other factors (sediment composition

and texture, temperature, light penetration, salinity, oxygenation, food supply,

hydraulic energy, and sedimentation rates) play a significant role in the distribu-

tion of animals and their traces. Because each specific sedimentary environment

is normally associated with a package of factors that controls the distribution of

animals and the preservation of their traces in that environment, the develop-

ment of ichnofacies models linking trace fossil assemblages with sedimentary

facies (Frey & Pemberton 1984) seems entirely logical. It should be borne in

TRACE FOSSILS OF THE SARDINIA BAY FORMATION Da

Fig. 12. Weathered surface of a mudrock unit showing disseminated pyrite now altered to lim-

onite (dark nodules) and short, gently curved burrows with rounded ends (arrowed), which

resemble Planolites. Note that the burrow fill is almost indistinguishable from the substrate;

the burrow margins are revealed only by the effects of differential weathering.

mind, however, that ichnofacies models are likely to be less reliable in dealing

with ancient sedimentary rocks than is the case in younger strata and sediments,

where the organisms responsible for producing the traces can be more readily

identified and their behaviour more easily correlated with changes in the govern-

ing conditions.

SKOLITHOS ICHNOFACIES

The Skolithos ichnofacies model described by Seilacher (1967) and Frey &

Pemberton (1984: 198-199) is typified by the presence of burrows with rein-

forced wall linings and a predominance of vertical shafts. Characteristic ichno-

genera are Skolithos, Ophiomorpha and Diplocraterion. The wall linings indicate

burrowing in soft sediment. Sedimentary structures associated with this ichno-

facies include near-horizontal lamination, ripple laminae, trough cross-bedding,

and flaser-, wavy- and lenticular-bedding. Biogenic structures are subordinate.

The Skolithos ichnofacies is believed to indicate high current and wave energy

regimes in shallow marine settings, ranging from shallow shelf and shoreface to

estuarine and tidal-flat environments.

Nap ANNALS OF THE SOUTH AFRICAN MUSEUM

CRUZIANA ICHNOFACIES

The Cruziana ichnofacies model (Seilacher 1967; Frey & Pemberton 1984:

199-200; Ekdale et al. 1984: 194-196) is typified by unconsolidated substrate at

the time of bioturbation, and is indicative of moderate to low energy levels in

shallow marine environments. Cruziana, Thalassinoides, and Planolites are

characteristic trace fossils. Horizontal shafts predominate over vertical shafts,

and the substrate tends to be totally bioturbated with few relict sedimentary

structures.

TRACE FOSSIL ASSOCIATIONS IN THE SARDINIA BAY FORMATION

Three trace fossil associations can be distinguished in the Sardinia Bay

Formation.

Association I

This trace fossil association, which includes traces resembling Ophiomor-

pha, Thalassinoides, Diplocraterion, Chondrites, and possibly Cruziana, occurs

in the interval 41-60 m (see Fig. 2). Sedimentary structures predominate over

biogenic ones in this interval and are indicative of relatively high wave and

current energy at the time of deposition. Both vertical and horizontal traces are

present. Vertical components appear to be more abundant (see Fig. 3), and

many of the burrows are lined with a dark pyritic sand. The pyrite is evidently

diagenetic in origin (Shone 1983: 115) and could have formed in response to

local reducing conditions associated with originally organic-rich linings.

Association 2

A second trace fossil association, consisting of Planolites-like traces in a

dark pyritic mudrock, occurs in the interval 60-73 m (see Fig. 2). The apparent

paucity of burrows and low faunal diversity suggest a fairly inhospitable environ-

ment. The presence of disseminated pyrite in this interval could indicate anoxic

and possibly hypersaline conditions (Shone 1983), both of which might inhibit

colonization by sediment-dwelling organisms.

Association 3

This association, which occurs in the Upper Sardinia Bay Formation in the

interval 125-155 m (see Fig. 2), consists of intensely bioturbated muddy sand-

stones and siltstones in which traces resembling Skolithos and Fascifodina occur.

There are few relict sedimentary structures. Burrowing appears to have taken

place in unconsolidated substrate.

Discussion

None of the Sardinia Bay Formation trace fossil associations fit comfortably

into the models proposed by Frey & Pemberton (1984). The first Sardinia Bay

association (41-60 m) contains elements of both the Skolithos and Cruziana

TRACE FOSSILS OF THE SARDINIA BAY FORMATION 23

ichnofacies models, fitting best into the Skolithos ichnofacies despite the absence

of Skolithos burrows and the presence of Cruziana-like traces. The third Sar-

dinia Bay association (125-155 m) fits best into the Cruziana ichnofacies model

but lacks Cruziana itself and is characterized by Skolithos! Both Skolithos and

Cruziana ichnofacies models indicate a shallow marine origin for the Sardinia

Bay strata but details of the ichnofacies models are poorly matched with details

of the Sardinia Bay association.

The difference extant between the ideal ichnofacies models and trace fossil

associations in the Sardinia Bay Formation could reflect broad aspects of the

depositional environment. Detailed facies descriptions, interpretations, and palaeo-

geographic reconstructions have been described elsewhere (Shone 1983, 1987).

The purpose of this paper is to document and describe the Sardinia Bay trace fossils.

The proposed depositional model is, however, sufficiently unconventional to

warrant brief consideration. The Sardinia Bay Formation is believed to have

been deposited in a transgressing epeiric sea. Epeiric seas formed by flooding of

intracontinental areas may have been quite common in the past. Their extreme

shallowness over large areas may have led to frictional damping of tides and cur-

rents in inner epeiric shelf reaches (Shaw 1964; Irwin 1965; Hallam 1981), reducing

circulation to the point where anoxic hypersaline conditions prevailed. The inner-

most shelf areas might therefore be characterized at best by a restricted biota, at

worst by no biota at all. Mudrock units in the Sardinia Bay Formation containing

disseminated pyrite, a possible indicator of anoxic and hypersaline conditions (cf.

Berner et al. 1979), do in fact include very few identifiable trace fossils. What is

puzzling, however, is the repeated superposition of trace-free mudrock units

with bioturbated sandstones (see Fig. 2). Such rapid vertical facies changes are

probably typical of epeiric sea deposits. Smail fluctuations in relative sea-level

might accomplish large lateral shifts in the boundaries of depositional environ-

ments, because of the extremely low bottom gradients involved. The repetition

of certain facies (see Fig. 2), in particular the innermost shelf mud facies, could

reflect small changes in relative sea-level.

The ichnofacies models proposed by Frey & Pemberton (1984) are based in

part on more conventional sedimentation models.

The sedimentary structures in trace fossil association 1 indicate moderate

wave and current activity, which could have been responsible for restricting both

the number of sediment-dwelling organisms and the degree of bioturbation. The

ichnofaunal diversity was not so restricted, however, which suggests that the

sediments hosted a variety of animals during fair-weather periods. Association 1

could be representative of the zone of wave-break and unfettered tidal current

flow in an ancient epeiric sea.

Association 2 (mudrock with barely discernible Planolites traces, dissemi-

nated pyrite) appears to have been deposited in a zone of restricted circulation.

Increasingly anoxic conditions in such quiet reaches of an epeiric sea would

eventually prohibit colonization by sediment dwellers and eliminate the possi-

bility of soft-substrate burrowing.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Association 3 appears to have developed in an intermediate zone, where

damping of tidal currents and open sea waves was only partial. Anoxic con-

ditions could have arisen in this zone as a result of reduced circulation, and this

could be reflected in the restricted ichnofauna (Skolithos and ? Fascifodina). The

extensive bioturbation that occurred probably indicates long-term burrowing in

a low-energy environment in which ichnocoenoses were seldom displaced. Bed-

ding traces formed here would have a very low preservation potential because

of the high degree of bioturbation. This places Skolithos out of the high-energy

clean-sand facies with which it is normally associated, but in other respects

provides a far better explanation for the Sardinia Bay trace fossil associations.

ACKNOWLEDGEMENTS

I wish to thank the University of Port Elizabeth for a research grant that

helped to defray incidental costs. Two anonymous referees suggested a number

of improvements now incorporated in the manuscript, which was typed by

Shelagh Matthews.

REFERENCES

ALLEN, J. R. L. 1980. Sand waves: a model of origin and internal structures. Sedimentary

Geology 26 (1-3): 281-328.

BERNER, R. A., BALDWIN, T. & HOLpDREN, G. R. 1979. Authigenic iron sulfides as palaeo-

salinity indicators. Journal of Sedimentary Petrology 49 (4): 1345-1350.

Cocks, L. R. M., Brunton, C. H. C., ROwELL, A. J. & Rust, I. C. 1970. The first Lower

Palaeozoic fauna proved from South Africa. Quarterly Journal of the Geological Society of

London 125 (4): 583-603.

De Swarpt, A. M. J. & RowseLL, D. M. 1974. Note on the relationship between diagenesis

and deformation in the Cape Fold Belt. Transactions of the Geological Society of South

Africa 77 (3): 239-245.

EKDALE, A. A., BROMLEY, R. G. & PEMBERTON, S. G. 1984. Ichnology: trace fossils in sedi-

mentology and stratigraphy. SEPM Short Course 15. Tulsa, Oklahoma: Society of Econ-

omic Palaeontologists and Mineralogists.

FLower, R. H. 1955. Trails and tentacular impressions of orthoconic cephalopods. Journal of

Paleontology 29: 857-867.

Frey, R. W. & PEMBERTON, S. G. 1984. Trace fossil facies models. Jn: WALKER, R. G. ed.

Facies models: 189-207. 2nd ed. St. Johns, Newfoundland: Geoscience Canada. (Reprint

Series 1.)

GoLpRING, R. 1964. Trace fossils and the sedimentary surface in shallow-water marine sedi-

ments. In: VAN STRAATEN, L. M. J. U. ed. Deltaic and shallow marine deposits. Develop-

ments in Sedimentology 1: 136-143.

HALLAM, A. 1981. Facies interpretation and the stratigraphic record. San Francisco: W. H.

Freeman & Co.

HANTZSCHEL, W. 1975. Trace fossils and problematica. 2nd ed. In: TEICHERT, C. ed. Treatise

on invertebrate paleontology, Part W, Miscellanea, Supplement 1. Boulder, Colorado:

Geological Society of America; Lawrence, Kansas: University of Kansas Press.

Irwin, M. L. 1965. General theory of epeiric clear water sedimentation. Bulletin of the

American Association of Petroleum Geologists 49 (4): 445-459.

NicHoitson, H. A. 1873. Contributions to the study of the errant annelides of the older

Palaeozoic rocks. Proceedings of the Royal Society 21: 288-290.

Oscoop, R. G. 1970. Trace fossils of the Cincinnati area. Palaeontologia Americana 6 (41):

281-444.

TRACE FOSSILS OF THE SARDINIA BAY FORMATION D5

RADWANSKI, A. 1977. Present-day types of trace in the Neogene Sequence; their problems of

nomenclature and preservation. Jn: Crimes, T. P. & Harper, J. C. eds. Trace fossils 2:

227-263. Liverpool: Seel House Press.

RicuTerR, R. 1927. Die fossilen Fahrten und Bauten der Wiirmer, ein Uberblick iiber ihre

biologischen Grundformen und dehren geologische Bedeuting. Paldontologische Zeit-

schrift 9: 193-240.

Rust, I. C. 1967. On the sedimentation of the Table Mountain Group in the Western Cape

Province. Unpublished D.Sc. thesis, University of Stellenbosch.

Rust, I. C. 1973. The evolution of the Paleozoic Cape Basin, southern margin of Africa. In:

Nairn, A. E. M. & STEHLI, F. G. eds. The ocean basins and margins. 1. The South Atlan-

tic: 247-276. New York: Plenum Press.

Rust, I. C. 1977. Evidence of shallow marine and tidal sedimentation in the Ordovician

Graafwater Formation, Cape Province, South Africa. Sedimentary Geology 18 (1-3):

123-133.

SEILACHER, A. 1955. Spuren und Fazies in Unterkambrium. /n: SCHINDEWOLF, O. H. & SEIL-

ACHER, A. (ed.) Beitrage zur Kenntnis des Kambriums in der Salt Range (Pakistan). Aka-

demie fiir Wissenschaften und der Literatur, Mainz. Abhandlungen der Mathematisch-

Naturwissenschaftlichen Klasse—10: 11-143.

SEILACHER, A. 1967. Bathymetry of trace fossils. Marine Geology 5 (5-6): 413-428.

SHaw, A. B. 1964. Time in stratigraphy. New York: McGraw-Hill.

SHONE, R. W. 1978. Giant Cruziana from the Beaufort Group. Transactions of the Geological

Society of South Africa 81 (3): 327-329.

SHONE, R. W. 1979. Sardinia Bay excursion. Jn: Rust, I. C. ed. Excursions guide book 1-4,

Geokongres ’79: 171-180. Geological Society of South Africa.

SHONE, R. W. 1983. The geology of the Sardinia Bay Formation. Unpublished Ph.D. thesis,

University of Port Elizabeth.

SHONE, R. W. 1987. A resedimented conglomerate and shallow-water turbidite from the Sar-

dinia Bay Formation, Lower Table Mountain Group. South African Journal of Geology 90

(1): 86-93.

Smmpson, S. 1956. On the trace-fossil Chondrites. Quarterly Journal of the Geological Society

of London 112 (4): 475-499.

TANKARD, A. J., Jackson, M. P. A., Eriksson, K. A., Hosppay, D. K., HUNTER, D. R. &

Minter, W. E. L. 1982. Crustal evolution of southern Africa. 3.8 billion years of earth

history. New York: Springer-Verlag.

WEIMER, R. J. & Hoyt, J. H. 1964. Burrows of Callianassa major Say, indicators of littoral

and shallow neritic environments. Journal of Paleontology 38 (4): 761-767.

j ig

Ht ii

a D rh Pal |

ra foo ta ;

‘ 1 [ ’ i

my at

| Aa ee i s

f , ‘

. if : hy

| j

| 5

. ‘fe i :

4 i ’ H

a ent

; ie he ay ra

\ i t Wi ,

\ y yl v

i] i 1 0 wy

a i 1 i aor he

4 ¢ 4 hin

| i

| ‘

i. ih

| : ;

| A i i 7

fee j

, j 0 Nae i :

I ‘ i}

}

} y

be e ' : 4 i

aS } Ro

1 f

: E

ES r

f Ue, 7 7

“heh

min :

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

ete.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. ‘. . . the Figure depicting C. namacolus ...: *. . . in C. namacolus (Fig. 10). . .’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by

initials or full names

e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviaticn to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

RUSSELL W. SHONE

TRACE FOSSILS OF THE ?7EARLY

ORDOVICIAN SARDINIA BAY FORMATION,

TABLE MOUNTAIN GROUP

—e—— ae

x VOLUME 101 PART 3 AUGUST 1991 ISSN 0303-2515

; EQTHSONIAN

LIBRARIES

OF THE SOUTH AFRICAN

0 aS BUM

CAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.

2. LAYOUT should be as follows:

(a) Centred masthead to consist of

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes . . .’

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 +#£4Band

August 1991 Augustus

Part 3 Deel

A NEW SPECIES OF CALANUS

(COPEPODA, CALANOIDA)

FROM SOUTH AFRICAN WATERS

A. H. B. DE DECKER, B. Z. KACZMARUK

G. MARSKA

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

il, HEB, 5-9), HID, 2S, 8, tap.) SUES, 5, 7-9).

GG, tsa); WOED), ©, COD, 7), IOGED), IMD, 8. 7, C.'.),

MD), 15(C=5), 240, 8), Ti, BGS), 3AG), 23

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 123 7

Printed in South Africa by In Suid-Afrika gedruk deur

ithe Rusticassbress;. Rtye. letds: Die Rustica-pers, Edms., Bpk.,

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D661

A NEW SPECIES OF CALANUS (COPEPODA, CALANOIDA)

FROM SOUTH AFRICAN WATERS*

A. H. B. DE DEcKErR}, B. Z. KACZMARUK & G. MARSKA

Department of Marine Biology, South African Museum, Cape Town

(With 7 figures and 2 tables)

[MS accepted 15 October 1989]

ABSTRACT

A new calanid species belonging to the Calanus helgolandicus species-group is described.

Calanus agulhensis sp. nov. is reported from the Agulhas Bank waters and its seasonal

distribution is briefly presented.

CONTENTS

PAGE

ECHEROOCREE CERO TE oe ee eee gg tein eee ee Ae ea Ue TA fin 8 ee 27

Matera ban GuMmethOGS so. 3 2 sc pas reise anne hee a en eiaee 2 oe 28

SEE DX CHINOILY aire 0 ete, um Pra aa oR NE Chace gears, Ue, po

PFISERAO ELIOT Aetna spline nee eee Race Ny BUS a yee 39

SS TRERTETI BVsfee fo octets os eh ee alae cae ore 5 ch aCH Neue eid ar 42

PRERMOWLECOCIECINGS 2p cveiies aaah hide ee eh svn un oe Ree ae, Rania 42

RGHOTERISES, Beth eB ha) nah aia Gases Oe RUE eee eke eae ae 42

INTRODUCTION

The major part of research on the taxonomy of the family Calanidae in the

Southern Hemisphere has been performed by Brodskii (e.g. 1959, 1961, 1964,

1965, 1972, 1976). He divided (1959) the superspecies Calanus finmarchicus s.1.

of Yashnov (1958) into a number of taxa of specific and lower rank. Small

morphological differences, together with differences in geographical distribu-

tion, formed the basis for his classification within the genus Calanus s.s. The

species and varieties of Calanus that Brodskii recognized in the Southern

Hemisphere are C. australis Brodskii, 1959, C. chilensis Brodskii, 1959, as well

as C. australis atlanticus Brodskii, 1959, and C. australis pacificus Brodskii,

1959. Although Brodskii’s work undoubtedly increased our knowledge on

calanids, his classification (e.g. Brodskii 1976) raised controversy (see Bradford

& Jillett 1974; Bradford 1988).

Calanus finmarchicus (Gunnerus, 1765) was first reported from South

African waters by Cleve (1904: 185), who described the species as ‘common

south of the Cape Colony, rare east and west’. This was confirmed by De

* This paper is partially based on an earlier manuscript by Dr A. H. B. De Decker.

+ Dr De Decker died on 3 January 1986.

WJ)

Ann. S. Afr. Mus. 101 (3), 1991: 27-44, 7 figs, 2 tables.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Decker (1964: 18) who found that ‘a fairly numerous colony is consistently

present over the Agulhas Bank’ and that ‘off the West Coast and off Natal it

occurs only at certain times and in small numbers’.

Seno et al. (1963) reported 4 915 individuals of C. finmarchicus from the

northernmost station (37°43’S 20°05’E) occupied during an Africa—Antarctica

transect of the 1956-57 ‘Umitaka-Maru’ Expedition. Although beyond the

continental shelf, this station is located in an area influenced by the Agulhas

Current system. The area directly south of South Africa is known for the forma-

tion of eddies and rings as a consequence of the Agulhas Current retroflexion

and considerable latitudinal variation in the position of the Subtropical Con-

vergence (e.g. Bang 1970; Lutjeharms 1985). Therefore, transport of Agulhas

Bank water and its plankton away from the shelf is not unexpected.

Carter (1978) found the species to be rare off Durban but occurring fairly

frequently 150 km further south, off Port Edward.

The northernmost record of Calanus populations off South Africa is at 30°S

(Port Nolloth) in the Atlantic Ocean and at 28°S (Cape St Lucia) in the Indian

Ocean (unpublished data).

Various authors, e.g. Brodskii (1959), Yashnov (1972), and Fleminger &

Hulsemann (1977), have presented convincing evidence that C. finmarchicus

and C. helgolandicus (Claus, 1863) are indeed separate species and that both

species are restricted to the North Atlantic.

De Decker (1973) first considered the South African specimens to be

C. finmarchicus s.1., but subsequently (De Decker 1984: 320) used the name

C. finmarchicus s.\. in referring to the South African form to ‘designate a form

of uncertain taxonomic status showing morphological resemblances with both

Calanus australis (Brodskii, 1959) and Calanus pacificus pacificus (Brodskii,

1959)’. He suggested that it might represent a separate taxon. Hutchings (1985)

reported the species off Cape Town under the name C. finmarchicus v. australis.

De Decker (pers. comm. in Bradford 1979) indicated that the South

African form could not be identified with any described species of Calanus.

Careful comparison with material from New Zealand, Australia and California

have revealed differences of the same order as those used in Brodskii’s system.

These differences, together with the geographical isolation of the population,

permit its recognition as a distinct species.

MATERIAL AND METHODS

Material for the study was obtained from monthly routine surveys of the

then Division of Sea Fisheries around the Cape, during the period July

1963—June 1965. The sampling stations were located at +20-mile intervals

(Fig. 1). The three eastern transects, viz. lines 72, 76 and 80, covered the

western part of the Agulhas Bank. Regular plankton sampling was done only

during daylight. Plankton samples were taken with a Nansen vertical closing net

(mouth opening 70 cm) of 0,200-mm mesh size and preserved in 4 per cent

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 29

routine stations

type locality

e—e selected transects

o—__9_____+>——_-®

20 Elands)Bay A

33° iar A .

‘ e

e c e

e e

0 e

e e

: Plettenberg Bay

3 3 2 Cape Infanta St.A625

52 1

e e a rs : ;

35° e e e i : 2 N a8

4 e \ i

e e e . ; ‘ A 80

ane | 76% B

72a) A

« O z 1 Ss

be t u 5s A

e ' H

e 4 U L

e A G

, .)

18° 20° oP oa?

Fig. 1. Routine monthly stations and the type locality of Calanus agulhensis sp. nov.

formaldehyde solution. The net was built according to specifications provided in

the Discovery Reports (Kemp ef al. 1929). At the vast majority of stations a

single haul from 100 m depth to the surface was made.

A total of 1 840 samples were checked for the presence of the species.

Adult females, adult males and copepodites V were counted separately. How-

ever, total numbers of adults are presented here, as numbers of males were

insignificant at all stations. For the purpose of the present study, data from only

three transects, viz. lines 20, 52 and 80, are presented.

TAXONOMY

The extensive material studied by Brodskii (1964, 1967) from the Southern

Hemisphere was mostly collected by the Soviet Antarctic Expedition of

1955-1958. However, the only region sampled in the Atlantic Ocean was off the

South American coast (Brodskii 1964). Nevertheless, he implied distribution of

C. australis in South African waters, although he never mentioned having

specimens from this region.

In addition to Brodskii’s studies, occurrences of copepods belonging to the

Calanus helgolandicus species-group have been reported elsewhere in the

Southern Hemisphere.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Farran (1929) found both sexes of a species recorded as C. finmarchicus in

material from New Zealand waters, and remarked on body size and the number

of teeth on the lamella of the first basipod of the fifth legs, without taking the

comparison any further. However, he conditionally recognized the southern

form as a separate species, pending the outcome of the C. finmarchicus versus

C. helgolandicus controversy. He speculated on the probability of a complete

separation of the two forms.

After examination of a number of females of ‘C. finmarchicus’ taken south

of Tasmania, Vervoort (1957) considered the differences between these and the

form from the Northern Hemisphere to be insufficient as a basis for the erection

of a new species.

Wiborg (1964) identified a few copepodites and adult females from Tristan

da Cunha as C. australis. He was unable to assign his specimens to either of

Brodskii’s (1959) two varieties of C. australis—cC. australis var. atlanticus and

C. australis var. pacificus, and remarked that Brodskii’s taxa ‘C. australis’,

‘C. pacificus’ and ‘C. chilensis’ were based on ‘very minute’ differences.

Deevey (1966) experienced difficulties when trying to assign New Zealand

specimens to Brodskii’s (1961) two varieties of C. australis. The females fitted

the description of C. australis var. atlanticus, whereas the males showed charac-

teristics of both C. a. var. atlanticus and C. a. var. pacificus.

Grice & Hulsemann (1967) found a single female specimen about mid-way

between South Africa and Australia (35°09’S 69°59’'E), which they assigned to

C. australis var. atlanticus Brodskii, 1959.

Vidal (1968) reported C. australis s.1. from the tropical waters off northern

Chile. He mentioned differences between his specimens and Brodskii’s (1959)

description of that species that seem to be of the same order as those on which

Brodskii recognized separate species.

Bradford (1979) could find no distinguishing characters between the females

of C. australis and C. chilensis.

Description of the new species, Calanus agulhensis, and comparative

remarks on affinities to closely related species follow.

Family Calanidae

Calanus agulhensis sp. nov.

Figs 2-7

Calanus finmarchicus (non Gunnerus) Cleve, 1904: 185. Send et al., 1963: 58, table 2. De

Decker, 1964: 7, 14, 18, 26, 31.

Calanus finmarchicus sensu lato De Decker, 1973: 213, 218; 1984: 315, 320, 321, 322, 323, 331.

Carter, 1978: 35, 45, 48, 52, 53, 69, 71.

Calanus finmarchicus var. australis Hutchings, 1985: 12, 29, 31, 37, 38, figs 9, 15, table 5.

Material

Type material from the Agulhas Bank is deposited at the South African

Museum, Cape Town.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 31

Holotype. SAM-—MC00001, adult female, dissected and mounted on

11 microscope slides. R.V. Africana II, Station A625, 34°12,5’S 23°24,5’E,

8 April 1960, depth 94—0 m.

Allotype. SAM—MC00002, adult male, dissected and mounted on 11 micro-

scope slides. Station data as for holotype.

Paratypes. SAM—MC00003, 148 adult females, 19 adult males, 184 juv-

eniles in 4 per cent formaldehyde solution in water with glycerine. Station data

as for holotype.

Description

Female (Figs 2—4). Total length 2,45-2,95 mm, mean 2,73 mm, n=50.

Ratio of prosome length to maximum width 3,00-—3,21, mean 3,06, n=9. Ratio

of prosome length to total length (including furca) 0,78—0,83, mean 0,80, n=9.

Head produced anterodorsally, ventrolateral margin slightly divergent in

the oral region. In dorsal aspect, forehead with sides anteriorly converging

towards a smoothly rounded apex; in lateral aspect evenly rounded, bulging in

front of the attachment of the rostral filaments. Posterior margin of head carrying

a small mid-dorsal knob, which in lateral aspect appears skewed posteriorly.

All five pedigerous segments clearly separated, with straight sides. Greatest

width of prosome near the suture between pedigerous segments 1 and 2.

Posterior projections of last pedigerous segment narrowly rounded in lateral

view; in dorsal view their dorsal and ventral margins converging in perpen-

dicular or slightly obtuse-angled directions to meet in a rounded tip, extending

over not more than one-third of the genital segment.

Genital segment in dorsal view nearly as broad as long, with moderately

bulging sides, its maximum width anterior of the middle; in lateral view with

pronounced ventral bulge culminating at the anterior third of the segment,

carrying the genital pore on its summit. Ventral surface of genital segment

posterior to genital pore straight or slightly convex in lateral view. Antral cover

with short straight sides joining distally in a semicircle. Spermathecae oblong to

bean-shaped; size variable; often distorted or dislodged. Spermatophore as

illustrated in Figure 4H.

Furcal rami about twice as long as broad, their inner margin with a row of

long, delicate hairs, which are often partly or wholly missing in preserved

specimens.

First antennae reaching as far as the tips of the furcal rami or no more than

one segment beyond. Second antennae, oral appendages and legs 1 to 4 of the

usual shape for the genus, although second maxillae appear stouter than usual,

with sides of the lobes touching over nearly their whole length.

Serrate lamina on inner margin of basipod 1 of leg 5 with 14-18 teeth

(mean 16, n=13). Some individual variation exists in the curvature of the

lamina; its median part usually bends on to the posterior surface of the joint and

turns backwards, so that its teeth become oriented at right angles to the

proximal and distal ones. Proximal teeth bent down in the shape of a hawk’s

oP) ANNALS OF THE SOUTH AFRICAN MUSEUM

C-D

0,2mm

Fig. 2. Calanus agulhensis, 9. A. Body, dorsal view. B. Body, lateral view. C. 2nd antenna. D. Mandible.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA

: Mg,

Kay

Fig. 3. Calanus agulhensis, 2. A. Mandible gnathobasis. B. 2nd maxilla. C. 1st maxilla.

D. Maxilliped. E. Ist leg. FF. Seminal receptacle, right lateral. G. Seminal receptacle,

ventral. H. Genital segment with spermatophore, right lateral.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Calanus agulhensis, 2. A. 2nd leg. B. 3rd leg. C. 4th leg.

D. 5th leg. E. Basipodal segment, inner margin, 5th leg.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 35

beak, ending in a sharp or blunted point; distal ones becoming progressively

straighter, tending towards the shape of an equilateral triangle. There are no

gaps between the bases of the teeth, which do not differ noticeably in length. In

a few cases the teeth were oval in shape instead of curved or triangular.

Basipod 2 of leg 5 with moderately convex inner margin. Ratio of endopod

length of leg 5 to the length of exopod 0,60—0,69 (mean 0,66, n= 13). Endopod

segment 1 with exterodistal corner produced into a sharp point; tip reaches the

suture between exopods segments 1 and 2. Endopod segment 3 reaching as far as,

or slightly beyond, insertion of first inner seta of exopod 3. Endopod segment 3

carrying 5—6 setae: 2 inner, 2 terminal and 1-2 outer ones (of 12 specimens

examined, five had one outer seta, five had two, and two had one on one leg and

two on the other). Great care was taken to ascertain that the absence of a second

seta was not due to damage. The terminal spine of exopod 3 as long as the

segment.

Male (Figs 5—7). Total length 2,74—3,00 mm (mean 2,78 mm, n=7). Ratio

of prosome length to maximum width 2,72. Ratio of prosome length to total

length, including furca: 0,78.

Forehead rounded, strongly protruding anterior to rostral filaments. Mid-

dorsal knob at the posterior margin of head more pronounced than in female; a

further mid-dorsal prominence opposite insertion of the second antennae.

All pedigerous segments clearly separated, but the separation between head

and first pedigerous segment becoming faint along the sides. Posterolateral

projections of the last prosomal segment extending over half the first urosomal

segment; their lateral profile evenly rounded, in dorsal aspect their margins

converging at an acute angle before joining in a blunt tip. Genital segment in

dorsal aspect widening posteriorly. Ratio of length of genital segment to the

length of urosome 0,31-0,32.

First antennae stretching beyond furca by 3-33 segments, segments 1 and 2

fused. Second antennae, oral appendages and legs 1 to 4 as in female, except

maxilliped, which carries on each of the two distal segments the enlarged,

retroflected and profusely plumose seta typical of male calanids.

Leg 5 with serrate lamina of basipod 1 less curved than in female, sometimes

nearly straight and with greater variation in size and shape of teeth, which tend to

be shightly more numerous: 16—20 (mean 18, n = 9); in some specimens they are

separated by narrow gaps. Distal margin of right basipod 2 of leg 5 reaching half-

way along the length of inner margin of left basipod 2.

Exopod of left leg 5 extending beyond furca by its distal segment and about

half segment 2. Segment 2 slightly shorter but considerably narrower than

segment 1, the width/length ratio being 3 to 4 in segment 1, 7 to § in segment 2,

and 3 to 3 in segment 3. Segment 3 with a small lateral spine and short and long

terminal seta, the latter very thin and easily lost in preserved specimens.

Endopod of left leg 5 excluding terminal setae, not extending beyond segment 1

of exopod; setation: 1, 1, 6.

ANNALS OF THE SOUTH AFRICAN MUSEUM

A-C

1mm

0,2mm

Fig. 5. Calanus agulhensis, O&. A. Body, dorsal view. B. Body, lateral view.

C. ist antenna. D. 1st antenna, segment 24. E. 1st antenna, segment 25.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 37

0,2mm A-E al

0,1mm F c

Fig. 6. Calanus agulhensis, 0. A. 2nd antenna. B. Mandible gnathobasis. C. Mandible.

D. 1st maxilla. E. Maxilliped. F. 2nd maxilla.

0,2mm

ANNALS OF THE SOUTH AFRICAN MUSEUM

C. 3rd leg. D. 4th leg. E. Right Sth leg. F. Left Sth leg.

Fig. 7. Calanus agulhensis, '. A. 1st leg. B. 2nd leg.

G. Basipodal segment, inner margin, Sth leg.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 39

Exopod of right leg 5, excluding terminal spine, extending only slightly

beyond segment 1 of left exopod. Endopod of right leg 5 extending to 4 or 3 of

the length of segment 3 of right exopod; setation: 1, 1, 6.

Remarks

Calanus agulhensis appears closely related to both C. australis and C. pacifi-

cus, with greater similarity to latter. With respect to body length, C. agulhensis is

among the smaller forms in the C. helgolandicus species-group, the average size

of which scarcely exceeds 3,00 mm in either sex. Both C. australis var. atlanticus

and C. pacificus var. pacificus Brodsku, 1965, belong to this group (Brodskii

1959).

The mean length of the males exceeds that of females. Brodskii (1961)

found this condition only in C. australis var. pacificus (females 2,95 mm and

males 3,16 mm) and C-. australis var. atlanticus (females 2,94 mm and males

3,05 mm). Deevey’s (1966) measurements of C, australis from New Zealand

waters confirmed this finding: mean length of females was 2,89 mm (range

2,40-3,50 mm), and of males 3,02 mm (range 2,42—3,35 mm).

Calanus agulhensis differs from C. australis in the following: in the female

the first antennae are shorter; the curvature of the denticulate laminae on leg 5

is more pronounced; the exterodistal projection of endopod segment 1 of leg 5

reaches the suture between segments 1 and 2 of the exopod; segment 3 of the

exopod reaches or exceeds the insertion point of the proximal inner seta of

segment 3 of the exopod; the terminal seta of the exopod is not longer than

segment 3. In leg 5 of the male, the endopod does not reach beyond segment 1

of the left exopod, the right exopod reaches only slightly beyond segment 1 of

the left exopod.

The above-mentioned characters in which females of C. agulhensis differ

from those of female of C. australis are, however, identical in C. agulhensis

and females of C. pacificus. The similarity of the fifth leg of the male of

C. agulhensis to that of C. pacificus is striking, except for the presence of a

denticulate lamina on basipod 1.

DISTRIBUTION

The hydrology of the studied area has been described in detail by Orren

(1966), Bang (1970, 1973), Harris & Van Foreest (1978), Lutjeharms (1981),

and Swart & Largier (1987), to mention but a few. Water temperatures are

shown in Table 1.

The quantitative data confirmed previous records of the species and showed

that the Agulhas Bank waters are indeed the centre of distribution of the species

(Table 2); the number of animals decreased westwards. West of the Agulhas

Bank, the species was found irregularly throughout the year and in very low

numbers, and becoming rare at the two most offshore stations. This offshore

reduction in numbers was not observed over the Agulhas Bank, where the

species was evenly distributed.

ANNALS OF THE SOUTH AFRICAN MUSEUM

man

*sJ29SUBI] Pd}o9]9S Suoye suoijeys ye [ney od (v7) sy[npe pue (A) A Ssoypododoo Jo Joquinyy

T aTaViL

maN

mao Ot

“9peul [Ney OU sojedIpur — ,,

y s-00

y 6-00

Ve eth

A ee

yO Saas

x €-¢s

Wececs

Res

X $708

Pte

eeace

Ke 08

a3B1S “WIS

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA

TABLE 2

Water temperature (°C) at the time of plankton sampling along transect 80.

Depth Station no.

pont (m) 80-1 80-2 80-3 80—4 80-5

6 July 1963 0 15535 15,42 16,68 16,29 16,42

50 15523 14,10 15522 = 15,84

3 Aug. 1963 0 14,33 14,36 14,54 14,67 III

50 14,33 14,34 14,54 = 1557/9

7 Sept. 1963 0 14,99 14,79 16,21 16,21 16,33

50 14,14 14,19 14,65 = 16,21

4 Oct. 1963 0 16,06 7M) 7 3)1/ 16,98 16,64

50 ISS) 16,46 16,60 = 16,10

6 Nov. 1963 0 18,53 18,71 19,07 - 19,26

50 Oi 14,70 13,06 = 22

7 Dec. 1963 0 20,62 20,67 20,47 20,75 19,95

50 14,46 279 13}723) = 12,24

14 Jan. 1964 0 21,82 Jill) 21,84 22,09 228

50 12,60 12,89 WS = 15,34

8 Feb. 1964 0 21,96 22S 21,74 20,94 20,80

50 W237) 11,94 11,94 = 15599

24 Mar. 1964 0 21,60 21,70 21,40 21,20 =

9 Apr.1964 0 20,51 19,27 18,95 19,20 18,77

50 12,67 10,91 10,48 = 10,51

6 May 1964 0 18,66 17,97 17,91 18,03 18,01

50 11,85 10,94 i = 11,23

6June 1964 0 16,78 16,86 16,80 16,80 17,98

50 12,58 12,74 16,78 = 13,61

8July1964 0 14,83 14,99 15,29 16,30 16,10

50 10,81 14,99 15,18 = 15,50

9 Aug. 1964 0 14,90 15,58 15,71 16,24 16,54

50 14,61 15,42 15,19 = 16,43

5 Sept.1964 0 14,46 14,69 15,05 15,68 15,93

50 14,44 14,68 14,96 - --

8 Oct. 1964 0 16,49 16,77 16,71 18,25 17,53

50 15,39 15,64 15,32 = 15,73

4Nov. 1964 0 17,49 17,66 Gis 17,69 17,80

50 16,99 17,09 16,82 = 16,60

6Dec.1964 0 19,15 19,60 19,07 18,91 18,81

50 15,86 17,49 14,03 = 15,59

6 Jan. 1965 0 20,79 21,23 21,47 21,53 21,47

50 ea 18,15 16,83 = 18,70

3 Feb. 1965 0 21,53 21,65 DST 22,50 22,43

50 13,57 16,41 13,04 = 18,93

3Mar.1965 0 21,34 20,58 20,68 20,68 20,03

50 21,26 11,15 17,87 - 15,99

3 Apr.1965 0 21,15 20,87 20,88 21,35 21,58

50 12,48 20,32 13,44 ae 19,25

8 May 1965 0 18,94 18,60 18,42 18,04 18,35

50 18,49 lea 18,13 - 18,08

5June 1965 0 16,72 16,64 16,75 17,54 19,78

50 16,72 16,63 16,13 = 17,40

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

The seasonal occurrence of the species could only be investigated on stations

of transect 80, as the species was rare west of the Agulhas Bank. On this transect

a period of peak abundance was observed from August to November. At most

offshore stations moderate abundances were also observed in December. The

period of peak abundance applies to both the adults and copepodites V,

although, during these months, differences were found in the optimum

temperature ranges for adult and copepodite V stages.

During these months, adults were most abundant in water temperatures

ranging from 14,44°C to 19,00°C at 0 m and from 13,06°C to 16,03°C at 50 m

depth; corresponding temperatures for copepodites V were 14,9°C—22,18°C at

Om and 10,51°C-16,86°C at 50 m depth. It appears that copepodites V were

tolerant of wider temperature ranges than the adults.

SUMMARY

An isolated population of new Calanus species occurs over the Agulhas

Bank. Calanus agulhensis sp. nov. belongs to the C. helgolandicus species-

group. The species is closely related to the other calanid species from the

Southern Hemisphere, viz. C. australis. Calanus agulhensis shows a close

resemblance to C. pacificus, which is only found in the Northern Hemisphere.

ACKNOWLEDGEMENTS

The authors would like to thank Dr J. M. Bradford of the New Zealand

Division of Marine and Freshwater Science, Department of Scientific and

Industrial Research, Wellington, for making specimens of Calanus species from

New Zealand available to the late Dr De Decker. We are also indebted to

Dr K. Hulsemann of Biologische Anstalt Helgoland, Hamburg, Federal

Republic of Germany, for reviewing the manuscript and helpful and valuable

remarks.

REFERENCES

Banc, N. D. 1970. Dynamic interpretations of a detailed temperature chart of the Agulhas

Current retrofiexion and fragmentation area. South African Geographical Journal 52:

67-76.

Banc, N. D. 1973. The southern Benguela System: finer oceanic structure and atmospheric

determinants. Unpublished Ph.D. thesis, University of Cape Town.

BrRADForD, J. M. 1979. Zoogeography of some New Zealand neritic pelagic Crustacea and

their close relatives. Information Series. New Zealand Department of Scientific and

Industrial Research 137: 593-612.

BRADFORD, J. M. 1988. Review of the taxonomy of the Calanidae (Copepoda) and the limits

to the genus Calanus. In: BOXHALL, G. A. & SCHMINKE, H. K. eds. Biology of copepods.

Hydrobiologia 167/168: 73-81.

BrapForD, J. M. & JitteTT, J. B. 1974. A revision of generic definition in the Calanidae

(Copepoda, Calanoida). Crustaceana 27 (1): 5-16.

A NEW SPECIES OF CALANUS FROM SOUTH AFRICA 43

Bropskil, K. A. 1959. O filogeneticheskikh otnosheniyakh nekotorykh vidov roda Calanus

(Copepoda) severnogo i yuzhnogo polusharii. [On phylogenetic relations of some Calanus

(Copepoda) species of Northern and Southern hemispheres.|] Zoologicheskij Zhurnal 38

(10): 1537-1553. (In Russian.)

Bropskil, K. A. 1961. Comparison of Calanus species (Copepoda) from the Southern and

Northern hemispheres. Information Series. New Zealand Department of Scientific and

Industrial Research 33: 1-22.

Bropskul, K. A. 1964. Distribution and morphological features of the Antarctic species of

Calanus (Copepoda). Biological reports of the Soviet Antarctic Expedition (1955-1958). 2.

Issledovaniya Fauny Morei 2 (13): 194-256. (Translated by Israel Program for Scientific

Translations, 1966.)

Bropsku, K. A. 1965. Sistematika morskikh planktonnykh organismov i okeanologiya. [The

systematics of marine plankton organisms and oceanology.] Okeanologiya 5 (4): 557-591.

(In Russian.)

Bropsku, K. A. 1967. Distribution and size variablility of species of Calanidae (Copepoda) in

the Southern Hemisphere (from the collections of the Soviet Antarctic Expedition

1955-1958). Biological Reports of the Soviet Antarctic Expedition (1955-1958).

Issledovaniya Fauny Morei 4 (12): 190-219. (Translated by Israel Program for Scientific

Translations, 1968.)

Bropskil, K. A. 1972. Phylogeny of the family Calanidae (Copepoda) on the basis of a

comparative morphological analysis of its characters. Geographical and seasonal variability

of marine plankton. IJssledovaniya Fauny Morei 12 (20): 1-27. (Translated by Israel

Program for Scientific Translations, 1975.)

Bropskil, K. A. 1976. O reviziyak i ‘novykh’ klassifikasiyakh Calanoida (Copepoda). On the

revisions and the ‘new’ classifications of Calanoida (Copepoda). Issledovaniya Fauny Morei

18 (26): 5-10. (Russian with English summary. )

Carter, R. A. 1978. The distribution of calanoid Copepoda in the Agulhas Current system off

Natal, South Africa. Professional Research Series. National Research Institute for

Oceanology. Council for Scientific and Industrial Research 3: 1-165.

CLevE, P. T. 1904. Plankton of the South African seas. 1. Copepoda. Marine Investigations in

South Africa 3: 177-210.

De Decker, A. 1964. Observation on the ecology and distribution of Copepoda in marine

plankton of South Africa. Investigational Report. Division of Sea Fisheries, Union of South

Africa 49: 1-33.

De Decker, A. 1973. Agulhas Bank plankton. Jn: ZEITSCHEL, B. ed. Ecological studies

analysis and synthesis 3: 189-219. Berlin, Heidelberg, New York: Springer-Verlag.

De Decker, A. H. B. 1984. Near surface copepod distribution in the south-western Indian

and south-eastern Atlantic Ocean. Annals of the South African Museum 93 (5): 303-370.

DeEEvVEY, G. B. 1966. Seasonal variations in length of copepods in South Pacific New Zealand

waters. Australian Journal of Marine and Freshwater Research 17 (2): 155-168.

FARRAN, G. P. 1929. Crustacea. Part X. Copepoda. British Antarctic (‘Terra Nova’)

Expedition, 1910. Natural History Reports. Zoology 8: 203-306.

FLEMINGER, A. & HULSEMANN, K. 1977. Geographical range and taxonomic divergence in the

North Atlantic Calanus (C. helgolandicus, C. finmarchicus and C. glacialis). Marine

Biology 40: 233-248.

Grice, G. D. & HuLSEMANN, K. 1967. Bathypelagic calanoid copepods of the western Indian

Ocean. Proceedings of the United States National Museum 122 (3583): 1-67.

Harris, T. F. W. & VAN Foreest, D. 1978. The Agulhas Current in March 1969. Deep-Sea

Research 25 (6): 549-561.

Hutcuincs, L. 1985. Vertical distribution of mesozooplankton at an active upwelling site in

the southern Benguela Current, December 1969. Investigational Report. Sea Fisheries

Research Institute 129: 1-67.

Kemp, S., Harpy, A. C. & Mackintosu, N. A. 1929. Objects, equipment and methods.

Discovery Reports 1 (2): 141-232.

LUTJEHARMS, J. R. E. 1981. Features of the Southern Agulhas Current circulation from

satellite remote sensing. South African Journal of Science 77: 231-236.

LUTJEHARMS, J. R. E. 1985. Location of frontal systems between Africa and Antarctica: some

preliminary results. Deep-Sea Research 32: 1499-1509.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

OrreNn, M. J. 1966. Hydrology of the south west Indian Ocean. Investigational Report.

Division of Sea Fisheries, Republic of South Africa 55: 1-35.

SENO, J., Komaki, Y. & TAKEDA, A. 1963. Reports on the biology of the ‘Umitaka-Marw’

Expedition. Plankton collected by the ‘Umitaka-Marw’ in the Antarctic and adjacent

waters, with special references to Copepoda. Journal of the Tokyo University of Fisheries

49 (1): 53-62.

Swart, V. P. & Laraier, J. L. 1987. Thermal structure of Agulhas Bank water. In: PAYNE,

A. I. L., GULLAND, J. A. & Brink, K. H. eds. The Benguela and comparable ecosystems.

South African Journal of Marine Science 5: 243-253.

VeRvoorT, W. 1957. Copepods from Antarctic and Subantarctic plankton samples. Reports

B.A.N.Z. Antarctic Research Expedition (B) 3: 1-160.

VIDAL, J. W. 1968. Copepodos calanoideos epipelagicos de la Expedition Marchile II.

Gayana, Zoologia 15: 1-98.

WiporG, K. F. 1964. Marine copepods of Tristan da Cunha. Results of the Norwegian

Scientific Expedition to Tristan da Cunha 1937-1938 5 (51): 1-44.

YaASHNOV, V. A. 1958. Proiskhozhdenie vidov Calanus finmarchicus s.1. Origin of the species

Calanus finmarchicus s.1. Zoologicheskij Zhurnal 37 (6): 838-844. (In Russian. )

YaSHNOV, V. A. 1972. On the systematic status of Calanus glacialis, Calanus finmarchicus and

Calanus helgolandicus. Crustaceana 22 (3): 279-284.

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

fe SEBCIAL 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 see by

initials or full names

e.g. Du Toit but A.L. du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

+ 8: Joayte

A. H. B. DE DECKER, B. Z. KACZMARUK

G. MARSKA

A NEW SPECIES OF CALANUS

(COPEPODA, CALANOIDA)

FROM SOUTH AFRICAN WATERS

LXVOLUME 101 PART 4 SEPTEMBER 1991 | ISSN 0303-2515

ZAATHSON ane

Nov 21 1991

LIBRARIES

ANNALS

OF THE SOUTH

pans

CAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.

2. LAYOUT should be as follows:

(a) Centred masthead to consist of

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

; Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes. . .”

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

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

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

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

Note: no comma separating name and year

Pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 °#£zBand

September 1991 September

Part 4 Deel

DIE IN AFRIKA VORKOMMENDEN

COLOTES ERICHSON MIT EINFARBIG

METALLISCHEN FLUGELDECKEN

(COLEOPTERA, MALACHIIDAE)

von

W. WITTMER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

1, QOS, S29), SUD, 4-58 eo.) SES, 5, 79),

Al, ios, IED), 8, C2, 7), IOGES), IMCD, 5, 7, tori).

14(1=2), 15425) 242, 5) 27) 31123). 32G), 33)

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 124 5

Printed in South Africa by In Suid-Afrika gedruk deur

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

Old Mill Road, N’Dabeni, Cape Old Millweg, N’Dabeni, Kaap

D687

DIE IN AFRIKA VORKOMMENDEN COLOTES ERICHSON MIT

EINFARBIG METALLISCHEN FLUGELDECKEN

(COLEOPTERA, MALACHIIDAE)

(28. Beitrag zur Kenntnis der Fauna Afrikas)

von

W. WITTMER

Naturhistorisches Museum, Basel, Switzerland

(Mit 15 Abbildungen)

[MS accepted 1 December 1989|

ABSTRACT

The African species of Colotes with uniformly metallic elytra are revised, resulting in the

description of ten species new to science (C. cyaneovarius sp. nov., C. pseudochloropterus

sp. nov., C. pallidipes sp. nov., C. karkloofensis sp. nov., C. spinifer sp. nov., C. elongatipalpis

sp. nov., C. elongaticornis sp. nov., C. endroedyi sp. nov., C. hanangensis sp. nov. and

C. amplipalpus sp. nov.). A key to all presently known species of this group is provided, and

the maxillary palpi of all but two species are illustrated. Colotes jeanneli (Pic), C. innotatipennis

(Pic), C. evansi (Pic) and C. conradsi (Pic) are transferred from the genus Pseudocolotes to

Colotes, and P. reidi Pic is synonymized with C. innotatipennis (Pic). (In German.)

INHALT

PAGE

PTI LUN LNTGUET a cee coe ae Any ah cee Soe Os IIS As 45

Brix OMOMM Creare et orie Meemnee yA Sts Cones he apnea Sacto what 46

Bestimmunestabellemur@r@s soaaaaee aoe aecae ae eae 46

Beschncibumeemi.<.\ tierce. alee alte aie sa Ride eat coe ash 49

Paina COUN S CMY uci ea sees UN eie clea ye Wepre, Bis gees tlayecd acts 68

Iie ier pee ine Retr arie S care San ine minty ene Aer ane g 68

EINFUHRUNG

In dieser Arbeit wird versucht, eine Ubersicht zu geben, aller in Afrika vor-

kommenden Colotes-Arten mit einfarbig metallischen Fliigeldecken. Es lagen

mir die Typen von fiinf Arten vor, die dieses Merkmal aufweisen, wahrend

10 weitere vorgefunden wurden, die sich als neu fiir die Wissenschaft erwiesen.

Die weiteren aus Afrika gemeldeten Arten haben anders gefarbte Fligeldecken,

meist metallisch mit einem schmalen weiflich-gelben Seitensaum in der Mitte,

seltener vorwiegend gelbe oder schwarze Fliigeldecken. Das Material folgender

Museen konnte untersucht werden:

BM — British Museum (Natural History), London

MP — Muséum de Paris

NCI — National Collection of Insects, Pretoria

Ann. S. Afr. Mus. 101 (4), 1991: 45-69, 15 figs.

ANNALS OF THE SOUTH AFRICAN MUSEUM

NHMB — Naturhistorisches Museum, Basel

NMB —Nasionale Museum, Bloemfontein

SAM -—South African Museum, Cape Town

TMP —Transvaal Museum, Pretoria.

TAXONOMIE

BESTIMMUNGSTABELLE FUR OO

Halsschild eintarbig schwarz odermetallisch ..)5...... 4. ee Z

Halsschild einfarbig orange oder mit einem schwarzen Flecken oder

Band

Fuhlerglieder 1, 3 und 4 auf der Oberseite schwarz; 1 gegen die Spitze

schwach verbreitert, auf der Oberseite leicht eingedriickt; 3 und 4 deutlich

breiter als die folgenden. Kopf ohne Langsbeule. Hinterschienen ganz oder

teilweise gelb. Maxillarpalpen—Abb. 1A-B ...........................

Rn Teter ale, ra oe Oey See C. cyanopterus (Gorham, 1900) (see p. 49)

Fuhlerglieder 1 bis 4 gelb; 1 parallel, leicht abgeflacht; keine Glieder erwei-

tert. Langsbeule auf dem Kopf gegen den Halsschild deutlich verbreitert.

Hinterschienen schwarz. Maxillarpalpen— Abb. 2A-B ..................

BRE ie, SA ee Meaney teeee tis OR i ret eG CA C. cyaneovarius sp. nov. (see p. 49)

Halsschild orange mit einem schwarzen Flecken oder mit einem schwarzen

Lamesban@. . 266 6c ach seen es ee ede vee ees bee ~

Halsschild einfarbig orange . . 4 cc. cy ale ess od. ge oe ee 5

Kopf mit emer Langsbeule zwischen den Augen ...:.....-.5-.3===eee 5

Kopf ohne Langsbeule zwischen den Augen ..:.....:.52 2902 =e i

Langseindriicke des Kopfes gegen die Schlafen durch eine Leiste begrenzt,

oder an der Basis eine kurze, quere Beule. Maxillarpalpen mindestens zur

Halftehelll ¢ os. co.cc. 6 coe ees eek eS 2b ere a ee 6

Langseindriicke des Kopfes neben den Augen durchgehend, nirgends

unterbrochen. Maxillarpalpen fast vollstandig schwarz. Maxillar-

palpen— Abb. 3A—B. .. 065. sc ckeuaunagn. Sele oe + oe

Langseindrticke des Kopfes neben den Augen gegen die Schlafen durch

eine Leiste begrenzt; Langsbeule zwischen den Augen ungefahr in der

Mitte am breitesten, nach beiden Seiten hin schwach verschmalert;

Vorderstirne in der Mitte, fast am Vorderrande, mit einem kleinen spitzen

Dorn: Maxillarpalpen—Abb, 4A-B ......2...°.1..........

BR oe ry ihe RA a Heed C. pseudochloropterus sp. nov. (see p. 51)

Langseindrticke des Kopfes gegen die Schlafen nicht begrenzt, dafur

ungefahr in der Mitte am Innenrande der Augen eine kurze, quere Beule;

Langsbeule zwischen den Augen gegen den Halsschild regelmaig verengt;

Vorderstirne ohne kleinen spitzen Dorn in der Mitte. Maxillarpalpen—

ADD. DARIB 154 ee fees C. chloropterus Champion, 1922 (see p. 54)

10.

Hal.

12.

16.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 47

Vorderkopf ohne kleinen stumpfen Hocker tiber jeder Fuhlerwurzel...... 8

Vorderkopf mit einem kleinen stumpfen Hocker tiber jeder Fiihlerwurzel

= 0.16.0 0 67-0 Sen ye Saas are rarer ae C. conradsi (Pic, 1939) (see p. 66)

Fuhlerglied 2 kurzer als 3, oder so lang wie 3, einfach, nicht ausgehohlt .... 9

Fuhlerglied 2 langer als 3, auf der AufBenseite ausgehohlt. Maxillarpalpen

=== \|D/D) 5 C/A) 0) ae C. pallidipes sp. nov. (see p. 54)

Vorderrand des Vorderkopfes einfach, ohne Zahnchen in der Mitte;

Civ peusimicibeingedruckt . 4 see Ase oe el Re ie le ee oe ce 10

Vorderrand des Vorderkopfes mit einem kleinen spitzen Zahnchen in der

Mitte, Clypeus breit und ziemlich tief eingedriickt. Maxillarpalpen—

LNI01D.: VAN S3 0a eae eee ae ore C. karkloofensis sp. nov. (see p. 56)

Wiemigstens die’ Fuhlerglieder | bis4gelb.. 22... 2. eee. cece eee 11

Nur Fihlerglieder 1 bis 2 gelb. Maxillarpalpen— Abb. 8A-B .............

oe ek oie act a oe C. spinifer sp. nov. (see p. 58)

Wangen unter den Augen nicht ausgehohlt. Nur die ersten 4 Fuhlerglieder

CelbmGilied IauBben micht ausgerandet .... 62.2. $y. oe cee ree ee enue, 2

Wangen unter den Augen stark ausgehohlt. Fast die ganzen Fuhler gelb,

nur das letzte Glied angedunkelt, Glied 1 aussen stark ausgerandet. Maxil-

larpalpen—Abb. 9A-B............ C. elongatipalpis sp. nov. (see p. 58)

Fuhlerglied 1 an der breitesten Stelle wenig mehr oder etwas weniger als

doppeligcolonenl wie ander Basis. 25... be cee es ence ne sae 13)

Fuhlerglied 1 zur Spitze weniger stark verbreitert, gut dreimal so lang wie

Mmdoimonreitesien Stelle /Dreit .. 6... eke eee ee Suk eb ee eet ens enews 14

Fuhlerglied 2 so lang wie 3, ungefahr so breit wie dieses. Lange: 2,2 mm.

Fligeldecken starker punktiert, Zwischenraume glatt. Maxillarpalpen—

PNIO|D 5 L/S) 83) eae ee C. jeanneli (Pic, 1913) (see p. 62)

Fuhlerglied 2 viel ktirzer als 3 und viel schmaler als dieses. Lange: 3—3,5

mm. Fligeldecken weniger stark punktiert, Zwischenraume chagriniert.

Maxillarpalpen— Abb. 12A-B......... C. endroedyi sp. nov. (see p. 62)

Fuhlerglied 1 deutlich ktrzer als Glieder 2 bis 4. Maxillarpalpen—

ADO. SVE) 8) ee eee C. amplipalpus sp. nov. (see p. 67)

Fuhlerglied 1 fast so lang wie Glieder 2 bis 4. Maxillarpalpen—

/2\)0) 0% OVA Bai a a C. elongaticornis sp. nov. (see p. 61)

Flugeldecken sehr fein und dicht punktiert, fast matt, wenigstens die

Schenkel an der Basis mehr oder weniger dunkel ..................... 16

Flugeldecken sehr grob, wenig dicht punktiert, Beine einfarbig gelb.

Maxillarpalpen—Abb. 13A-B....... C. hanangensis sp. nov. (see p. 64)

Lange: 3—3,5 mm. Kopf von der Mitte der Augen nach vorne gelb, ohne

dunkle Flecken um die Futhlerwurzel. Maxillarpalpen gelb, Spitze des

letzten Gliedes angedunkelt, dieses mehr als doppelt so lang wie das vor-

letzte und ungefahr gleich breit. Maxillarpalpen— Abb. 12A-B...........

re ea hese hea kas C. endroedyi sp. nov. (see p. 62)

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

— Lange: 2,3 mm. Nur der Basalrand der Stirne schmal schwarz, um jede

Fuhlerwurzel ein dunkler Flecken. Maxillarpalpen gelb, letztes Glied der-

selben klein, viel schmaler als das vorletzte. Maxillarpalpen—Abb. 14A-—B

Be ential ca reed ah, Bie C. innotatipennis (Pic, 1932) (see p. 65)

BESCHREIBUNGEN

Colotes cyanopterus (Gorham, 1900)

Abb. 1A-B

Charopus cyanopterus Gorham, 1900: 75.

Colotes cyanopterus (Gorham) Champion, 1922: 339.

Bemerkungen

Champion (1922) hat bereits ausftihrlich tiber diese Art berichtet. Auer

dem Material im SAM und den 2 9 im BM, habe ich erst 1 © gesehen, das im

Januar 1979, in Winburg, Orange Free State, erbeutet wurde. Fur die Auf-

nahmen der Unter- und Oberseite der Maxillarpalpen mit dem Raster-

elektronenmikroskop (Abb. 1A—B) wurde dieses Exemplar verwendet.

Colotes cyaneovarius sp. nov.

Abb. 2A-B

Beschreibung

CO’. Vorderkopf gelb, Stirne ziemlich breit, Innenrand der Augen bis kurz

uber den Fuhlerwurzeln und Unterseite schwarz, die schwarze Farbung dringt

an der Spitze des Mittelwulstes kurz dreieckig in die gelbe Farbung ein; Fuh-

lerglieder 1 bis 4 oder 1 bis 5 gelb, restliche schwarz; Halsschild und Schildchen

schwarz, selten mit schwachem Metallschimmer. Fliigeldecken schwarz mit

blauem oder grunlichem Metallschimmer; die 4 vorderen Beine gelb, aber

Schenkel mit basaler schwarzer Farbung, die oft bis fast zu den Spitzen steigt;

Hinterbeine schwarz, nur die auBerste Spitze der Tibien und die Tarsen mit-

unter teils schwach aufgehellt.

Kopf mit den Augen so breit wie der Halsschild; Stirne neben den Augen

schwach breit langseingedriickt, dadurch der Langswulst zwischen den Augen,

der sich gegen die Stirne leicht verbreitert und bis auf die Hohe des Hinter-

randes der Augen reicht, noch besser abhebend; Vorderstirne zwischen den

Fuhlerwurzeln breit eingedruckt; Clypeus in der Mitte mit einem kleinen, fast

dreieckigen, queren Aufsatz; Maxillarpalpen (Abb. 2A-—B) gelb, an der Spitze

leicht gebraunt. Fuhler ca. ein Funftel kurzer als die Flugeldecken; Glied 1 sehr

lang und breit, flach, deutlich langer als 2 und 3 zusammen, Aufenrand fast

gerade, Innenrand vor der Mitte schwach, stumpfwinklig verbreitert; 2 bis 11

normal, der Innenrand von 4 bis 7 etwas starker gerundet als bei den ubrigen,

2 deutlich kurzer als 3. Halsschild breiter als lang (23 X 16); Seiten stark gerun-

det; Scheibe regelmaBig gewolbt, fein chagriniert. Fligeldecken nach hinten

leicht verbreitert, chagriniert.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN

Abb. 1. Maxillarpalpen von Colotes cyanopterus (Gorham), C. 163 xX.

A. Wahrscheinlich Oberseite. B. Wahrscheinlich Unterseite.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abb. 2. Maxillarpalpen von Colotes cyaneovarius sp. nov., CO. 163 X.

A. Oberseite. B. Unterseite.

@. Kopf einfarbig schwarz, Hintertarsen aufgehellt, sonst wie das &

gefarbt.

Lange: 2,5 mm.

Material

South Africa, Transvaal: Cycas Farm, 5. Nov. 1985, grass-netting

E-Y:2262, S. Endré6dy-Younga (Holotypus und 5 Paratypen—TMP, 3 Para-

typen—NHMB); E-Y:2297, S.Endrédy-Younga (3 Paratypen—TMP,

1 Paratypus—NHMB).

Verwandtschaft

Diese Art ist in der Farbung nur mit C. cyanopterus (Gorham) zu verglei-

chen, der wie C. cyaneovarius einen einfarbig schwarzen oder schwach

metallischen Halsschild besitzt. Von C. cyanopterus (Gorham) unterscheidet

sich die neue Art sehr wesentlich, vergleiche Bestimmungstabelle. Im Bau des

Kopfes hingegen, steht sie C. Jouwi Wittmer (1989) naher, ist jedoch von dieser

Art verschieden durch die hellen ersten Fuhlerglieder, den ganz dunklen Hals-

schild und die hellen Maxillarpalpen, die bei C. Jouwi verschieden geformt und

fast vollstandig dunkel sind.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 51

Colotes scotti Wittmer, 1953

Abb. 3A-B

Colotes scotti Wittmer, 1953: 88.

Bemerkungen

Von Lesotho (Basutoland) beschrieben, liegt nun die Art auch noch von

folgendem Fundort vor: Orange Free State, Golden Gate Highlands National

Park, SE 2828 Bc, 15.—16. Oct. 1985, ex Leucosidea sericea, 2 Ex. NMB, 2 Ex.

NHMB.

Fur die Aufnahmen der Maxillarpalpen (Abb. 3A-B) diente ein Exemplar

vom Golden Gate.

Abb. 3. Maxillarpalpen von Colotes scotti Wittmer, CG. 163 x. A. Oberseite. B. Unterseite.

Colotes pseudochloropterus sp. nov.

Abb. 4A-B

Beschreibung

CO. Kopf schwarz, mit 3 gelben Langsbandern; mittleres auf dem

Langswulst, gegen die Stirne ausgerandet und nach vorne bis zu der Oberlippe

reichend, je ein seitliches neben den Augen bis zu den Wangen; die schwarzen

Bander, die die gelben von dem mittleren trennen, sind im oberen Teil oft ver-

schmalert oder ganz unterbrochen; Maxillarpalpen gelb, Spitze des letzten

Gliedes auf der ganzen Breite schmal angedunkelt; Fihler schwarz, Glied 1

orange mit einem schwarzen Langsband auf der Innenseite, 2 und 4 gelb, letzteres

Sy ANNALS OF THE SOUTH AFRICAN MUSEUM

manchmal mit einem ganz kleinen dunklen Flecken an der Spitze oben, 3 gelb

mit der Spitzenhalfte schwarz, 5 an der Basis kurz gelb. Halsschild mit mehr

oder weniger breiten orangenen Seiten, in der Mitte durchgehend schwarz;

Schildchen schwarz; Flugeldecken dunkelblau metallisch; 4 Vorderschenkel bis

gegen die Knie schwarz, 4 Vordertibien gelb, 4 Vordertarsen mehr oder weniger

angedunkelt, Hinterbeine schwarz.

Kopf mit den Augen ein wenig schmaler als der Halsschild, Stirne neben

dem Langswulst schwach langseingedrtickt, dieser Langseindruck gegen die

Stirne und gegen die Schlafen durch eine schrage Leiste abgegrenzt; Langswulst

in der Mitte mehr oder weniger deutlich langseingedrtickt, der Wulst etwas vor

der Mitte am breitesten und nach beiden Seiten schwach verengt; fast am Vor-

derrand des Vorderkopfes in der Mitte ein kleiner, spitzer Dorn; Clypeus auf

der ganzen Breite tief eingedriickt; Maxillarpalpen (Abb. 4A—B) nach einem

Exemplar von Leydenburg. Fuhler ca. 10 Prozent ktrzer als die Fligeldecken;

Glied 1 ungefahr so lang wie 2 und 3, ziemlich breit, etwas abgeflacht, Seiten

leicht gerundet; 2 ein wenig kurzer als 4; 3 zur Spitze nach innen stark erweitert;

5 langer und etwas breiter als die folgenden, Spitze ganz wenig tber die Basis

von 6 vorgezogen. Halsschild breiter als lang (30 x 18); Seiten stark gerundet;

Basalrand in der Mitte schmal ausgerandet; Scheibe regelmaBig gewolbt, fein

chagriniert. Flugeldecken nach hinten leicht verbreitert, deutlich punktiert,

dazwischen chagriniert oder glatt.

2. Kopf einfach, einfarbig schwarz, Fuhlerglied 1 gelb mit einem schmalen

schwarzen Langsflecken auf der Oberseite, 2 bis 4 gelb, sonst wie das CO

gefarbt.

Ldnge: 2,8—3 mm.

Material

Transvaal: 33 km west of Volksrust, Rusthoek Farm, R543, 9. Okt. 1986,

W. Wittmer (Holotypus—TMP, 4 Paratypen—NHMB); 21-22 km south of

Leydenburg, 15. Nov. 1983, W. Wittmer (1 Paratypus—NHMB); Nelshoogte

Forest Station, 1. Dez. 1986, E-Y:2336, 2337, S. Endrédy-Younga (3 Para-

typen— TMP); Berlin Forest Station, carst plat., 8. Dez. 1986, E—Y:2368, 2365,

S. Endrédy-Younga (7 Paratypen—TMP). Natal: Royal Natal National Park,

1500 m, 8. Okt. 1986, W. Wittmer (1 Paratypus—TMP, 2 Paratypen—

NHMB); Cathedral Peak, Drakensberg, 6. Nov. 1981, J. & S. Klapperich

(12 Paratypen—NHMB); idem 29°00’S 29°27'E, 11.-12. Nov. 1981, S. J. van

Tonder, C. Kok (4 Paratypen—NCI); idem, Arensig Mt., 20. Dez. 1986,

P. Reavell (1 Paratypus—NHMB); Giant’s Castle Reserve, Drakensberg,

1 600 m, 29°16’S 29°31’E, 10. Okt. 1983, C. Bellamy (2 Paratypen—NHMB);

Oliviershoekpas, 8. Okt. 1986, W. Wittmer (2 Paratypen—NHMB). Orange

Free State: Golden Gate Highlands National Park, Drakensberg, 6. Nov. 1981,

J. & S. Klapperich (19 Paratypen—NHMB); Golden Gate Highlands National

Park, SE 2828 Bc, 15.-16. Okt. 1985, 13.-14. Nov. 1985, 3. Dez. 1985, ex

Leucosidea sericea (8 Paratypen—NMB, 1 Paratypus—NHMB).

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN

B. Unterseite.

llarpalpen von Colotes pseudochloropterus sp. nov., CO. 163 X.

A. Oberseite.

Max

Abb. 4.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

Verwandtschaft

Eine mit C. chloropterus Champion nahe verwandte Art, die sich

hauptsachlich durch die verschieden gebildeten Maxillarpalpen und dem ver-

schieden gebauten Kopf unterscheidet (vergleiche Bestimmungstabelle).

Colotes chloropterus Champion, 1922

Abb. 5A-B

Colotes chloropterus Champion, 1922: 340. Wittmer, 1956: 1075.

Bemerkungen

Diese Art wurde von Champion nach 2 9 aus Howick, Natal, beschrieben.

Vom gleichen Fundort: Howick, Umgeni River, Lions River District, N. Leleup

(TMP) besitze ich 1 C’, das ich dieser Art zuordnen kann. Des weiteren lag mir

Material von folgenden Fundorten vor: Orange Free State: Harrismith, south-

west foot of Platberg, 2 000 m, 20. Febr. 1984, 28°15’S 29°09’E, R. Oberprieler

(NCI und NHMB); Natal: Coleford Nature Reserve, 16. Nov. 1981, 29°57’S

29°27'E, S.J. van Tonder, C. Kok (NCI); Cathedral Peak, Drakensberg,

6. Nov. 1981, J. & S. Klapperich (NHMB, TMP); Giant’s Castle Reserve,

Drakensberg, 1 600 m, 29°16’S 29°31’E, C. Bellamy (NHMB).

Fur die Aufnahmen der Maxillarpalpen (Abb. SA—B) diente ein Exemplar

von Cathedral Peak.

Colotes pallidipes sp. nov.

Abb. 6A-B

Beschreibung

©’. Kopf schwarzlich, Vorderkopf in der Mitte kurz, oder bis fast zur Stirn-

basis, orange aufgehellt, Maxillarpalpen (Abb. 6A—B) orange, mit dem gr6Bten

Teil des vorletzten Gliedes dunkel; Fithler schwarzlich, erste 4 bis 6 Glieder

gelb; Halsschild orange mit einer kleinen dunklen Makel auf der vorderen

Halfte, den Basalrand schmal frei lassend, selten einfarbig orange; Schildchen

und Flugeldecken schwarz mit schwachem olivenem Metallschimmer; Beine

orange.

Kopf mit den Augen schmaler als der Halsschild; Stirne leicht gewolbt,

Vorderstirne gegen die Wangen ganz schwach erhoht, fein mikrochagriniert.

Fuihler nur wenig langer als doppelt so lang wie der Halsschild; Glied 2 etwas

langer als 4, auf der AuBenseite ausgerandet; 3 fast quadratisch, breiter als 4;

5 ein wenig langer und schmaler als 4. Halsschild breiter als lang (19 x 15);

Seiten gegen die Basis gerundet verengt, regelmaBig gewdlbt, wie der Kopf

chagriniert. Fliigeldecken hinter der Mitte am breitesten, dann gegen die

Spitzen gerundet verengt, ziemlich dicht und tief punktiert, Abstande der

Punkte kleiner bis gr6Ber als ihr Durchmesser, gefltgelt.

55)

iff

ion,

170 X

ite.

‘

B. Unterse

163K

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN

Maxillarpalpen von Colotes chloropterus Champ

A. Oberseite.

Abb. 5.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abb. 6. Maxillarpalpen von Colotes pallidipes sp. nov., CO. 163 X.

A. Oberseite. B. Unterseite.

2. Kopf schwarz, Wangen und Unterseite teils orange, Maxillarpalpen

orange, sonst wie das CO gefarbt. Fuhler einfach, Glied 2 nur wenig langer als 3.

Ungefltigelt, Schulterbeulen weniger ausgebildet als beim GC’.

Lange: 2 mm.

Material

Kenya: Marakwet, Elgeyo Escarpment, 2500 m, Mission de l’Omo,

1932-33 (Holotypus und 6 Paratypen—MP, 1 Paratypus—NHMB); Elgon Saw

Mill, Mt. Elgon, versan Est (Camp II), 2 470 m (3 Paratypen—MP, 3 Para-

typen—NHMB); Mt. Elgon, versan Est, prairies alpines, 3 000-4 000 m

(1 Paratypus— MP).

Verwandtschaft

Nahe verwandt mit C. jeanneli (Pic) und C. elongaticornis sp. nov. Die

neue Art unterscheidet sich durch das lange, am Aufenrande ausgerandete

Fuhlerglied 3 und die verschieden geformten Maxillarpalpen.

Colotes karkloofensis sp. nov.

Abb. 7A-B

Beschreibung

©. Kopf gelbbraun, an der Basis schwarz, schwarze Farbung neben den

Augen immer schmaler werdend bis fast zu den Wangen; Maxillarpalpen

(Abb. 7A-B) gelb, nur die Spitze ganz schwach dunkel; Fuhler schwarz, Unter-

seite der beiden ersten Glieder gelb; Halsschild schwarz, Seiten ziemlich breit

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN >)//

Abb. 7. Maxillarpalpen von Colotes karkloofensis sp. nov., &. 120 x.

A. Oberseite. B. Unterseite.

orange; Schildchen und Fligeldecken dunkelblau metallisch; 4 Vorderbeine

gelb, mit der Basis der Schenkel schwarz, Hinterbeine schwarz.

Kopf mit den Augen schmaler als der Halsschild; Stirne zwischen den

Augen flach; Vorderstirne mit einem kleinen Zahnchen in der Mitte am Vorder-

rand, Fuhlerwurzeln daneben erh6dht; Clypeus breit und tief eingedriickt.

Fuhler um ca. ein Funftel kurzer als die Fliigeldecken; Glied 1 lang, so lang wie

3 und 4 zusammen, gegen die Spitze nur wenig erweitert; 2 deutlich kirzer als 4;

3 langer als 4; 4 und 5 schwach plattgedriickt, so breit wie 3, breiter als die fol-

genden, 6 etwas langer als 5, langer als die folgenden. Halsschild breiter als lang

(28 x 19), Seiten stark gerundet, gegen die Basis starker als nach vorne; Basal-

rand vor dem Schildchen schwach ausgerandet, Scheibe regelmaig gewolbt,

schwach matt. Flugeldecken nach hinten leicht erweitert, zerstreut punktiert,

Abstand der Punkte meistens groBer als ihr Durchmesser, Zwischenraume fein

chagriniert.

Lange: 2,8 mm.

Material

South Africa, Natal: Karkloof Falls Nature Reserve, 22 km north of Pieter-

maritzburg, 6. Okt. 1986, W. Wittmer (Holotypus—NHMB).

Verwandtschaft

Diese Art ist neben C. spinifer sp. nov. zu stellen, der nachfolgend

beschrieben wird.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colotes spinifer sp. nov.

Abb. 8A-B

Beschreibung

©. Kopf gelb, Stirnbasis fast bis zur Mitte der Augen schwarz; Maxillarpal-

pen (Abb. 8A-B) orange mit leicht angedunkelten Spitzen; Fuhler schwarz,

Glieder 1 und 2 ganz gelb, 3 auf der Unterseite aufgehellt; Halsschild orange

mit einer quadratischen schwarzen Langsmakel, die weder den Vorderrand noch

die Basis erreicht; Schildchen schwarz; Fliigeldecken dunkelblau metallisch;

Beine orange.

Kopf mit den Augen schmaler als der Halsschild; Stirne mit 2 schwachen

Beulen, die in der Mitte durch einen seichten, kurzen Langseindruck von ein-

ander getrennt sind; Vorderstirne schwach flach eingedrtickt; Clypeus sehr kurz.

Fuhler ca. 10 Prozent kurzer als die Flugeldecken; Glied 1 fast parallel, ein

wenig langer als 3 und 4; 2 kurzer als 3, 3 bis 5 leicht verbreitert, ein wenig

langer als die folgenden. Halsschild breiter als lang (30 X 20); Seiten stark

gerundet, Basalecken fast vollstandig mit der Basis verrundet, Oberflache

gewirkt. Flugeldecken nach hinten leicht erweitert, grob punktiert, Abstand der

Punkte kleiner als ihr Durchmesser, Behaarung ziemlich lang, schrag abstehend.

Lange: 2,8 mm.

Material

South Africa, Natal: Giant's Castle Reserve, Drakensberg, 1 600 m,

9. Nov. 1983, C. Bellamy (Holotypus—NHMB).

Verwandschaft

Diese Art ist neben C. karkloofensis sp. nov. zu stellen, unterscheidet sich

aber durch ihre verhaltnismaBig grobe, lange, halb abstehende Behaarung auf

den Fliigeldecken, die starker als tblich punktiert sind, und die Form des

letzten Gliedes der Maxillarpalpen, die auf der einen Seite mit einem sehr

langen, kraftigen, gebogenen Fortsatz ausgestattet sind, wahrend auf der ent-

gegengesetzten Seite ein langer, leicht gebogener, feiner, teils durchsichtiger,

spatelfO6rmiger Fortsatz steht.

Colotes elongatipalpis sp. nov.

Abb. 9A-B

Beschreibung

©’. Kopf gelbbraun, Basis schwarz, ebenso neben den Augen bis zu den

Schlafen; Maxillarpalpen orange mit dunkler Spitze; Fiihler orange, nur das

letzte Glied mehr oder weniger gebraunt; Halsschild orange mit einem breiten

dunklen, griinlich schimmernden Langsband, das den Vorderrand erreicht,

nicht aber die Basis; Schildchen und Fligeldecken dunkelgriin metallisch; Beine

3y,

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN

Wi)

WA) Se

nov.,

nifer sp

ite

B. Unterse

Woe

Oberse

illarpalpen von Colotes sp

Max

Abb. 8.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abb. 9. Maxillarpalpen von Colotes elongatipalpis sp. nov., CO. 163 X.

A. Oberseite. B. Unterseite.

orange, nur die Hinterschenkel einfarbig schwarz, oder héchstens an der Basis

ein wenig aufgehellt.

Kopf mit den Augen schmaler als der Halsschild; Stirne zwischen den

Augen ziemlich flach; Wangen unter den Augen stark ausgehohlt; Maxillar-

palpen (Abb. 9A-B) nach einem Exemplar von Groot Brakrivier. Fihler um

ca. 10 Prozent ktrzer als die Flugeldecken; Glied 1 so lang wie 4 und 5; 2 deut-

lich kurzer als 3; 4 bis 6 ein wenig breiter als die folgenden. Halsschild breiter

als lang (28,5 xX 22); Seiten stark gerundet; Basalecken fast vollstandig mit der

Basis verrundet; Scheibe regelmaBig gewolbt, feinstens gewirkt, dazwischen

ganz feine Punkte sichtbar (64 x). Fliigeldecken nach hinten leicht verbreitert,

ziemlich dicht, etwas erloschen punktiert, Abstand der Punkte grofer als ihr

Durchmesser, Zwischenraume nicht ganz glatt.

2. Kopf vorwiegend schwarz, bei einem Exemplar sind die Wangen hell,

bei einem weiteren auch der vorderste Rand des Vorderkopfes und bei zweien

der groBte Teil des Vorderkopfes hell; Fihler schwarz, 4 bis 6 erste Glieder

gelb; Halsschild wie beim ©’, oder fast einfarbig, oder ein Flecken nur angedeu-

tet; Beine wie beim ©’, ausgenommen die Hinterschenkel, die an den Knien

kurz hell sind.

Lange: 3—3,2 mm.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 61

Material

South Africa, Cape Province: Algoa Bay, Dr. Brauns (Holotypus—TMP);

Buffalo Bay, 26. Nov. 1967, H. K. Munro (1 Paratypus OC’ —NCI); Groot Brak-

rivier (Mossel Bay), 6. Okt. 1984, W. Wittmer (7 Paratypen—NHMB).

Verwandtschaft

Diese Art ist neben C. innotatipennis (Pic) zu stellen, sie ist jedoch viel

groBer und unterscheidet sich leicht durch die unter den Augen stark ausgehohl-

ten Wangen und die verschieden gebauten Maxillarpalpen.

Colotes elongaticornis sp. nov.

Abb. 10A-B.

Beschreibung

CO. Kopf orange, Basis nicht ganz bis zur Mitte der Augen schwarz, oder

die schmalen, nach vorne verschmalerten schwarzen Streifen bis zum Clypeus;

Maxillarpalpen (Abb. 10A-—B) gelb; Fiihler dunkel, die ersten 4 bis 5 Glieder

gelb; Halsschild orange mit einer kleinen, fast runden dunklen Makel, mehr

gegen den Vorderrand als gegen die Basis gelegen; Schildchen und Flugel-

decken mit dunkelgriinem Metallschimmer; Beine gelb, nur die Hinterschenkel

auf der basalen Halfte angedunkelt.

Kopf mit den Augen schmaler als der Halsschild, Stirne leicht gewolbt,

kaum sichtbar mikrochagriniert, fast glatt (64 x). Fuhler ca. 2,2 mal langer als

der Halsschild; Glied 1 sehr lang, ein wenig langer als 2 und 3, gegen die Spitze

—

©)

lon]

Abb. 10. Maxillarpalpen von Colotes elongaticornis sp. nov.,

B. Unterseite.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

allmahlich verbreitert; 2 fast so lang wie 3, jedoch ein wenig breiter. Halsschild

breiter als lang (20 X 15,5); Seiten stark gerundet, von der Mitte zur Basis

starker als nach vorne; Scheibe regelmaBig gewolbt, etwas deutlicher als der

Kopf chagriniert. Flugeldecken hinter der Mitte am breitesten, dann gegen die

Spitze gerundet verengt, dicht und tief punktiert, Abstande der Punkte kleiner

als ihr Durchmesser, Zwischenraume glatt, Fligel vorhanden.

Q. Bisher unbekannt, dtrfte ungefltiigelt sein.

Lange: 2 mm.

Material

Athiopien: Mt. Maigudo, R. Linnavuori (Holotypus—NHMB); Belleta

Forest, 13.-14. Juni 1963, R. Linnavuori (1 Paratypus CO —NHMB).

Verwandtschaft

Diese Art ist neben C. pallidipes sp. nov. zu stellen, sie unterscheidet sich

jedoch durch das nicht ausgehohlte Fuhlerglied 2 und durch Fuhlerglied 3, das

fast so lang ist wie 2. Bei C. pallidipes ist Fihlerglied 3 breiter als 2, kurzer als

dieses und fast quadratisch.

Colotes jeanneli (Pic, 1913) comb. nov.

Abb. 11A-B

Pseudocolotes jeanneli Pic, 1913: 231.

Pseudocolotes jeanneli var. testaceiceps Pic, 1913: 232. (syn. nov.)

Bemerkungen

Im MP befindet sich eine langere Serie von Syntypen, die alle von Kijabé

(Kikuyu Esct.) stammen, daraus wurde ein Lectotypus gewahlt, wahrend die

ubrigen als Paralectotypen bezeichnet wurden. Das C& mit hellem Kopf hat Pic

als C. jeanneli var. testaceiceps beschrieben, diese Varietat ist als Synonym zu

betrachten.

Fur die Abbildung 11A—B der Maxillarpalpen wurde ein Paratypus aus

dem MP verwendet.

Colotes endroedyi sp. nov.

Abb. 12A-B

Beschreibung

CO’. Vorderkopf gelb, von der Mitte der Augen bis zur Basis schwarz; Maxil-

larpalpen (Abb. 12A-—B) gelb mit schwarzer Spitze; Fuhler schwarz, vier erste

Glieder und 5 auf der Unterseite gelb; Halsschild einfarbig orange oder mit

einer schwarzen, etwas verschwommenen Makel, die weder den Vorderrand

noch den Basalrand erreicht; Schildchen schwarz; Fligeldecken dunkelblau

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 63

SSSSSSSSS

Abb. 11. Maxillarpalpen von Colotes jeanneli (Pic), O&. 163 x.

A. Seitlich. B. Unterseite.

metallisch; Beine gelb, Schenkel an der Basis mehr oder weniger breit, bei den

hinteren bis kurz vor den Knien, schwarz.

Kopf mit den Augen kaum merklich schmaler als der Halsschild; Stirne und

Vorderstirne leicht gewolbt, feinstens gewirkt, fast matt. Fuhler verhaltnis-

mafBig kurz, ca. 2,1 mal langer als der Halsschild, Glied 1 fast dreieckig, ein

wenig kurzer als 3 und 4; 2 sehr kurz; 3 so lang wie 4; 3 und 4 ein wenig breiter

als die folgenden, die an Breite leicht abnehmen. Halsschild breiter als lang

(29 x 22); Seiten stark gerundet; Scheibe regelma%ig gewolbt, fein gewirkt.

Fligeldecken zur Spitze leicht erweitert, ziemlich dicht, etwas erloschen

punktiert.

@. Kopf schwarz, nur die Wangen in der Nahe der Mandibelbasis kurz auf-

gehellt, Makel auf dem Halsschild gut ausgebildet, sonst wie das C’ gefarbt.

Lange: 3-3,5 mm.

Material

South Africa, Cape Province: Bothastrand, ground and vegetation, 6. Okt.

1984, 34°03'S 22°18’E, E-Y:2137, S. Endr6édy-Younga (1 Holotypus co’, 1 Para-

typus 9 —TMP; | Paratypus O'—NHMB).

Verwandtschaft

Diese Art ist neben C. elongatipalpis sp. nov. zu stellen und 4hnlich

gefarbt, unterscheidet sich aber durch die breiteren Fuhler und das viel kurzere,

fast dreieckige Glied 1, das oben nicht ausgehohlt ist, und durch die verschieden

gebildeten Maxillarpalpen.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abb. 12. Maxillarpalpen von Colotes endroedyi sp. nov., CO. 163 xX.

A. Oberseite. B. Unterseite.

Colotes hanangensis sp. nov.

Abb. 13A-B

Beschreibung

OC’. Kopf, Halsschild und Beine gelb-orange, an der Stirnbasis unter dem

Halsschildvorderrand durchscheinend, eine kleine dunklere Stelle in der Mitte;

Fuhler gelb, die letzten 4 bis 5 Glieder kaum merklich braunlich; Schildchen

und Flugeldecken schwarz, kaum metallisch schimmernd; Maxillarpalpen

(Abb. 13A—B) gelb, Spitzen kaum gebraunt.

Kopf mit den Augen schmaler als der Halsschild; Stirne bis zur

Vorderstirne ganz leicht gewolbt, glatt. Fuhler um ca. ein Viertel ktrzer als die

Fligeldecken; Glied 1 langer als 2 und 3; 2 und 3 ungefahr gleich lang und

gleich breit. Halsschild breiter als lang (20 x 15), Seiten gerundet, Basalecken

mit der Basis verrundet, glatt. Fliigeldecken nach hinten leicht erweitert, grob

und tief punktiert, dazwischen fast glatt.

Q. In der Farbung mit dem CO tibereinstimmend, ausgenommen der ein-

farbig schwarze Kopf und der Halsschild, der Tendenz zu Verdunklung zeigt;

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 65

AB - B

Abb. 13. Maxillarpalpen von Colotes hanangensis sp. nov., O. 163 x.

A. Oberseite. B. Unterseite.

beim einen Exemplar ist eine verschwommene Langsmakel angedeutet und

beim anderen ist sie deutlich sichtbar.

Lange: 2 mm.

Material

Tanzania: Mt. Hanang, versand Sud, 2 500 m, 26. Mai. 1957, haut taillis

avec Protea, P. Basilewsky, N. Leleup, Holotypus © und 1 Paratypus Q

(Musée R. de I’Afrique Central), 1 Paratypus 9 (NHMB).

Verwandtschaft

Diese Art konnte ich erst jetzt mit Colotes jeanneli (Pic) vergleichen, fir

die ich sie gehalten hatte (Wittmer 1962: 263). Sie stellte sich als neu heraus und

ist mit C. jeanneli (Pic), sowie mit C. pallidipes sp. nov. nahe verwandt. Sowohl

durch den einfarbig gelb-orangenen Halsschild und fast ebenso einfarbigen

Kopf, sowie durch die verschieden gebauten Maxillarpalpen ist sie von diesen

beiden Arten leicht zu trennen.

Colotes innotatipennis (Pic, 1932) comb. nov.

Abb. 14A-B

Pseudocolotes innotatipennis Pic, 1932: 6.

Pseudocolotes reidi Pic, 1945-48: 62. (syn. nov.)

Bemerkungen

Ich hatte Gelegenheit Material von C. reidi (Pic), das Evers mit dem Holo-

typus vergleichen konnte, dem MHolotypus von C. innotatipennis (Pic)

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abb. 14. Maxillarpalpen von Colotes innotatipennis (Pic),O. 163 x. A. Oberseite.

B. Unterseite.

gegenuberzustellen. Die beiden Arten stimmen tberein, so daB C. reidi (Pic) in

Synonymie verfallt.

Fur die Photographien der Maxillarpalpen (Abb. 14A—B) wurde 1 O’ von

Mt. Selinda (NHMB) verwendet.

Material

Mozambique and Natal.

Colotes conradsi (Pic, 1939) comb. nov.

Pseudocolotes conradsi Pic, 1939: 166.

Pseudocolotes conradsi var. notaticollis Pic, 1939: 169.

Pseudocolotes conradsi var. insignatus Pic, 1939: 169.

Bemerkungen

Diese in der Farbung sehr variable Art liegt in einer Serie im Material des

MP vor. Die @ sind meistens dunkler gefarbt als die CO’, manchmal einfarbig

schwarz. Es handelt sich um eine weitere Art mit grob punktierten Flugel-

decken, deren G’ durch eine abweichende Kopfform gekennzeichnet sind. Uber

jeder Fihlerwurzel befindet sich ein kleiner stumpfer Hocker, die Fuhlerglieder

2 und 3 sind praktisch gleich lang, wobei 2 ein wenig breiter ist als 3.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 67

Abb. 15. Maxillarpalpen von Colotes amplipalpus sp. nov., &. 170 x.

A. Oberseite, letztes Glied leicht defekt. B. Unterseite.

Colotes amplipaipus sp. nov.

Abb. 15A-B

Beschreibung

©. Kopf schwarz, Wangen orange, Vorderstirn mit einer kleinen, runden,

kaum angedeuteten Aufhellung in der Mitte; Maxillarpalpen (Abb. 15A-B)

braun, gerundet, nach hinten gerichteter Teil des vorletzten Gliedes

schwarzlich; Fihler schwarzlich, 3 erste Glieder hell; Halsschild orange mit

einem dunklen Langsflecken in der Mitte, der den Vorderrand beruhrt, nicht

aber den Basalrand; Schildchen und Fligeldecken schwarz, letztere kaum wahr-

nehmbar metallisch, Schenkel orange, Schienen und Tarsen leicht angedunkelt.

Kopf mit den Augen schmaler als der Halsschild; Stirne und Vorderstirne

leicht gew6lbt, einzelne Punkte erkennbar, dazwischen glatt. Fuhler um ca. ein

Finftel kiirzer als die Fligeldecken, Glied 1 fast so lang wie 2 bis 4; 2 deutlich

kurzer als 3. Halsschild breiter als lang (19 x 13); Seiten stark gerundet; Basal-

ecken nicht ganz mit der Basis verrundet, regelmaBig gewolbt, wie der Kopf

skulptiert. Fligeldecken nach hinten leicht erweitert, Punktierung grob und tief,

Zwischenraume glatt.

Lange: 2 mm.

Material

Tanzania: Magamba, Usambara, Nov. 1959 (Holotypus—NHMB).

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Verwandtschaft

Die Art ist neben C. jeanneli (Pic) zu stellen, aber durch das kitrzere

Fuhlerglied 2 und die Maxillarpalpen, vergleiche Abbildung 11A—B, zu unter-

scheiden.

Incertae sedis

Colotes evansi (Pic, 1930) comb. nov.

Pseudocolotes evansi Pic, 1930: 77.

Bemerkungen

Das Holotypus 9 befindet sich im BM und konnte nicht gedeutet werden,

weil das O fehlt.

DANSAGUNGEN

Folgenden Kolleginnen und Kollegen, die mir durch Ubermittlung von

Material behilflich waren, danke ich verbindlichst fiir ihre Hilfe: Dr. M. Bran-

cucci, Basel; Mrs. M. A. Cochrane, Cape Town; Dr. S. Endrédy-Younga,

Pretoria; Dr. S. Louw, Bloemfontein; Dr. J. J. Menier, Paris; Mr. R. Ober-

prieler, Pretoria; Mrs. E. R. Peacock, London; Dr. V. B. Whitehead, Cape

Town.

Zu besonderem Dank bin ich auch Herrn Dr. R. Guggenheim, Leiter des

Laboratoriums fir Raster-Elektronenmikroskopie, Geologisch-Palaontolo-

gisches Institut der Universitat Basel, sowie seinem Mitarbeiter Herrn

M. Duggelin, fur die sorgfaltigst ausgefuhrten Aufnahmen verpflichtet.

LITERATUR

CHAMPION, G. C. 1922. Revision of the S. African species of Dinometopus, Troglops, Chali-

corus, Colotes, Helcogaster, and the allied genera, with an account of their

accessory O’-characters (Coleoptera). Annals and Magazine of Natural History (9) 10 (58):

309-358.

GorHaM, H. S. 1900. Descriptions of new genera and species of Coleoptera from South and

West Africa, of the section Serricornia, and of the families Erotylidae, Endomychidae and

Languridae. Annals and Magazine of Natural History (7) 5 (25): 71-94.

Pic, M. 1913. Collections recueillies par MM. Alluaud et Jeannel dans l’Afrique Orientale.

Diagnoses préliminaires de Coléoptéres, Malachids, Dasytides, Hylophilides. Bulletin du

Muséum d histoire naturelle, Paris 1913 (4): 231-232.

Pic, M. 1930. Malacodermes exotiques. Echange (hors-texte) 46: 77-92.

Pic, M. 1932. Contribution a l’étude de la faune du Mozambique. Voyage de M.P. Lesne

(1928-1929). Memdérias e estudos do Museu zooldgico da Universidade de Coimbra

(Série 1) 63: 3-10.

Pic, M. 1939. Malacodermes exotiques. Echange (hors texte) 55: 165-172.

Pic, M. 1945-48. Malachiidae nouveaux de |’Afrique australe. Entomologische Blatter fiir

Biologie und Systematik der Kafer 41-44: 62-64.

WitlMeR, W. 1953. 5. Beitrag zur Kenntnis der Malacodermata Afrikas. Proceedings of the

Royal Entomological Society of London (B. Taxonomy) 22 (5-6): 85-94.

Witrmer, W. 1956. 11. Beitrag zur Kenntnis der Malacodermata Afrikas (Col.). Entomo-

logische Arbeiten aus dem Museum Georg Frey 7 (3): 1072-1088.

COLOTES MIT EINFARBIG METALLISCHEN FLUGELDECKEN 69

WitrMER, W. 1962. Mission zoologique de l’I.R.S.A.C. en Afrique orientale (P. Basilewsky et

N. Leleup, 1957). LXXVII—Coleoptera Drilidae, Cantharidae, Malachiidae, Dasytidae et

Prionoceridae. Annales Musée royale de l'Afrique central (Zool.) 110: 259-270.

Witmer, W. 1987. Die Gattung Colotes Erichson in Athiopien, Somalia und Eritrea und

Beschreibung einer neuen Gattung (Coleoptera: Malachiidae). (26. Beitrag zur Kenntnis

der Fauna Afrikas). Annali del Museo civico di storia naturale di ‘Giacomo Doria’

86: 627-655.

Wittmer, W. 1989. Uber die Gattung Colotes Erichson (Coleoptera: Malachiidae) im siid-

lichen Afrika—Arten mit metallischen Fliigeldecken und hellen Seiten. (31. Beitrag zur

Kenntnis der Fauna Afrikas). Navorsinge van die Nasionale Museum Bloemfontein 6 (4):

111-149.

1 maar),

cn ny par tPA

yay

;

i} |

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc:

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: SO.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

Gee the ricure 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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

W. WITTMER

DIE IN AFRIKA VORKOMMENDEN

COLOTES ERICHSON MIT EINFARBIG

METALLISCHEN FLUGELDECKEN

(COLEOPTERA, MALACHIIDAE)

ANNALS

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes... .’

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 + #£Band

March 1992 Maart

Part 5 Deel

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

BARREMIAN REPRESENTATIVES OF THE

AMMONITE FAMILY ANCYLOCERATIDAE

Gib s/t

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

LAGER, 5-0), GED, 425, 8 enn), 6deS, 5, 7-9).

GG, tat), A=), & OCB, MD), MOOS), 1D, S, 7), G.t.,),

14(1-2), 15(4-5), 24(2, 5), 27, 31(1-3), 32(5), 33,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 128 8

Printed in South Africa by In Suid-Afrika gedruk deur

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

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D999

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,

SOUTH AFRICA.

BARREMIAN REPRESENTATIVES OF THE AMMONITE

FAMILY ANCYLOCERATIDAE GILL, 1871

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 40 figures)

[MS accepted 17 December 1990]

ABSTRACT

The Barremian representatives of the family Ancyloceratidae in Zululand are described.

These include Crioceratites (C.) yrigoyeni (Leanza, 1970), C. (C.) australis sp. nov., Acrioceras

(A.) zulu sp. nov. and Acrioceras sp. The genus Cryptocrioceras Aguirre Urreta, 1981 (type

species C. yrigoyeni (Leanza, 1970)), is regarded as a synonym of Crioceratites s.s., as are the

supposedly geographically separated Boreal Paracrioceras and Tethyan Emericiceras, respect-

ively. The temporal and geographic association of certain species of Acrioceras with

Crioceratites, as well as similarities in the early spiral whorls, suggests that they may represent

dimorphic pairs. Recognition of this type of dimorphism would simplify systematics of the

family Ancyloceratidae but, as yet, it cannot be proven unequivocally.

Two specimens from Zululand are similar to material from Patagonia described under the

genus Hemihoplites by Riccardi & Aguirre Urreta (1989).

CONTENTS

PAGE

IRE N ER OREN EY ETON Sas os cio re one slerec's wi sate touiectitcie Moe salsoae ae acetinas seemed se uaaiemeena es 12

OC AHIOMNOMSICCIMENS: «262 onc ac sessisns seanicnisee bata sinawdeeibcinsatin dame dsteatcoeacacte ates 72

ELGG. LOGE TSS cat Seade sca Gh Oa ene aa Sts ae Ey MARTE ne nee a sient am aa eR Laer 12

BYE BSIOMSTOMSPECIMENS cee. 22 8c seete ec ccaceect eae SGMRC Noob eis ne eves Sas cesleeiels sei 73

RSEUPUSRCMCCREIEMOL OD YO ce Sasa sa cdot 5c sited eden Ne disco Mele eninislsciy nae souialalslacie owen poe eckial 73

SUSE MM ARIE ANAC OM LOL OG Ya. i5 323 creo wicie nse ioeitisls bev Slap wi cite <cta ce ew cls ots coin s'oulaavesla aries 73

Discussion on shell shape, dimorphism and systematics in Crioceratitinae

AMEE NTIGVIOCCEALMAC cet e ee tee ccd niauiele wows ntas SS asnsionsee see sos aoa 129

OD GMRT RUSCH eee eRe aN a adres acacia arated warded es Seaneten ements 132

WO OMICS CINE INES nese cece acres Orc aiid nse Sauls Moblartamaudetina ak wecsGeaiiate 132

RRM CS prea ee eters seo ah Re ete ation hinid aie w Gala ab Hen maeameney sais Se cttentnaenaaees 32.

Ann. S. Afr. Mus. 101 (5), 1992, 71-138, 40 figs.

WZ ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

The majority of South African material belonging to the family Ancylocera-

tidae occurs in the Aptian of Zululand, and was described previously by Klinger

& Kennedy (1977). Barremian representatives of the family are here described

for the first time; these are all from the same (and only Barremian) outcrop,

locality 170 of Kennedy & Klinger (1975: 302), 30 m of cliff and gully sections

2 km north-west of the Mlambongwenya Trading Store, and include the oldest

ammonites known from the Zululand Group.

This locality has yielded rich ammonite faunas consisting of abundant Hete-

roceratinae (Klinger 1976; Klinger et al. 1984; Aguirre Urreta & Klinger 1986)

and Haplocerataceae (Kennedy & Klinger 1979). The palaeobiogeographic affi-

nities of these heteroceratine faunas with those of the Caucasus and southern

Patagonia are striking (Klinger 1990). Haploceratid faunas similar to those from

the Barremian of Zululand have recently been described from southern Pata-

gonia by Riccardi et al. (1987). The recent discovery at locality 170 of the sup-

posedly endemic Patagonian genus Hatchericeras Stanton, 1901, by Kennedy &

Klinger (1990) has further emphasized the close palaeobiogeographical affinities

of the Barremian ammonite faunas of Zululand with those of the Austral Basin

of southern Patagonia.

The ammonite family Ancyloceratidae is represented in the Barremian of

Zululand by numerous crioceratitid specimens, some of which are identical to

material described under the name Cryptocrioceras by Aguirre Urreta (1981),

and acrioceratid specimens best referred to the genus Acrioceras Hyatt, 1900.

Two specimens resemble material referred to Hemihoplites Spath, 1924, by

Riccardi & Aguirre Urreta (1989) and are herein tentatively referred to the

family Ancyloceratidae.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of the

material studied:

BMNH Natural History Museum, London

SAM South African Museum, Cape Town

FIELD LOCALITIES

Details of the field locality mentioned in the text are given by Kennedy &

Klinger (1975) and Aguirre Urreta & Klinger (1986); fuller descriptions of the

section are deposited in the Palaeontology Department of the Natural History

Museum, London; Geological Survey, Pretoria; and the South African Museum,

Cape Town.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 73

DIMENSIONS OF SPECIMENS

All dimensions given below are in millimetres. D = diameter, Wb = whorl

breadth; Wh = whorl height, U = umbilical diameter. Figures in parentheses are

ratios expressed as a percentage of the total diameter.

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916) reviewed by Kullmann &

Wiedmann (1970) is followed here. I=internal lobe; U=umbilical lobe;

L = lateral lobe; E = external lobe.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA Cuvier, 1797

Class CEPHALOPODA Zittel, 1884

Order AMMONOIDEA Zittel, 1884

Suborder ANCYLOCERATINA Wiedmann, 1966

Superfamily ANCYLOCERATACEAE Gill, 1871

Family Ancyloceratidae Gill, 1871

Systematics in the family Ancyloceratidae Gill, 1871, are confusing from

subfamilial, through generic, and down to specific level. The family Ancylocera-

tidae is generally taken to consist of the subfamilies Crioceratitinae Wright,

1952, and Ancyloceratinae Gill, 1871 (Wright 1952, 1957; Wiedmann 1962;

Thomel 1964; Kakabadze 1981), but may also include the subfamily Helicancyli-

nae Hyatt, 1894 (Casey 1961: 76; Wright 1981: 171; Aguirre Urreta 1986),

Protancyloceratinae Breistroffer, 1947 (Wiedmann 1973: 314; Immel 1978: 23),

Leptoceratinae Manolov, 1962 (=Leptoceratoidinae Thieuloy, 1966), and

Karsteniceratinae Immel, 1987 (p. 118), and tentatively Heteroceratinae

Hyatt, 1900 (Immel 1978: 23). Dimitrova took an extreme view (1970: 76) and

divided the superfamily Ancylocerataceae into three different families,

Himantoceratidae Dimitrova, 1970, Ancyloceratidae Meek, 1876, and

Protacrioceratidae Dimitrova, 1970; fortunately this view has found little

support.

Our material primarily addresses the relationship between the often

strongly ornamented Barremian crioceratitids referred to Emericiceras in the

Tethyan Realm, Paracrioceras in the Boreal Realm, and Cryptocrioceras in the

South Gondwanid Region (Zululand, Mozambique and southern Patagonia)

respectively, and Crioceratites s.s., and their relationship to the co-occurring

ancyloceratid genus Acrioceras. This latter relationship has bearing on the val-

idity of dividing the family Ancyloceratidae into the subfamilies Crioceratitinae

and Ancyloceratinae.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Subfamily Crioceratitinae Wright, 1952

Genus Crioceratites Leveillé, 1837

[= Emericiceras Sarkar, 1954; Paracrioceras Spath, 1924; Sornayites Wiedmann,

1962; Cryptocrioceras Aguirre Urreta, 1981]

Type species. Crioceratites duvalii Leveillé (1837: 313, pl. 22 (fig. 1a—b)).

Discussion

Typical representatives of the genus Crioceratites with loose, crioceratitid

coiling have not yet been found in Zululand. Instead, closely coiled forms, some

of which had been described under the generic name Cryptocrioceras in Pata-

gonia by Aguirre Urreta (1981) occur. In this discussion we concentrate mainly

on the systematic position and affinity of ‘Cryptocrioceras’ within the genus

Crioceratites.

The genus Cryptocrioceras was introduced by Aguirre Urreta (1981: 114)

for Mantelliceras yrigoyeni Leanza (1970: 249, pl. 42 (figs 1-2), pl. 43 (fig. 1))

(the type species) and Acanthoceras hoggi Leanza (1970: 244, pl. 40 (figs 1-3),

pl. 41 (fig. 1)), two species erroneously considered by Leanza to be members

of the family Acanthoceratidae, and of Cenomanian age. Freely translated,

the diagnosis of Cryptocrioceras given by Aguirre Urreta (1981: 114) is as

follows:

Of medium size, coiling is evolute with a dorsal zone of impression.

The whorl section changes from subtrapezoidal to subquadrate. The

ornamentation of the inner whorls consists of strong, trituberculate ribs

and intercalatories. Both types pass over the flanks on the whole

surface. On the outer whorls all the ribs are the same with smooth

bullae. The suture line consists of bifid saddles and trifid lobes, asym-

metrical; the lateral lobe is very large and well developed.

Examination of the suture line clearly shows that Mantelliceras yrigoyeni, of

which Acanthoceras hoggi is a synonym, is a typical quadrilobate member of the

suborder Ancyloceratina. Also, subsequent detailed stratigraphic collecting in

Patagonia has shown the age to be Barremian rather than Cenomanian. Crypto-

crioceras yrigoyeni is a member of the Hatchericeras patagonense assemblage

Zone, which occurs above the Favrella wilckensi assemblage Zone and below

the Colchidites assemblage Zone (see e.g. Riccardi 1984a, 1984b; 1988: 57,

table 15; Riccardi et al. 1987: 120; Riccardi & Aguirre Urreta 1989: 448).

Aguirre Urreta (1981: 116) referred Cryptocrioceras to the subfamily Crio-

ceratitinae, and considered the genus to have affinities with Crioceratites

Leveillé and Paracrioceras Spath.

Discovery in Zululand of ‘Cryptocrioceras’ yrigoyeni and of another (new)

species that could be referred to ‘“Cryptocrioceras’, raises doubts whether it

really is necessary to separate this group of crioceratitids from Crioceratites s.s.,

and whether the apparently endemic, Southern Hemisphere occurrence of

‘Cryptocrioceras’ is not a taxonomic artefact.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 75

Interpretation of the genus Crioceratites is beset with many difficulties, the

main problem being whether it is possible, and if so how, to reconcile the names

applied to Boreal and Tethyan crioceratitid populations. Monographic descrip-

tions from Boreal and Tethyan regions respectively (e.g. Neumayr & Uhlig

1881; Von Koenen 1902; Sarkar 1955; Thomel 1964) create the impression that

there are distinct, geographically separated, Boreal and Tethyan crioceratitid

populations. This impression is further strengthened by the apparent parallel

(?synchronous) phylogenetic development within Crioceratites leading to similar,

strongly tuberculate on the inner whorls, yet geographically distinct lineages,

e.g. Paracrioceras in the Boreal, and Emericiceras in the Tethyan Realm.

To this apparent synchronous development of geographically separate but mor-

phologically related forms, Cryptocrioceras from the South Gondwanid

Region (southern Patagonia, Zululand and southern Mozambique) can now be

added.

Crioceratites s.1. shows a wide range of morphological variation over its geo-

graphic and stratigraphic distribution during the Hauterivian and Barremian.

The genus is most probably interpreted too widely but, as yet, no satisfactory

subdivision has been proposed. Several species groups (e.g. Uhlig 1883; Sarasin

& Schondelmayer 1902; Kilian in Roman 1938: 352; Sarkar 1955; Thomel 1964;

Thieuloy & Thomel 1964; Immel 1978, 1979b) and genera or subgenera, e.g.

Emericiceras Sarkar, 1954, and Paracrioceras Spath, 1924, and phylogenetic

lineages (e.g. Wiedmann 1962) have been recognized within Crioceratites but,

unfortunately, many of these are either poorly defined or open to question. A

full discussion of these attempted subdivisions of the genus Crioceratites is

beyond the scope of the present investigation, and we limit ourselves to the

relevant sections that have bearing on the identity of ‘Cryptocrioceras’ .

Within the genus Crioceratites, the closest allies to “‘Cryptocrioceras’, espec-

ially the type species C. yrigoyeni, are the strongly trituberculate forms that

have been placed in Emericiceras Sarkar, 1954, in the Tethyan Realm and Para-

crioceras Spath, 1924, in the Boreal Realm, as was initially indicated by Aguirre

Urreta (1981: 117).

The status of Emericiceras and Paracrioceras varies according to different

authors. According to Sarkar (1954, 1955, 1977: 260) and mainly French authors

(e.g. Thomel 1964, 1981), Emericiceras is retained for the strongly tuberculate

Mediterranean (Tethyan) crioceratitids and Paracrioceras for the Boreal equiva-

lents (Sarkar 1977: 260). Wright (1957: L208) included Emericiceras in the

synonymy of Crioceratites with a question mark, but maintained Paracrioceras as

a separate genus.

Wiedmann (1962: 112) regarded both Emericiceras and Paracrioceras as

junior synonyms of Crioceratites and, in addition, included Pseudothurmannia as

a subgenus in Crioceratites s.1., a view followed here. The most significant aspect

of Wiedmann’s (1962) work was that he recognized a trend towards recoiling

(Fig. 1), and the concurrent reduction or simplification of ornamentation on the

outer whorls in some of the Crioceratitinae, leading to near-ammonitic end-

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

HAUTERIVIAN BARREMIAN

Fig. 1. Suggested phylogeny of major branches of Crioceratites to illustrate the trend towards

recoiling, according to Wiedmann (1962). 1. C. (C.). duvali. 2. C. (C.) nolani. 3. C. (C.)

emerici (= ‘Emericiceras’). 4. C. (Sornayites) paronai. 5. C. (C.) majoricensis. 6. C. (Pseudo-

thurmannia) angulicostatus. 7. Hemihoplites feraudianus. 8. C. (Pseudothurmanni) balearensis

(= ‘Balearites’). 9. C. (Ps.) balearis ibizensis. (Redrawn after Wiedmann 1962, text-fig. 35.)

members such as Pseudothurmannia, Hemihoplites and ‘Sornayites’ (Fig. 1), a

trend also recognized by Thieuloy (1964) (Fig. 3).

Rawson (1975: 279), commenting on Spath’s (1924: 279) reference to the

‘emerici’ group in his brief discussion on Paracrioceras, regarded Emericiceras as

a junior subjective synonym of Paracrioceras and suggested that the latter might

be regarded as a subgenus of Crioceratites (see also Rawson & Mutterlose 1983:

138; Rawson 1983: 498).

Immel (1978) followed Wiedmann’s (1962) views in regarding both Para-

crioceras and Emericiceras as synonyms of Crioceratites. Further, Immel (1978,

1979b), on the basis of the phylogenetic trend towards recoiling, was able to

recognize five Mediterranean and three Boreal species groups in Crioceratites

s.l. (Fig. 2).

Immel’s (1978, 1979b) (Fig. 2) grouping of Crioceratites is here used merely

as a convenient framework for discussing some of the affinities of the Gon-

dwanid crioceratitid ‘Cryptocrioceras’. A full discussion of the merits and

demerits of Immel’s species grouping is beyond the scope of the present

discussion.

In the Mediterranean region, the group of Crioceratites (C.) nolani bears

some resemblance to ‘Cryptocrioceras’. It is a long-ranging group, occurring

CRETACEOUS FAUNAS FROM SOUTH AFRICA a

Mediterranean Boreal

Tal Od Si

<|5 <

= =

: ae EE

<|5 is x

28 es —— o| 2

Ss s

(e)

way

©! ok

=|> e|/|<

> 9 >| =

a fa

£ !

a| = =D

rls OE

fe) Ss

= (e)

Fig. 2. Arrangement of Crioceratites species groups according to Immel (1978, 1979). 1. C. (C.)

nolani. 2. C. (C.) quenstedti. 3. C. (C.) majoricensis. 4. C. (C.) barremensis. 5. C. (C.) hilde-

siensis. 6. C. (C.) fissicostatus. 7. C. (C.) denckmanni. 8. C. (Pseudothurmannia) mortileti.

(Redrawn after Immel 1979, text-fig. 3.)

from the Lower Hauterivian to the Upper Barremian, which could make it a

potential ancestral group. In the group of Crioceratites (C.) nolani, the species

closest to ‘Cryptocrioceras’, especially its type species, are Crioceratites (C.)

emerici Leveillé (1837: 314, pl. 23 (fig. 1) (see Immel 1978: 35, table 4a) and

C. (C.) thiollierei (Astier, 1851: 18, pl. 5 (fig. 7) (see Immel 1978: 35, table 4b),

both from the Lower Barremian. These are typical ‘Emericiceras’. Crioceratites

(C.) thiollierei (including Emericiceras ottohaasi Sarkar (1955: 95, pl. 6 (fig. 5)

and Emericiceras thiollierei Ast. sp. var. multicostata Sarkar (1955: 93, pl. 4

(fig. 22), pl. 10 (fig. 6)) is closest to ‘Cryptocrioceras’ yrigoyeni. The style of

ornament, consisting of strong trituberculate ribs, separated by variable inter-

mediaries that themselves may bear tubercles, and the formation of loops

between the tubercles, are all features found in ‘Cryptocrioceras’ yrigoyeni. The

modes of coiling, however, are totally different. Coiling in Crioceratites (C.)

thiollierei is distinctly crioceratitid, loose, and some specimens uncoil in the

adult stage (Thomel 1964: 35, text-fig. 4) and may even form a recurved hook

(Immel 1978: 40) (Fig. 4).

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

L. NEOCOMIAN HAUTERIVIAN BARREMIAN

Fig. 3. Suggested phylogeny of major branches of Crioceratites to illustrate trend towards

recoiling according to Thieuloy (1964). 1. Protancyloceras kurdistanense. 2. Himantoceras tri-

nodosum. 3. Crioceratites (C.) nolani. 4. C. (C.) duvali. 5. C. (C.) villersianus. 6. C. (C.)

majoricensis. 7. C. (Pseudothurmannia) angulicostata. 8. C. (Emericiceras) emerici. (Redrawn

after Thieuloy 1964, text-fig. 3.)

As far as the strong trituberculation of the major ribs and the low number

of intermediaries on the inner whorls are concerned, ‘Cryptocrioceras’ finds its

closest match in the group of Crioceratites (C.) barremense (Kilian, 1895).

Species referable to this group include C. (C.) barremense (Kilian, 1895: 978;

see also Simionescu 1900: 14, pl. 1 (figs 4-5)), C. (C.) thomeli Immel, 1978

(= Crioceratites (Emericiceras) collignoni Thomel, 1964: 33, pl. 6 (fig. 4), text-

fig. 3), C. (C.) alpinus (d’Orbigny, 1850: 100; see also Cottreau 1937: 63, pl. 78

(figs 16-17)), and C. (C.) janus Thieuloy, 1979: 310, pl. 2 (figs 1-4)). The

Upper Barremian occurrence of this species group also matches that of ‘Crypto-

crioceras’. Unfortunately, very little is known of the overall (adult) shell of these

species. From the available figures, however, it does seem that at least C.

(C.) alpinus, C. (C.) thomeli and, to a lesser extent, C. (C.) barremense have

open, crioceratitid coiling, quite unlike that of ‘Cryptocrioceras’. Crioceratites

(C.) janus has the closest coiling of the group and shows the distinct change

from major trituberculate ribbing on the phragmocone to simple ribbing on the

body chamber. All the specimens of C. (C.) janus figured by Thieuloy (1979)

are small in comparison to the other species in this group, and we are not sure if

these are all juveniles or possibly micromorphs.

In the group of C. (C.) quenstedti (Ooster, 1860), C. (C.) binelli (Astier,

1851: 14, pl. 2 (fig. 2); see also Sarkar 1955: 57, pl. 2 (figs 4, 6); Thomel 1964:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 79

es?

cadaaienne Cee

Fig. 4. Crioceratites (C.) thiollierei (Astier, 1851). Specimen with fully developed uncoiled

stage; collections Staatliches Museum fiir Naturkunde, Stuttgart, cat. no. 6076, from Escrag-

nolles, France. Total length 95 cm. (Photograph supplied by courtesy of Dr G. Dietl.)

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

28, pl. 3 (figs 2—3)) is similar to Crioceratites (C.) australis sp. nov. in the very

short trituberculate stage on the inner whorls. Crioceratites (C.) binelli may also

develop constrictions on the outer whorls, as in Crioceratites (C.) australis, albeit

associated with major ribs. The ribs on the outer whorls of C. (C.) binelli

diverge from radially elongated umbilical tubercles. This type of ornament on

the outer whorls of C. (C.) binelli is unknown in ‘Cryptocrioceras’.

Although similarities in style of ornament can be found between ‘Crypto-

crioceras’, especially the type species ‘C.’ yrigoyeni, and some of the above-

mentioned Mediterranean species of Crioceratites, none of these possesses the

combination of close coiling plus the strong trituberculate ornamentation of the

early whorls of ‘*C.’ yrigoyeni.

Close coiling (albeit not with a dorsal zone of impression) associated with

strong ornamentation of the inner whorls is known in the Boreal ‘Paracrioceras’,

especially in the group of Crioceratites (C.) denckmanni (Miller, 1892) (sensu

Immel 1978: 63). This species group includes C. (C.) denckmanni (Miller, 1892:

18; see Von Koenen 1902: 261, pl. 11 (figs 1-2)) and C. (C.) stadtlaenderi

(Miller, 1892: 19; see Von Koenen 1902: 274, pl. 6 (fig. 6)), both from the

Middle Barremian of north-west Germany. Crioceratites (C.) denckmanni is also

known from Western Morocco.

Again, none of the above-mentioned Boreal crioceratitids has quite the

close coiling and inner trituberculate ornamentation of ‘Cryptocrioceras’.

A similar, though not identical, combination of these features is, however,

found in a very poorly known group of crioceratitids for which Wiedmann (1962:

140 footnote) proposed the subgenus Sornayites (type species Emericiceras

paronai Sarkar, 1955 = C. emerici in Parona 1898: 142, pl. 17 (fig. 6)). Accord-

ing to Wiedmann (1962: 140), Sornayites represents another branch within

Crioceratites in which recoiling had taken place (see also Fig. 1). It differs from

the other recoiled groups, however, in that trituberculation persists to the outer

whorls. Wiedmann (1962: 140 footnote) thought that ‘Sornayites’ originated in

the group of Crioceratites emerici. Apart from the type species, Wiedmann also

included Crioceratites edouardi (Honnorat-Bastide, 1889: 462, pl. 11) and Crio-

ceratites crioceroides (Torcapel, 1884, pl. 8 (fig. 1)) in Sornayites. (Sarkar (1955)

and Busnardo (1970) regarded C. edouardi and C. crioceroides as representa-

tives of Pseudothurmannia.)

Thomel (1965: 419) regarded the name Sornayites as superfluous and pre-

ferred to refer to it as the species group of C. edouardi—a view supported by

Immel (1978: 11). ‘Sornayites’ is poorly known from the Lower Barremian of

southern France, northern Italy, Romania, and southern USSR (fide Wiedmann

1962: 141 footnote). [Dr G. Delanoy, letter 4.01.1990, informed H.C.K. that

‘Sornayites’ edouardi may occur as early as Upper Hauterivian—which widens

the gap between Sornayites and Cryptocrioceras even further. He has also found

forms resembling Cryptocrioceras in a condensed sequence in the Upper Bar-

remian of south-east France.] The combination of close coiling and strong

trituberculation of ‘Sornayites’ are very similar to those of ‘Cryptocrioceras’,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 81

except that in the latter coiling tends to become even closer and there is a reduc-

tion of trituberculate ornamentation from the inner whorls outwards. It would

seem possible to regard ‘Sornayites’ in the Lower Barremian as ancestral to

‘Cryptocrioceras’ in the Upper Barremian but it cannot be proven. Even if this

suggested phylogenetic sequence is not wholly true, the features of ‘Cryptocrio-

ceras are by no means so unique as to merit generic separation from

Crioceratites s.1. The features of ‘Cryptocrioceras’ are as far removed from the

mainstream of Crioceratites as are ‘Emericiceras’ or ‘Paracrioceras’ .

In the discussion on possible dimorphism (see p. 129), we also note that

‘Cryptocrioceras’ occurs with Acrioceras in Zululand—a situation similar to that

found in Boreal and Tethyan Europe where ‘Emericiceras’ and ‘Paracrioceras’

also occur associated with Acrioceras. Studies on the subfamily Heteroceratinae

by Klinger et al. (1984), Aguirre Urreta & Klinger (1986), and Klinger (1990) all

point to strong palaeobiogeographic affinities between these heteromorph

ammonite faunas of the Caucasus, Zululand and Patagonia. With this in mind,

there can be no serious palaeobiogeographical objections to referring ‘Sorna-

yites’ and ‘Cryptocrioceras’ to Crioceratites s.s.

Even though it seems possible to connect ‘Cryptocrioceras’, via ‘Sornayites’ ,

with the genus Crioceratites, the morphological similarities and geographic prox-

imity with the predominantly Malagasy crioceratitid genus Menuthiocrioceras

Collignon, 1949, cannot be disregarded.

This apparently endemic genus was erected by Collignon (1949: 75) (as a

subgenus of Crioceratites) with type species Crioceras (Menuthiocrioceras)

lenoblei Collignon (1949: 76, pl. 11 (4) (fig. 1, 1a)) from the Upper Hauterivian

of Belohasifaka (Cercle de Sitampiky), Madagascar. In addition to the type

species, C. (M.) besairiei Collignon (1949: 77, pl. 10 (3) (fig. 1, la, 1b)) and

C. (M.) hourcgqi Collignon (1949: 78, pl. 12 (5) (figs 2, 2a, 3, 3a)), both from the

Upper Hauterivian of the same locality, were referred to Menuthiocrioceras.

According to Collignon (1949), special features of Menuthiocrioceras

included the whorl section, which is as wide as high to slightly wider than high,

the contiguous whorls with a dorsal zone of impression such as found in Lyto-

ceras, trituberculate ornamentation, and lack of an uncoiled section.

Collignon (1949) commented on the similarities between Menuthiocrioceras

and the Boreal crioceratitids from northern Germany, but also noted similarities

to Pseudocrioceras abichi (Batsevich & Simonovich, 1873). Finally, Collignon

(1949) also noted that the continuity of the ribbing over the venter were features

more ancyloceratid than crioceratitid.

In the same paper, Collignon (1949: 79, pl. 12 (5) (fig. 4, 4a)) described

Crioceras sp.? This specimen has very strongly trituberculate major ribs and we

are not sure why Collignon did not refer this specimen to Menuthiocrioceras as

well. This specimen is, to all appearances, identical to the inner whorls of

‘Cryptocrioceras’ yrigoyeni.

Later, Collignon (1962) described additional material from the Upper Hau-

terivian, which he referred to Menuthiocrioceras. These include M. colcanapi

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Collignon (1962: 78, pl. 202 (figs 918-919)), M. kuntzi Collignon (1962: 80,

pl. 206, figs 920-921)), M. ampakabense Collignon (1962: 82, pl. 207 (figs

922—923)), M. aontzyense Collignon (1962: 82, pl. 207 (fig. 924)), M. sornayi

Collignon (1962: 84, pl. 208 (figs 925-926), p. 86, pl. 209 (fig. 927)), M. sarkari

Collignon (1962: 88, pl. 210 (fig. 928)), and M. mahafalense Collignon (1962:

90, pl. 211 (fig. 929)). These specimens show the extreme variability of Menu-

thiocrioceras as interpreted by Collignon. Menuthiocrioceras is very large by

normal crioceratitid standards—with estimated phragmocone diameters of

30-40 cm (see e.g. M. sornayi in Collignon 1962, pl. 208 (fig. 925)), with orna-

ment varying from densely ribbed (e.g. M. kuntzi) to very faint as in M. sarkari.

Flared trituberculate ribs may occur in large fragments of M. sornayi (e.g.

pl. 209 (fig. 927)); others seem to have only faint trituberculate ribbing. Distinct

constrictions occur in M. ampakabense.

In addition, Collignon (1962: 20, pl. 431 (fig. 821)) also described Menu-

thiocrioceras sp. indet. from the Lower Valanginian of Antsalova.

The genus Menuthiocrioceras is known mainly from Madagascar, but was

also recently recorded from Indonesia (Skwarko & Thieuloy 1989). Crioceratites

diamantensis (Gerth, 1925) (see Gerth 1925: 115, pl. 1 (figs 3-4); Giovine 1950:

53, pl. 5 (fig. 1), pl. 6 (figs 2-3); Riccardi 1988 pl. 7 (figs 1-2)) from the Upper

Hauterivian—Lower Barremian of the Neuquén Basin of Patagonia, which also

has close coiling, may also belong to Menuthiocrioceras.

In view of the variation shown in Malagasy Menuthiocrioceras, it is difficult

on morphological grounds alone, to find any valid reasons for maintaining

‘Cryptocrioceras’ and Menuthiocrioceras separate. A problem in this connection

is that we do not have any material between the last occurrence of Menuthio-

crioceras in the Upper Hauterivian, and the first occurrence of “Cryptocrioceras’

in the upper part of the Lower Barremian or lower Upper Barremian—but this

is also the case between ‘Sornayites’ and ‘Cryptocrioceras’ and thus cannot alone

be regarded as a convincing argument.

The main problem in relating Menuthiocrioceras to ‘Cryptocrioceras’, or to

the rest of the crioceratitids for that matter, lies in the close coiling of the

former. The very earliest forms of Menuthiocrioceras in the Lower Valanginian

(see e.g. Collignon 1962: 20, pl. 181 (fig. 821)—this could possibly be an

uncoiled neocomitid and not a crioceratitid (Dr P. F. Rawson pers. comm.)),

are already closely coiled with a distinct dorsal zone of impression, as also are

the better known forms from the Upper Hauterivian. If Menuthiocrioceras is to

be considered as ancestral to Crioceratites—as tentatively suggested by Wied-

mann (1973)—or possibly to ‘Cryptocrioceras’, an intermediate group of

uncoiled forms has to be envisaged near the Hauterivian—Barremian boundary

to connect with loosely coiled Crioceratites gr. ex. C. emerici. As far as we

know, however, the general trend in the Crioceratitinae is towards recoiling (cf.

Wiedmann 1962; Thieuloy 1964; Figs 1, 3) rather than uncoiling. This would

seem to count against the possibility that Menuthiocrioceras may be regarded as

a direct ancestor of Crioceratites gr. ex. C. emerici (and subsequent ‘Emerict-

CRETACEOUS FAUNAS FROM SOUTH AFRICA 83

ceras’ and ‘Paracrioceras’). This leaves us with the possibility that Menuthio-

crioceras connects directly with ‘Cryptocrioceras’. This would not involve any

drastic uncoiling in the Lower Barremian, neither would it involve dramatic

change in ornament. It would, however, suggest that ‘Cryptocrioceras’ evolved

independently from Menuthiocrioceras, but in parallel with Emericiceras in the

Tethyan Realm and Paracrioceras in the Boreal Realm, from Crioceratites s.s.—

indeed a remarkable case of convergence.

Thus, although Menuthiocrioceras is morphologically very similar to

‘Cryptocrioceras’, it seems preferable from phylogenetic, palaeobiogeographic

and taxonomic points of view to separate them, and to regard Cryptocrioceras as

a true Crioceratites with affinities to the groups of ‘Emericiceras’ and ‘Paracrio-

ceras’, and possibly ‘Sornayites’. The exact systematic position of Menuthio-

crioceras within the Crioceratitinae remains an enigma and it is accorded separ-

ate generic status.

The ornamentation of ‘Cryptocrioceras’ resembles that of Pseudocrioceras

(type species Scaphites abichi Batsevich & Simonovich, 1873). Pseudocrioceras

was introduced by Spath (1924: 78) by merely indicating the type species. The

most comprehensive discussions are by Kakabadze (1978, 1981). Pseudocrio-

ceras has an uncoiled straight shaft after the planispiral section, which clearly

distinguishes it from ‘Cryptocrioceras’.

Relationships between ‘Cryptocrioceras’ and Pedioceras Gerhardt, 1897

(type species Pedioceras cundimarcae Gerhardt, 1897: 172, pl. 4 (fig. 7)), are

difficult to formulate because Pedioceras is poorly defined. We do not know if

the various species and figured specimens referred to Pedioceras by different

authors (e.g. Gerhardt 1897; Royo y Gémez 1945; Yenne 1949) in fact all

belong to the same genus. If Pedioceras is interpreted in terms of Gerhardt’s

(1897: 170-171) diagnosis and the type species P. cundimarcae only, it appears

that Pedioceras lacks differentiation into major and minor ribs on the inner

whorls, and is thus easily distinguished from ‘Cryptocrioceras’. The specimen

figured by Yenne (1949, pl. 102 (fig. 1)) appears to be a Crioceratites.

The genus Hoplitocrioceras Giovine, 1950 (type species (by monotypy)

Hoplitocrioceras gentilii Giovine, 1950, pl. 4 (figs 1-2)), from the Neuquén

Basin of Argentina, is a doubtful evolute crioceratitid, being based on a single

specimen lacking the inner whorls.

Occurrence

Crioceratites has a world-wide distribution through the Hauterivian to the

Upper Barremian. It is best documented from the Tethyan and Boreal regions

of Europe but is also known from the Caucasus (e.g. Drushchits & Kudryav-

tseva 1960; Kotetishvili 1970; Kakabadze 1981); in Africa from Morocco (Roch

1930), Egypt? (H. Douvillé 1916), Somalia? (Tavani 1942), Tanzania? (Spath

1930: 135), Madagascar? (Collignon 1949), Mozambique (Férster 1975), and

Zululand (herein); southern Patagonia, Argentina (Austral Basin) (Leanza

1970; Aguirre Urreta 1981); Neuquén Basin (Gerth 1925; Weaver 1931; Giovine

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

1950, 1952; Camacho & Riccardi 1978; Riccardi 1988); Colombia? (Etayo Serna

1968); Cuba (Myczynski 1977); Antarctica (Thomson 1971, 1974, 1980, 1983):

Japan (Obata et al. 1975; Obata & Ogawa 1976; Obata et al. 1976; Obata et al.

1982; Obata & Matsukawa 1984); Arctic Canada (Jeletzky 1964: 58); and ques-

tionably Indonesia (Skwarko & Thieuloy 1989).

Crioceratites (Crioceratites) yrigoyeni (Leanza, 1970)

Figs 5B, 6-16, ?30E-F, ?31F-J, 32A, 40

Acanthoceras hoggi Leanza, 1970: 244, fig. 40 (1-3).

Mantelliceras yrigoyeni Leanza, 1970: 249, fig. 42 (1-2), fig. 43 (1).

Crioceratites (Emericiceras) cf. thiollierei (Astier) Forster, 1975: 150, pl. 2 (fig. 2).

Emericiceras Kennedy & Klinger, 1975: 274.

?Emericiceras sp. Blasco, Nullo & Ploszkiewicz, 1980: 46, pl. 2 (fig. 8).

Cryptocrioceras yrigoyeni (Leanza) Aguirre Urreta, 1981: 117, pls 1-4.

Type

Holotype is the original of Mantelliceras yrigoyeni Leanza (1970, fig. 42

(1-2), fig. 43 (1)), housed in the collections of the Servicio Geoldégico Nacional,

Argentina, no. 12483, from the lower part of the Rio Belgrano Formation, at

Chorrillo Rivera, Santa Cruz Province, Argentina.

Material

BMNH Cs80003—4, C80039, C80055—80089, SAM-—PCZ8424a—b, PCZ8425a—

b, PCZ8430-—8431, PCZ8433—8435, PCZ8437-—8438, PCZ8445-8448, PCZ8453, all

from locality 170, cliff and gully sections 2 km north-west of Mlambongwenya

Trading Store on the north side of the stream, Makatini Formation, Barremian

I-II. Some of the specimens are precisely located within this cliff section: SAM-—

PCZ8447 is from Bed 17; PCZ8446 from Bed 18; PCZ8431 and PCZ8435 from

Bed 19; PCZ8437 from Bed 21; PCZ8445 from Bed 23; and PCZ8425a—b and

PCZ8453 from Bed 25.

Dimensions

Specimen D Wb Wh Wb/Wh U

C80066 45 15 (@Ci,7) Ss (200) 0,69 156333)

C80057 26 8,6 (33,0) 10 (38,5) 0,86 11 (42,3)

C80069 38 —(crushed) — — 15 (9,5)

C80060 35 11 (31,4) 135(67.2) 0,85 16 Gow)

C80055 26,6 11,5 (43,2) 11,4 (42,9) 1,00 11 (41,4)

Description

Our material of this species is very variable as far as coiling of the inner

whorls, nature and strength of ornament throughout, and inflation of the whorl

section are concerned. Coiling on the inner whorls is crioceratitid, with major

trituberculate and intermediary, tuberculate or non-tuberculate ribs, becoming

closer coiled on the outer whorls where the ribs become non-tuberculate and

uniform.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 85

Coiling on the inner whorls is very variable, ranging from open crioceratitid

with the whorls not touching at all, as in PCZ8447 (Fig. 30E), through slightly

less open with only the third whorl being detached from the preceding one, as in

PCZ8425b (Figs 16, 31F-I), to close crioceratitid, as in PCZ8433 (Fig. 31J),

PCZ8424a—b (Fig. 32A), C80056, and C80055 (Fig. 6A—B). The ventral spines

of one whorl generally touch a shallow impressed zone of the preceding whorl,

but the main surface of the dorsum and venter do not seem to have been in

contact. This appears also to be the case in later whorls, the dorsum of which is

also impressed, and which also shows what appear to be depressions to house

spines (e.g. Fig. 10D).

The whorl section varies from compressed to depressed, with the greatest

breadth at the umbilical spine. In general, ornament is strongest in inflated indi-

viduals and weakest in compressed ones; it will be convenient to describe the

species in these terms.

10mm

Fig. 5. A. Crioceratites (C.) australis sp. nov., BMNH C80054. B. Crioceratites (C.) yrigoyeni

(Leanza, 1970), BMNH C80057. Suture lines.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Crioceratites (C.) yrigoyeni (Leanza, 1970). A-B. BMNH C80055. x 1,5.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

i (Leanza, 1970). A, C, E. BMNH C80075.

D. BMNH C80066. All x 2.

.) yrigoyen

tes (C

Fig. 7. Criocerati

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

—

Fig. 8. Crioceratites (C.) yrigoyeni (Leanza, 1970). A, D-E. BMNH C80057.

B-C. BMNH C80073. F. BMNH Cs80004. All x 2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

TX IV “PpO00O8D HNN O-V “(COL6L “ezueaq) mMadosud (‘D) saiwso00uD

Gee aging,

‘6 “SI

ANNALS OF THE SOUTH AFRICAN MUSEUM

i (Leanza, 1970). A-B. BMNH C80058.

C-E. BMNH C80075. All x 2.

igoyen

tites (C.) yr.

Fig. 10. Criocera

CRETACEOUS FAUNAS FROM SOUTH AFRICA 91

Fig. 11. Crioceratites (C.) yrigoyeni (Leanza, 1970). A, E-F. BMNH Cs80081.

B-D. BMNH C80078. G-I. BMNH C0080. J. BMNH C80059. K-M. BMNH Cs0079.

A, E-F, J x 3; B-D, H-I x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

toceras (A.)

i (Leanza, 1970). BMNH C80060.

igoyen

Dxe2s

tes (C.) yr

Cx 4

(Leanza, 1970). BMNH C80003. C. Acr

iocerati

igoyeni

A-B xX 1

tites (C.) yr

iocera

Cr.

BMNH Cs80007. D. Cr

Fig. 12. A-B.

zulu sp. nov.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 93

Inflated forms: The innermost whorls are smooth and with rounded

whorl section but, at a diameter of c. 4-5 mm, the whorl section is distinctly

coronate, depressed as seen in C80055 (Fig. 14) and PCZ8425b (Fig. 16). There

are about 20-25 ribs per whorl. These arise on the umbilical wall and bear faint

conical to radially elongate tubercles on the inner edge of the umbilical wall.

Very strong, rounded tubercles occur at or slightly below midflank. When well

preserved, these are revealed as the bases of short, wedge-shaped spines;

from them, the ribs pass straight across the flanks and bear prominent ventro-

lateral tubercles that are the bases of sharp spines. Already at this diameter,

the lower two rows of tubercles are more prominent on some ribs than on

others.

With increase in size, the umbilical tubercles migrate to a lower-lateral pos-

ition, and the other two rows to an inner and outer ventrolateral position,

respectively. Differentiation in strength of ribbing becomes more prominent

with the addition of secondary ribs that are very variable both in form and

number, as shown by Figures 6-13.

At one extreme is a single, relatively weak rib bearing no, or only small,

ventrolateral spines. Others may bear umbilical/lateral and upper ventrolateral

tubercles or lower and upper ventrolateral tubercles only, with up to four minor

ribs between the major in some individuals. Furthermore, the numbers of sec-

ondary ribs may be either variable or constant in different individuals and has no

apparent taxonomic significance. Attendant on this proliferation of ribs is a

change in their direction from prorsiradiate to radial in some specimens, and

from straight to convex in all. Between tubercles, ribs are generally single but,

in some specimens where preservation is good, distinct looping is visible, and up

to four ribs may connect between the tubercles on the flanks. In some individ-

uals (e.g. Fig. 12D), very long, hollow septate spines survive, whereas in others

the ribs efface over the siphonal region.

At a diameter of c. 45 mm, tuberculation begins to decline markedly in

C80055 (Fig. 6) and there follows a stage of dense, relatively even, flexuous

ribbing with single, long ribs and some intercalated ribs, occasional weak,

radially elongated bullae low on the flank, no obvious tubercles (but a thickened

rib) at the ventrolateral shoulder, and ribs that broaden and weaken across the

siphonal area. Some specimens, e.g. C80060, remain strongly tuberculate to

greater diameters.

Compressed forms: We have juveniles only of this type, e.g. PCZ8433

(Fig. 31J), C80007 (Fig. 30F), and C80073 (Fig. 8B—C), and they show the same

basic pattern of ornament as already described, differing in a higher rib density,

generally weaker ribs, elongate bullae rather than conical umbilical/lateral

tubercles, weaker lower ventrolateral tubercles that are radially elongate,

although retaining strong ventral tubercles on the major ribs. Minor ribs tend to

bear only ventral tubercles or none, and ribbing is commonly very subdivided at

the ventral midline. In a few specimens the ribbing is particularly flexuous and

may be rursiradiate. Occasional pairs of ribs loop to the larger ventral spines.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

The suture line of juvenile specimens is shown in Figures 5B and 15. The

external lobe (E) is relatively small, the lateral lobe (L) large and asymmetri-

cally trifid, the umbilical (U) small and trifid. The saddle E/L is large and bifid,

and L/U and U/I smaller but also bifid.

Discussion

None of our specimens is as large as the Patagonian specimens figured by

Aguirre Urreta (1981), but the similar inner whorls leave no doubt that they all

belong to the same species. Our material is significant in that it shows the

extreme variation, not only in strength of ornament on the inner whorls, but

also the differing ratios of major and minor ribs during the course of ontogeny

within the same, and also between different specimens. None of our specimens

show the adult, body-chamber modifications seen in the Argentinian material

with the development of bullae on the ribs and even partial uncoiling in some

(e.g. Aguirre Urreta 1981, pl. 4 (fig. b)).

. Crioceratites (C.) australis sp. nov. described below (see p. 98), differs from

C. (C.) yrigoyeni in having a tuberculate stage of much shorter duration, and

the ornament is never as coarse as in the latter. Also, C. (C.) australis develops

distinct constrictions on the outer whorls; these have never been observed in

C. (C.) yrigoyeni.

The specimen described and figured by Collignon (1949: 79, pl. 12 (5)

(fig. 4, 4a)) from the Upper Hauterivian of Belohasifaka as Crioceras sp.?, is

indistinguishable from our material. As discussed above (p. 81), the affinities of

this specimen are obscure. It may be a fragment of Menuthiocrioceras.

Crioceratites (C.) yrigoyeni has ornament on the inner whorls similar to that

of some Mediterranean ‘Emericiceras’ species of some authors (especially Sarkar

1954, 1955; Thomel 1964, 1981). Thomel (1964: 30) recognized six species

groups in ‘Emericiceras’; of these, ‘E.’ gr. ex. barremense (Kilian, 1895) and ‘E.’

gr. ex. thiollierei (Astier, 1851) are closest to C. (C.) yrigoyeni. Both are Upper

Barremian. Our inflated forms of C. (C.) yrigoyeni with strong ornamentation

and few intermediary ribs are closest to ‘E.’ gr. ex. barremense, whereas the

compressed forms with more intermediate ribs are closest to ‘EF.’ gr. ex.

thiollierei.

Immel (1978: 52) included the following species in the group of Crioceratites

barremense: C. (C.) barremense (Kilian, 1895), C. (C.) thomeli Immel, 1978

(= Crioceratites (Emericiceras) collignoni Thomel, 1964: 33, pl. 6 (fig. 4), text-

fie. 3), and C. alpinus’ (d’Orbigny, 1850) (See Cottrean 19377 63a spines

(figs 16-17)). In all three species the trituberculate ribs are dominant over the

intermediaries, if present. However, as far as can be seen from the illustrations,

they all appear to have open crioceratitid coiling. One specimen figured by

Kakabadze (1981, pl. 15) as Paracrioceras barremense, however, has close

coiling as in C. (C.) yrigoyeni, and is difficult to separate satisfactorily on the

basis of the figured material, as is the specimen figured by Kakabadze (1981,

pl. 3 (fig. 4)) as Paracrioceras dolloi (Sarkar).

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 13. Crioceratites (C.) yrigoyeni (Leanza, 1970). A-B. BMNH Cs80065; x 1.

C-D. BMNH C80039; x 2.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

4 a!

Fig. 15. Crioceratites (C.) yrigoyeni

(Leanza, 1970). Suture lines. A. SAM-—

PCZ8433. B. BMNH C80055. A xX 8,2;

Bx:

Fig. 14. Crioceratites (C.) yrigoyeni

(Leanza, 1970). Reconstruction of whorl

section of BMNH C8s0055. x 2,1. C)

Fig. 16. Crioceratites (C.) yrigoyent

(Leanza, 1970). Whorl section of

SAM-PCZ8425b.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 97

Aguirre Urreta (1981: 119) also compared C. (C.) yrigoyeni with Criocera-

tites hoheneggeri (Uhlig, 1883) (in Thomel 1964: 32, pl. 6 (figs 2-3)), but the

coiling in this species is also more open crioceratitid than in C. (C.) yrigoyeni.

Crioceratites janus Thieuloy (1979: 310, pl. 2 (figs 1-4)), from the Upper

Barremian of the Southern Vercors Mountains and referred to the group of

C. barremense by Thieuloy, is very similar to C. (C.) yrigoyeni. It has strong

trituberculate ribs on the inner whorls, separated by single intermediaries,

but this mode of ornament soon gives way to slightly flexuous, sometimes

bifurcating ribbing. Coiling in C. janus is also relatively tight and comparable

to that of C. (C.) yrigoyeni, and not at all as loosely coiled as in typical

C. barremense, C. thomeli or C. alpinus. Unfortunately, all available figured

specimens of C. janus are small, and we do not know if they are juveniles or

small adults.

Crioceratites (C.) thiollierei (Astier, 1851: 18, pl. 5 (fig. 7)) resembles

our compressed forms as far as the ornament is concerned, but has distinct

crioceratitid coiling (see e.g. Thomel 1981: 49, fig. 85). According to Thomel

(1964: 30) the species in the group of C. thiollierei grow to very large size—

up to 1m in diameter, and may uncoil (Thomel 1964: 35, text-fig. 4)

and even form a recurved hook, as shown in a large specimen housed

in the collections of the Staatliches Museum fir Naturkunde, Stuttgart

(Fig. 4).

The ornamentation on the phragmocone of Pseudocrioceras, best known

from the Caucasus (see e.g. Rouchadze 1933, 1938; Kakabadze 1978, 1981) is

very similar to that of C. (C.) yrigoyeni but, as pointed out above and by Kaka-

badze (1978, 1981), the genus has distinct ancyloceratid uncoiling.

Unfortunately, the material from Alexander Island, Antarctica, initially

described and figured by Thomson (1974: 12, pl. 2e, text-fig. 4a) as Emericiceras

(?) sp., and subsequently as Paracrioceras (Thomson 1983: 410, text-fig. 2), is

too poorly preserved for definite identification, but it does superficially resemble

C. (C.) yrigoyeni.

The early whorls of C. (C.) yrigoyeni are indistinguishable from those of

coarsely ornamented forms of Acrioceras zulu sp. nov. described below (see

p. 110). In fact, small, criocone inner whorls could be referred to either genus

(see e.g. Figs 30-31). What is more, they occur together. This association and

its implications are discussed below.

The specimen described and figured from the Neuquén Basin of Patagonia

as Paracrioceras cf. P. emerici Lev. by Giovine (1950: 59, pl. 5 (fig. 5)) is too

poorly preserved for definite identification.

Occurrence

Lower—Middle Barremian of Patagonia, Hatchericeras patagonense

assemblage Zone. In Zululand, C. (C.) yrigoyeni occurs below the zone of

abundant Colchidites vulanensis australis Klinger et al., 1984, Barremian I.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crioceratites (Crioceratites) australis sp. nov.

Figs 5A, 17-24

Type

Holotype is BMNH C80037 from Bed 2 at locality 170, cliff and gully sec-

tions, 2 km north-west of Mlambongwenya Trading Store, on the north side of

the stream, Makatini Formation, Barremian I.

Material

Eighteen specimens, BMNH C80035, 80038, 80040-80054, and SAM-—

PCZ8452, all from the same locality as the holotype. SAM—PCZ8452 is from the

basal bed, Bed 1, of the cliff section.

Description

Loosely coiled but with the later whorls at least in contact, the dorsal

impressed zone being distinct, if shallow. The whorl section is typically com-

pressed, trapezoidal, with a broad dorsum, broadly rounded umbilical

shoulders, flattened convergent flanks, rather narrowly rounded ventrolateral

shoulders, and flattened venter. The greatest breadth is close to the umbilical

shoulder.

At the smallest diameter visible (30 mm), ornament consists of distant

umbilical bullae giving rise to groups of two to three ribs with three or four single,

non-bullate ribs between. All ribs are delicate, narrow, rursiradiate and rather

crowded. They flex backwards and are feebly convex to straight on the flanks,

broadening towards the ventrolateral shoulder. The ribs, which arise from bullae,

are commonly reunited at a conical, lower ventrolateral and clavate upper ventro-

lateral tubercle; other ribs bear rounded tubercles on the ventrolateral edge, and

all ribs are interrupted on the venter at a smooth, siphonal band.

Only one specimen, the holotype C80037 (Figs 17, 18A—C), shows the

tuberculate early stage described above. In other specimens, only the second

growth stage is visible; here the whorl section is more compressed (whorl

breadth to height ratio is 0,7). There are six to nine ribs in a distance equal to

the whorl height. These arise at the umbilical shoulder as the feeblest of bullae.

They are recti- to feebly rursiradiate, and broaden markedly from the umbilicus

towards the ventrolateral shoulder. Varying from straight to flexuous, they ter-

minate in transversely elongated feeble tubercles on either side of a smooth

siphonal area up to a whorl height of 15-17 mm, and are mostly long and

simple. There is, however, a range of individuals of uniform to variable rib

strength and direction. Variation is also pronounced in the shape of the ventral

tubercles (they are sometimes conical), and in the prominence and distinctness

of the smooth siphonal area. Both tubercles and smooth zone become less

prominent as size increases, disappearing at different diameters from individual

to individual, and generally associated with a rounding of the venter

(Fig. 19A—C).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 99

In our large specimens, ribs may branch conspicuously (Fig. 23) and gener-

ally decline in prominence as size increases. Associated with this decline is the

appearance of broad, deep, flexuous, prorsiradiate constrictions, bounded in

front and behind by parallel ribs, which give even fragments an immediately

recognizable and characteristic appearance (Figs 17, 20B, 21E, 22A, 23A, 24B).

The suture is intricately subdivided, with a large, deeply incised external

lobe (E) in which there is a large median element. The lateral lobe (L) is large

and asymmetrically trifid; the umbilical lobe (U) is smaller but also asymmetri-

cally trifid; and the internal lobe (1) is intricately subdivided. The saddle E/L is

Fig. 17. Crioceratites (C.) australis sp. nov. Holotype, BMNH C80037. x 1.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

large and asymmetrically bifid with deep incisions; L/U is smaller but also asym-

metrically bifid (Fig. 5A).

Discussion

The (ontogenetically) short trituberculate stage and the appearance of

prominent constrictions on the outer whorls is characteristic of this species, and

easily separates it from Crioceratites (C.) yrigoyeni, as well as from the majority

of other Crioceratites species.

Amongst the Mediterranean species, C. (C.) binelli (Astier, 1851) (see e.g.

Sarkar 1955: 57, pl. 2 (figs 4, 6); Thomel 1964: 28, pl. 3 (figs 2-3); Immel 1978:

43, table 6a) bears some resemblance to C. (C.) australis in the relatively tight

coiling, short duration of the trituberculate stage on the inner whorls, and in the

presence of occasional constrictions. In C. (C.) binelli, however, the constric-

tions are associated with thickened ribs that may bear umbilical and ventral

tubercles.

Amongst the Boreal species, C. (C.) australis can be compared with C. (C.)

fissicostatus (Roemer, 1841) (see e.g. Von Koenen 1902: 233, pl. 12 (fig. 2),

pl. 22 (figs 1-2)) and especially C. (C.) fissicostatus Roemer var. minor von

Koenen (1902: 236, pl. 23 (figs 1-2) (=?C. (C.) aequicostatus (von Koenen,

1902) fide Immel 1978: 63)) as far as the fine ornamentation on the inner whorls

is concerned. As far as can be seen, however, the Boreal species show distinct

uncoiling on the outer whorls.

Crioceratites diamantensis Gerth, 1925 (p. 115, pl. 1 (figs 3-4); Giovine

1950: 53, pl. 5 (fig. 1), pl. 6 (figs 2-3); Riccardi 1988, pl. 7 (figs 1-2)), from the

Upper Hauterivian—Lower Barremian of the Neuquén Basin of Patagonia, bears

some resemblance to Crioceratites (C.) australis in the close coiling and presence

of constrictions on the outer whorls. An inner, trituberculate stage has not yet

been found in C. diamantensis and we are not quite sure of its taxonomic pos-

ition. We suspect that it may be closer to Menuthiocrioceras than to Crioceratites

S.S.

Occurrence

As yet, C. (C.) australis is only known from the first division of the Bar-

remian of Zululand.

Subfamily Ancyloceratinae Gill, 1871

Genus Acrioceras Hyatt, 1900

[= Aspinoceras Anderson, 1938; Mesocrioceras Breistroffer, 1952;

Protacrioceras Sarkar, 1955; Paraspinoceras Breistroffer, 1952;

Subaspinoceras Thomel et al., 1987]

Type species.. Ancyloceras tabarelli Astier (1851: 19, pl. 7 (fig. 9)), by orig-

inal designation of Hyatt (1900: 588).

101

CRETACEOUS FAUNAS FROM SOUTH AFRICA

. nov. A-C. BMNH C80037, the holotype.

D-F. BMNH C80041. All x 1.

is sp

. 18. Crioceratites (C.) austral.

Fig

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. Crioceratites (C.) australis sp. nov. A-B. BMNH C80052. C-D. BMNH C80038.

E. BMNH C80035. All x 1.

103

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘TX TV

‘nynz (‘y) svaavouoy paqqu Ajauy Jo yUourseI] JO uonIsod sajesIpul MOITY ‘90080 HNN

9-V ‘Aou

‘ds sy

DAISND (°D) $a]NVsVIOUD

07 ‘314

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 21. Crioceratites (C.) australis sp. nov. A-B. BMNH C80040. C. BMNH C80041.

D. BMNH C80049. E. BMNH C80035. Note dorsal zone of impression in A and distinct con-

strictions in E. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 105

Fig. 22. Crioceratites (C.) australis sp. nov. A. BMNH C80046. B. BMNH C80047.

C. BMNH C80038. Note constriction in A and dorsal zone of impression in C.

All x 1.

B-C. Inner whorls of

A. BMNH C80047.

ANNALS OF THE SOUTH AFRICAN MUSEUM

nov.

BMNH C8s0047. D. BMNH C8s0040. All x 1.

Fig. 23. Crioceratites (C.) australis sp.

106

CRETACEOUS FAUNAS FROM SOUTH AFRICA 107

Discussion

As in the case of Crioceratites, recent reviews of the group of small ancylo-

ceratids, variously referred to as Acrioceras, Aspinoceras, Paraspinoceras,

Subaspinoceras, Mesocrioceras or Protacrioceras by Sarkar (1955), Wiedmann

(1962), Thomel (1964), Breskovski (1966), Dimitrova (1967), Kakabadze

(1981), Thomel et al. (1987, 1990), and Vasicek & Michalik (1988), confuse

rather than clarify taxonomic issues.

The most comprehensive discussion of the genus in the wider sense (as

Acrioceras s.1.) is by Sarkar (1955) and, recently, in the restricted sense (as

Acrioceras s.s.) is by Thomel et al. (1990).

Sarkar (1955) believed that Acrioceras could be divided into four sub-

genera, based primarily on the mode of coiling, i.e. acrioceratid or aspinocera-

tid, and whether major ribs, if present, are or are not trituberculate. His scheme

is as follows:

1. Acrioceras s.s Hyatt, 1900 (type species Ancyloceras tabarelli Astier, 1851:

19, pl. 7 (fig. 9))—coiling acrioceratid, major ribs trituberculate.

2. Acrioceras (Paraspinoceras) Breistroffer, 1952 (type species Ancyloceras pul-

cherrimum dOrbigny, 1840: 495, pl. 121 (figs 3-7))—coiling acrioceratid,

major ribs, if present, non-tuberculate; crozier Hamulina-like.

3. Acrioceras (Aspinoceras) Anderson, 1938 (type species Aspinoceras hamlini

Anderson, 1938: 207, pl. 60 (figs 1-2))—-coiling aspinoceratid, major ribs non-

tuberculate.

4. Acrioceras (Protacrioceras) Sarkar, 1955 (type species Ancyloceras ornatum

d’Orbigny, 1850: 101)—coiling aspinoceratid, major ribs trituberculate.

This classification has been modified to a greater or lesser extent by various

subsequent authors.

Casey (1960: 18) did not regard Aspinoceras as an ancyloceratid, but rather

saw it as a morphological type from which the *. . . Crioceratitidae and Hetero-

ceratidae may have diverged’. Wiedmann (1962: 143) only accepted Acrioceras

s.s. and A. (Aspinoceras), and regarded Ancyloceras meriani Ooster (1860: 35,

pl. 39 (figs 1-7)) as a connecting link between these two subgenera. This view

was followed by Immel (1978: 73) who pointed out that Hamites phillipsi Bean

(in Phillips 1829, pl. 1 (fig. 30)), the type species of Spath’s (1924: 78) genus

Hoplocrioceras, is a typical example of Aspinoceras. According to the rules

of priority, A. (Aspinoceras) should be regarded as a junior synonym of

A. (Hoplocrioceras). This view was followed by Kakabadze (1981) and recently

also by Vasicek & Michalik (1988).

Thomel (1964) followed Sarkar’s (1955) views, and later Thomel et al.

(1987) added the genus Subaspinoceras (type species Ancyloceras mulsanti

Astier, 1851: 18, pl. 6 (fig. 8)) for the European equivalents of the North

American genus Aspinoceras.

Recently, Thomel et al. (1990) have clarified some of the problems sur-

rounding Acrioceras s.s. on the basis of a large collection of specimens from the

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 24. Crioceratites (C.) australis sp. nov. A. BMNH C80052. B. BMNH C8s0035. Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 109

south-east of France. They are fully aware of the extreme intraspecific variation

in Acrioceras, and that many of Sarkar’s (1955) names are superfluous. Instead,

they only consider six names sufficiently well understood to qualify as valid

species in the biological sense. These include Acrioceras meriani (Ooster),

A. monopujaae Sarkar, A. ramkrishnai Sarkar, A. sarasini Sarkar, A. tabarelli

(Astier), and A. terveri (Astier).

Thomel et al. (1990) did not regard Protacrioceras as ancestral to Acrio-

ceras; instead, they (p. 97) derived Acrioceras in the Upper Hauterivian (Zone

of Sayni) from the group of Paraspinoceras pulcherrimum, via Acrioceras

meriani. Two distinct acrioceratid lineages can be traced from A. meriani: (1)

A. meriani—A. tabarelli—A. terveri, and (2) A. sarasini-A. ramkrishnai—A. mono-

pujaae. In the former there is a distinct trend towards increase in size of the

recurved hook in relation to the criocone whorls, but coiling remains more or

less acrioceratid. In the latter lineage, especially in the Upper Barremian group

of A. monopujaae, the shaft becomes incurved and the hook open, resulting in

an aspinoceratid ‘Protacrioceras’-type of coiling.

Dimitrova’s (1967) interpretation of Protacrioceras is unusual. Protacrio-

ceras tzankovi (Dimitrova, 1967: 55, pl. 21 (fig. 1, 1a), pl. 22 (fig. 1)) is a

gigantic ancyloceratid and certainly does not belong here in the sense of Sarkar

(1955).

Epacrioceras Egoian, 1974 (type species Epacrioceras rarum Egoian, 1974:

225, figs 1-2), is an Upper Aptian homoeomorph of Acrioceras, thus far only

known from the Western Caucasus. Ornament is virtually identical to that of

Acrioceras, but the suture has a distinct bifid lateral lobe (L) in contrast to the

trifid condition in Acrioceras.

Rawson (1975) suggested that Acrioceras and Emericiceras—Paracrioceras

might be dimorphic. Details of this are discussed below (see p. 129).

Our Barremian material from Zululand shows that details of ornamenta-

tion are very variable but that, at specific level at least, acrioceratid and

aspinoceratid coiled forms may be quite distinct, albeit possibly contemporary.

All our specimens are tuberculate at some stage or other and are thus referred

to Acrioceras S.s.

Occurrence

Acrioceras s.1. ranges from the Upper Hauterivian to the Upper Barremian,

and occurs in both the Tethyan and Boreal realms. It is best known from West

and Central Europe, but has also been recorded from the United States of

America (California and Oregon) (Anderson 1938; Murphy 1975), Canada

(Jeletzky 1964), Colombia (Etayo Serna 1968), Madagascar (Collignon 1962),

Antarctica (Thomson 1974), Japan (Obata et al. 1976; Obata & Ogawa 1976),

and Indonesia (as Hoplocrioceras—Skwarko & Thieuloy 1989).

Reports of Acrioceras from the Lower Aptian (cf. Wright 1957: L211;

Drushchits & Kudryavtseva 1960: 294; Kakabadze 1981: 96) have to be viewed

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

with suspicion, as is discussed below (p. 129) in the section on dimorphism.

Records from Australia (Kakabadze 1981: 96) cannot be confirmed.

Acrioceras (Acrioceras) zulu sp. nov.

Figs 25-28, 29A-B, ?C, 30A-D, ?E-F, 31A-D, ?E, ?F-J, 32B—C, 33-34, 39

Type. Holotype is BMNH C80009 from locality 170, cliff and gully sections

2 km north-west of the Mlambongwenya Trading Store, on the north side of the

stream, Makatini Formation, Barremian I.

Material

Fifteen specimens: BMNH C80005a—c, C80006-—80008, SAM-—PCZ8423,

PCZ8426, PCZ8428—-8430, PCZ8440-1, PCZ8443, PCZ8449, PCZ8451 and

PCZ8666, all from the same locality as the holotype; PCZ8441 is from Bed 25;

PCZ8423 is from Bed 1.

Description

The shell is small, ancyloceratid, with an initial variably coiled spiral

section, followed by a straight shaft. None of our specimens possesses a com-

plete crozier, but PCZ8423 (Fig. 33A—B) shows part of a short hook, as does

C80009 (Fig. 34), suggesting an overall length of c. 50 mm at maturity.

The earliest developmental stages are best seen in C80005c (Fig. 28A), and

PCZ8443 (Fig. 30A—D). The former shows the globose protoconch 0,5 mm in

diameter, in close contact with half a whorl of phragmocone. Coiling is very

variable —open with the early whorls not in close contact in C80005c (Fig. 28A)

and tight with the whorls in close contact in PCZ8443 (Fig. 30A—D). The whorl

section of the early spiral varies from compressed in C80005c to depressed in

PCZ8443 (Figs 28A, 30A—D). The dorsum is slightly concave.

Ornament on the spire and shaft is extremely variable. At one extreme, as

represented by C80005c (Fig. 28A), a compressed individual, there are numer-

ous (70-80) fine, delicate, flexuous ribs per whorl. Mere striae on the dorsum,

they strengthen across the umbilical wall (where they are distinctly rectiradiate)

and are concave across the umbilical shoulder. Most ribs are single, although a

[ U

10mm s\

Fig. 25. Acrioceras (A.) zulu sp. nov. Suture line. SAM—PCZ8668.

7% X HV ‘Cl pure > ur yey) YM vy UT BuTTI09

asoo] aavdwioD “q-e¢0008) HNN °C 850008) HNN ‘O “9500089 HNING ‘A °S00089 HNWd ‘Vv ‘sou ‘ds nynz (‘y) spsavouay “97 “BIq

Ebt

CRETACEOUS FAUNAS FROM SOUTH AFRICA

ANNALS OF THE SOUTH AFRICAN MUSEUM

‘Cc X Ylog

‘600080 HNING ‘edAiojoH{ ‘g ‘JUOWRUIO s]e[NOIOGNIIq ‘YeoM YUM susUtIDedS ‘q—eCGQDD8D HNINA “V “Aou ‘ds njnz (‘y) spsav0uoy “LZ “814

Siaaitendiindneeennnemnne

CRETACEOUS FAUNAS FROM SOUTH AFRICA

iil)

D-E xX 3,5.

Fig. 28. Acrioceras (A.) zulu sp. nov. A. BMNH C80005c. B-C. BMNH C80005b. D-E. BMNH C80005b. A-C x 2

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 29. A-B. Acrioceras (A.) zulu sp. nov. Coarsely ornamented, sextuberculate

shaft. SAM-—PCZ8666. C. Coarsely ornamented criocone whorls that could

belong to either Acrioceras (A.) zulu sp. nov. or coarsely ornamented form of

Crioceratites (C.) yrigoyeni (Leanza, 1970). SAM-—PCZ8667. Both x 1.

few branch into two or three at the shoulder. They are slightly prorsiradiate on

the flanks, feebly convex on the inner flank, concave on the outer flank, and

quite markedly projected on the ventrolateral shoulder. Narrow at the umbili-

cus, the ribs thicken and strengthen across the flanks. On some ribs there is a

slight indication of strengthening at the lower ventrolateral position, although no

clearly defined tubercle develops. In contrast, ventral, transversely elongated

tubercles are well developed, and some bear indications that they are in fact the

bases of short spines. Tubercle strength varies from mere swellings to marked,

sharp excrescences. As size increases, there is a tendency towards a pattern of a

strongly tuberculate rib followed by three to five weakly tuberculate ribs. In a

few cases a pair of flank ribs connect to one of these more pronounced

tubercles. All ribs are broad and low and feebly convex across the venter.

Less compressed forms, such as C80005a (Fig. 26C), are more robustly

ribbed and with a distinct rursiradiate style and stronger suggestion of a lower

ventrolateral tubercle. These specimens form a link to the coarsely ribbed form

illustrated by C80009 (Fig. 27B) and PCZ8443 (Fig. 30A—D). In these, differen-

tiation of ribbing occurs—the major ribs bearing variably developed umbilical

tubercles, very strong ventrolateral and strong ventral tubercles.

This range of variation is also seen on the shafts of our specimens. In the

compressed variety, ribbing is dense and prorsiradiate, and only ventral

tubercles, linking two to three ribs, occur. On the more robustly ornamented

specimens (Fig. 29A—B), differentiation of ribbing is more pronounced, and dis-

tinct umbilical, ventrolateral and ventral tubercles are developed. In some, e.g.

PCZ8441 (Fig. 33C—D) every rib on the shaft is tuberculate.

CRETACEOUS FAUNAS FROM SOUTH AFRICA US)

Discussion

As here interpreted, this species is very variable, so much so that, in terms

of ornament on the spiral whorls, we can speak of compressed, bi- or incipiently

quadrituberculate, finely ribbed forms, e.g. C80005a—c (Fig. 26), ‘normally’

ornamented (C80009, Fig. 27B), and coarsely ornamented forms (PCZ8666,

Fig. 29A—B). As far as the coiling of the spiral section is concerned, we can

speak of tightly coiled ancyloceratid (C80005a, Fig. 26C), and loosely coiled

aspinoceratid or crioceratitid forms (C80005c (Fig. 26A) and PCZ8451). As far

as Ornament on the shaft is concerned, we can distinguish bi- (C80005c,

Fig. 28A), quadri- (C80008) and sextuberculate (PCZ8666, Fig. 29A—B) forms.

Variation is so extensive that virtually every specimen merits description as

a separate species, if species concepts of most previous workers on this genus

were applied (which is more or less what Sarkar (1955) did in his description of

the French material).

Acrioceras zulu clearly belongs to the group of Acrioceras tabarelli (Astier,

1851: 19, pl. 7 (fig. 9)), but it connects morphologically with Paraspinoceras pul-

cherrimum (d’Orbigny, 1840: 495, pl. 121 (figs 3-7)) via the compressed, finely

ribbed, bituberculate forms.

Acrioceras tabarelli is a very variable species, as noted by Thomel (1964: 41

‘d'une forme tres plastique’) and Thomel ef al. (1990). Sarkar (1955: 102-112)

tried to overcome this problem by naming virtually every individual specimen,

e.g. Acrioceras tabarelli var. sarasini, var. uhligi, Acrioceras cf. tabarelli sp.

forme nrs 1—8. Thomel et al. (1990) provided excellent figures to illustrate the

wide extent of intraspecific variation in A. tabarelli. Even so, their extensive use

of open nomenclature in the captions to their excellent figures clearly shows how

difficult it is to put a definite specific or subspecific name to every specimen,

even if it is completely preserved. Apart from two formal subspecific names,

A. tabarelli tabarelli and A. t. sugrivai, Thomel et al. (1990) used the following

combination of names in the explanation to their figures:

Acrioceras aff. tabarelli (Astier)—pl. 2 (fig. 11).

Acrioceras gr. tabarelli (Astier)—pl. 3 (fig. 1), pl. 4 (figs 4-6), pl. 8 (fig. 7),

plots: 2).

Acrioceras gr. tabarelli (Astier) aff. uhligi Sarkar—pl. 3 (fig. 2).

Acrioceras entre tabarelli (Astier) et sarasini (Sarkar)—pl. 3 (figs 7-8), pl. 5

(fig. 6), pl. 9 (figs 3-4).

Acrioceras entre sarasini Sarkar et tabarelli (Astier)—pl. 4 (fig. 1), pl. 6

(figs 4-5).

Acrioceras entre tabarelli (Astier) et terveri (Astier)—pl. 4 (figs 8-9).

Acrioceras entre tabarelli (Astier)—terveri (Astier) et uhligi Sarkar—pl. 4

(fig. 10).

Acrioceras gr. tabarelli (Astier) comparer 4 Acrioceras cf. tabarelli Astier formes

4 et 5 de Sarkar—pl. 10 (fig. 4).

Acrioceras sp. entre gr. tabarelli (Astier) et uhligi Sarkar—pl. 11 (fig. 6).

ANNALS OF THE SOUTH AFRICAN MUSEUM

116

‘¢ xX dG: X O'7 X d-A “d-V “200080 HNN ‘A “Lep8ZOd-WVS ‘A ‘Aou “ds njynz ("y) spsaoouny

1O (OL6I “ezuRaT) 1MakosUA (*D) SajvsaIOUD JO S[IOYM BUOdOIID ALR “J-A “€pPYZOd-WVS ‘Aou ‘ds njnz (‘y) svsavousy “q-V ‘0¢ “34

117

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘OSCV8ZOd-WVS I

‘O€P8SZOd-WVS ‘Aou “ds njnz (‘y) spdao0uoy¢ “A ‘Ory8ZOd-WVS ‘Aou

‘TX f H-O'b X I A-V ‘€€r8ZOd-WVS ‘f

‘aou ‘ds njnz (‘Y) spsav0ldy JO sjIoyM ouOdOTIOR ApIeA ‘f-y

‘ds njnz (‘Y) spdav0uoy JO sjioyM suOsOLIO ALIeq “q-V ‘IE ‘sly

A ‘(OL6L ‘ezueay) makosiud (Dp) saj1jwv1a90lUD 10

ANNALS OF THE SOUTH AFRICAN MUSEUM

118

Toodee < O-V.

‘6998ZOd-NVS ‘Aou ‘ds njnz (“y) svsav0uoy “D-G “9-P/TP8ZOd-WVS ‘(OLOT ‘ezURIT) Juadosud (‘D) saiNMsav0UID “YW °ZE “BIA

CRETACEOUS FAUNAS FROM SOUTH AFRICA 119

Fig. 33. Acrioceras (A.) zulu sp. nov. A-B. SAM—PCZ8423, specimen with part of final hook

preserved. C-D. SAM-PCZ8441. Part of shaft with prominent sextuberculate ornament.

Both xX 2.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 34. Acrioceras (A.) zulu sp. nov.

Silicone mould of holotype,

BMNH Cs80009.

None of the European specimens has quite as coarse ornamentation as our

most strongly ornamented specimens; at the other extreme, none of the Zulu-

land specimens is as finely ribbed and non-tuberculate as Paraspinoceras

pulcherrimum. Acrioceras (A.) zulu ranges morphologically from coarsely orna-

mented, sextuberculate forms, much more coarsely ornamented than any

specimens of A. tabarelli known, to finely ornamented, bi- or incipiently quadri-

tuberculate forms, although never as finely ornamented as Paraspinoceras

pulcherrimum.

According to Thomel et al. (1990), Acrioceras tabarelli is restricted to the

Lower Barremian and A. (Paraspinoceras) pulcherrimum ranges through most

of the Upper Hauterivian and Lower Barremian (Thomel 1964: 73, table 5).

Acrioceras zulu, in contrast, is only known from the Upper Barremian of Zulu-

land. Thus, on stratigraphic grounds alone, separation of A. (A.) zulu from

A. tabarelli and A. pulcherrimum seems justifiable.

Acrioceras terveri (Astier, 1851) is a related Lower Barremian species that

differs mainly in the strong development of the shaft and crozier in relation to

the criocone section. Some specimens of A. terveri, figured by Thomel et al.

(1990, e.g. pl. 13 (fig. 1)), are as strongly ornamented as some of our specimens

of A. (A.) zulu (e.g. Figs 29, 33).

Acrioceras monopujaae Sarkar, 1955, appears to be restricted to the Upper

Barremian, according to Thomel et al. (1990). It is also very variable and

CRETACEOUS FAUNAS FROM SOUTH AFRICA 1A

Thomel et al. (1990) recognized four different subspecies—A. monopujaae

monopujaae Sarkar, 1955, A. monopujaae minor Thomel et al., 1990, A. m.

multicostatum Sarkar, 1955, and A. m. yvanii Thomel et al., 1990. Coiling in

A. monopujaae appears to be predominantly aspinoceratid (of the type of Prot-

acrioceras in the sense of Sarkar 1955), compared to distinctly acrioceratid in

A. (A.) zulu. This suffices to distinguish between the two species.

The criocone whorls of coarsely ornamented variants of A. (A.) zulu are

indistinguishable from those of Crioceratites (C.) yrigoyeni (Leanza) (see

p. 84). Implications of these similarities are discussed below (p. 129).

Occurrence

Upper Barremian (Barremian I-II) of Zululand.

Acrioceras sp.

Bigs 35

Material

A single specimen, SAM—PCZ8442, from the basal bed, Bed 1, at locality

170, cliff and gully sections 2 km north-west of Mlambongwenya Trading Store,

on the north side of the stream. Makatini Formation, Barremian I.

Description

The specimen is about 45 mm long, and consists of part of the recurved

crozier and an incomplete impression of the early whorls and succeeding curved

shaft, partially preserved in sparry calcite.

The earliest whorls are smooth, with a rounded whorl section. They are fol-

lowed by a stage with single ribbing and no visible lateral tuberculation. As far

as can be seen, only ventral tubercles are developed on the coiled part of the

phragmocone. At the end of the curved shaft, at or near the end of the phrag-

mocone, a single pair of tubercles are visible on the venter.

Ornament on the rest of the recurved crozier consists of rounded, prorsi-

radiate ribs only, with no sign of tuberculation. The ribs arise on the umbilical

wall, sweep forwards across the flanks, becoming progressively thicker towards

the venter, and cross the latter without interruption. Some bifurcate at the

umbilical edge.

Discussion

This species clearly differs from Acrioceras (A.) zulu in having a totally dif-

ferent mode of ornamentation, smaller size, and aspinoceratid, rather than

acrioceratid coiling. Even though the criocone part is not fully preserved, that

section is much smaller in relation to the rest of the shell than in Acrioceras (A.)

zulu. This relationship is perhaps comparable to that of Adouliceras to Ancylo-

ceras as far as general proportions are concerned.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Acrioceras sp. SAM—PCZ8442. x 2.

Acrioceras (Aspinoceras) sp. aff. brevis (d’Orbigny) in Thomel (1964: 48,

pl. 7 (fig. 11)) is very similar in overall shape, but the French specimen lacks

tubercles.

Protacrioceras hourcqi Collignon (1962: 95, pl. 214 (figs 936—7)) from the

Barremian of Antsalova, Madagascar, has coiling and relative proportions

similar to our specimen, but has a trituberculate phragmocone. Given more

material, it may be possible to connect our bituberculate specimen to the sextu-

berculate Malagasy specimen, very much in the same way as shown by the range

of variation in Acrioceras (A.) zulu. For the present, a specific identification of

the Zululand specimen is not possible.

Occurrence

Upper Barremian (Barremian I) of Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 123

INCERTAE SEDIS

Genus Hemihoplites Spath, 1924

[= Matheronites Renngarten, 1926]

Type species. Ammonites feraudinaus d’Orbigny, 1841, by original desig-

nation of Spath (1924: 84).

Discussion

Two specimens in our collections can be tentatively identified with material

from Patagonia, described recently by Riccardi & Aguirre Urreta (1989) under

the generic name of Hemihoplites Spath, 1924. Our material does not contribute

in any constructive way to the discussion of whether Hemihoplites should be

retained in a separate family Hemihoplitidae Spath, 1924, as proposed by

Wright (1957, 1981) and followed by Riccardi & Aguirre Urreta (1989), or

whether it in fact should be regarded as a recoiled crioceratitid, and included in

the family Ancyloceratidae, as suggested by Wiedmann (1962, 1966) (see also

Bogdanova 1971). We consequently refer our material to the genus Hemi-

hoplites but leave the allocation at family level open.

Occurrence

According to Riccardi & Aguirre Urreta (1989), material that could be

referred to Hemihoplites has been described from the Barremian and/or Lower

Aptian of Mallorca, France, Italy, Bulgaria, Yugoslavia, Caucasus and Crimea,

and questionably Mexico (Imlay 1938) and Canada. In southern Patagonia,

Hemihoplites occurs in the Hauterivian and Barremian. Skwarko & Thieuloy

(1989) recently described Hemihoplites from Indonesia.

Hemihoplites sp. cf. H. ploszkiewiczi Riccardi & Aguirre Urreta, 1989

Figs 36A-B, 37, 38

Compare:

Femtihoplites ploszkiewiczi Riccardi & Aguirre Urreta, 1989: 458, pl. 52

(figs 4-9), text-figs 3d—e, 5g—h, 6.

Material

BMNH C80033, from locality 170, cliff and gully sections, 2 km north-west

of Mlambongwenya Trading Store, on the north side of the stream, Makatini

Formation, Barremian I.

Description

The specimen is a wholly septate fragment with a maximum whorl height of

31 mm. Coiling appears to have been moderately evolute, with a deep, narrow,

impressed dorsum (Fig. 38). The whorl section is compressed, with an estimated

whorl breadth to height ratio of 0,5. The umbilical wall is quite low, flat and

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

sloping; the umbilical shoulder is abruptly rounded, the flanks flattened and con-

vergent, shoulders bluntly rounded, and the venter narrow and broadly

rounded. The greatest breadth is close to the umbilical shoulder.

Ornament consists of low, narrow, dense, crowded, rounded ribs. On the

preserved fragment, nine ribs extend to the umbilicus, where they are low and

wide. They sweep forwards to a point below midflank, where they split into two,

rarely three, with an occasional intercalated rib, so that there are more ribs on

the outer flank than on the inner. At the point of splitting, the ribs flex back-

wards, and are convex across the middle of the flanks, concave on the outer

flank, and strongly projected on the ventrolateral shoulders. The ribs pass

straight across the venter, weakening to the point of virtual effacement over the

siphonal line.

The sutures are deeply and intricately incised (Fig. 37). The external lobe

(E) is broad, with narrow incisions and a prominent median element; the lateral

lobe (L) is large and asymmetrically trifid; the umbilical lobe (U) is small and

imperfectly exposed; E/L is broad and asymmetrically bifid; and L/U is smaller

and more symmetrically bifid.

Discussion

Our specimen resembles Hemihoplites ploszkiewiczi Riccardi & Aguirre

Urreta, 1989, from the Hauterivian of Patagonia, as far as the flexuous ribbing is

concerned, but differs in being much more compressed (Wb/Wh = 0,5 compared

to 0,8). Given more material it may prove to be a new species.

We initially thought that this specimen might be a relatively late, closely

coiled Pseudothurmannia, perhaps allied to C. (P.) angulicostata (d’Orbigny,

1840) (see e.g. Lapeyre 1974: 82, pl. 1 (figs 1-9)), C. (P.) balearis (Nolan, 1894)

(see e.g. Wiedmann 1962: 128, pl. 8 (fig. 5), pl. 9 (fig. 1)), or C. (P.). moritilleti

(Pictet & Loriol, 1858) (see e.g. Wiedmann 1962: 132, pl. 7 (fig. 5); Sornay

1968: 4, pl. 1 (fig. 1), text-fig. 1). In all these C. (Pseudothurmannia) species,

however, some of the ribs arise in groups from elongated umbilical nodes on the

outer whorls. These are lacking in the present specimen and we provisionally

accept Riccardi & Aguirre Urreta’s (1989) allocation of the material to

Hemihoplites.

Occurrence

Barremian I of Zululand.

Hemihoplites sp. cf. H. varicostatus Riccardi & Aguirre Urreta, 1989

Fig. 36C—E

Compare:

Hemihoplites varicostatus Riccardi & Aguirre Urreta, 1989: 452, pl. 51 (figs 1-9),

pl. 52 (figs 1-3), text-figs 3a—c, 4, 5a—f.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 125

Cc D E

Fig. 36. A-B. Hemihoplites sp. cf. H. ploszkiewiczi Riccardi & Aguirre Urreta, 1989.

BMNH C80033. C-E. Hemihoplites sp. cf. H. varicostatus Riccardi & Aguirre Urreta, 1989.

BMNH C80034. Both x 2.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

10mm

Fig. 37. Hemihoplites sp. cf. H. ploszkiewiczi Riccardi & Aguirre Urreta, 1989.

Suture line of BMNH C80033.

‘2

Y UV

Oo 5

Fig. 38. Hemiholpites sp. cf. H. ploszkiewiczki

Riccardi & Aguirre Urreta, 1989. Whorl

section of BMNH C80033.

CRETACEOUS FAUNAS FROM SOUTH AFRICA IBF,

Lito.

Fig. 39. Reconstruction of Acrioceras (A.) zulu sp. nov.

Material

One specimen only, BMNH C80034 from locality 170, cliff and gully sec-

tions, 2 km north-west of Mlambongwenya Trading Store, on the north side of

the stream, Makatini Formation, Barremian I.

Dimensions

Specimen D Wb Wh Wb/Wh U

C80034 AVS) 8,4 (32) 12,5 (47) 0,67 5,2 (20)

Description

The specimen is beautifully preserved, retaining replaced shell and part of

the body chamber. Coiling is involute, with a small, shallow umbilicus (20% of

the diameter). The umbilical wall is flat, and inclined outwards; the umbilical

shoulder is abruptly rounded. The whorl section is compressed (whorl breadth

to height ratio is 0,67), with the greatest breadth below midflank. The inner

flanks are gently rounded, the outer flanks flattened and convergent, the ventro-

lateral shoulders rounded and the rather narrow venter somewhat flattened.

Ornament consists of dense, broad, relatively strong, flexuous crowded ribs.

Twenty-four ribs per whorl arise at the umbilical seam as mere striae. They are

prorsiradiate across the umbilical wall and strengthen into feeble bullae at the

ANNALS OF THE SOUTH AFRICAN MUSEUM

128

‘(OL6L ‘ezuRa) Makosi (*D) saivsad014D JO UONONI\SUODEY “Op “SIF

aig=!

CRETACEOUS FAUNAS FROM SOUTH AFRICA 129

umbilical shoulder. On the flank, the ribs are at first narrow and prorsiradiate;

they split into pairs below midflank, with occasional ribs intercalated, giving a

total of perhaps 40-50 ribs per whorl on the outer flank. The ribs are convex

across the midflank and concave on the outer flank, and cross the venter with a

distinct forward flexure and ventral convexity. Initially weak, the ribs strengthen

and broaden from inner flank to ventrolateral shoulder, although weakening and

broadening over the siphonal line.

The sutures are not exposed.

Discussion

Our specimen resembles the juvenile specimens of Hemihoplites varico-

status Riccardi & Aguirre Urreta, figured by Riccardi & Aguirre Urreta (1989,

text-fig. Sa—f) as far as ornament is concerned, but differs in being much more

involute. The umbilical diameter of the Patagonian specimens ranges from 28 to

40 per cent of the total diameter, compared to only 20 per cent in the Zululand

specimen. For the present it is best to refer to the specimen as Hemihoplites sp.

cf. H. varicostatus.

Superficially similar material has recently been recorded from Indonesia

(Skwarko & Thieuloy 1989: 31, pl. 4 (figs 2-4)) as Hemihoplites taminabuan-

enesis, where it is dated as early Barremian.

Hemihoplites sp. cf. H. varicostatus differs from H. sp. cf. H. ploszkiewiczi

(see p. 123) in its coarser ribbing, especially the blunt ventral development of

the ribs, as well as the presence of feeble bullae. Given a larger suite of speci-

mens, these two individuals might prove to be different developmental stages or

variants of the same species.

Occurrence

Barremian I of Zululand.

DISCUSSION ON SHELL SHAPE, DIMORPHISM AND SYSTEMATICS

IN CRIOCERATITINAE AND ANCYLOCERATINAE

Rawson (1975: 282) suggested that Ancyloceratidae with ancyloceratid/aspi-

noceratid coiling (herein referred to as acrioceratid), and larger, crioceratitid

forms might be dimorphic. Related to this, he observed that:

1. Spath (1924) grouped large crioceratitid and smaller aspinoceratid forms in

one genus—Hoplocrioceras; this cuts across normal taxonomic procedure of

separating forms with different coiling.

2. Thomel (1964) pointed to the similarity between the ontogeny and ornamen-

tation of some Tethyan crioceratitids and acrioceratids.

3. The type species of Paracrioceras is associated with body chambers of Acrio-

ceras in the English Snettisham Clay (of the Boreal Realm)—a situation com-

parable to that found in Tethyan faunas as described by Thomel (1964).

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Spath’s (1924) observations may be ignored, because they are subjective,

personal taxonomic views. If Spath really intended uniting forms with different

coiling, it was certainly not with dimorphism in mind.

Thomel (1981) subsequently added to his initial views, commenting on the

similarity between Acrioceras tabarelli (Astier) and Emericiceras ottohaasi

Sarkar, and Paraspinoceras dilatatum (d’Orbigny) and Crioceratites duvali

Leveillé, respectively, but he never actually referred to it as a manifestation of

dimorphism.

Immel (1978: 28), in his revision of the Hauterivian—Barremian Criocerati-

tinae, totally ignored the issue of dimorphism, but disagreed with Spath’s (1924)

views that forms with aspinoceratid and crioceratitid coiling should be referred

to the same genus (Hoplocrioceras). Immel made his views on dimorphism in

general quite clear in a recent discussion (Immel 1987); this may explain the

total omission of the subject in his (1978) work on the Crioceratitinae. Kaka-

badze (1981: 75-76) referred to Rawson’s (1975) statement, but regarded the

similarities in early whorls of acrioceratid/aspinoceratid and crioceratitid forms

as being due to ‘phylogenetic proximity’, rather than dimorphism.

Surprisingly enough, Thomel et al. (1990) scarcely touched on the topic of

dimorphism in their monographic treatment of Acrioceras from the south-east of

France. In the morphometric analysis of Acrioceras, they only referred briefly to

dimorphism (p. 49) without any in-depth discussion.

In Zululand, Crioceratites (C.) yrigoyeni (Leanza) and C. (C.) australis were

found at the same locality as Acrioceras (A.) zulu and Acrioceras sp. As we have

noted above in the systematic descriptions, the inner whorls of Crioceratites (C.)

yrigoyeni are indistinguishable from the spiral whorls of coarsely ornamented

variants of Acrioceras (A.) zulu. Some of these specimens also occur together in

the same nodules. It would be tempting to further link Crioceratites (C.) austra-

lis with the finely ribbed variants of Acrioceras (A.) zulu as a dimorphic pair. In

fact, part of a finely ribbed fragment of Acrioceras (A.) zulu is preserved in

BMNH C80046 (Fig. 20), a body chamber fragment of Crioceratites (C.) austra-

lis. But if we accept these as a dimorphic pair, it would mean formally

separating finely and coarsely ribbed Acrioceras (A.) zulu. On purely morpho-

logical grounds, separation of finely and coarsely ribbed Acrioceras (A.) zulu is

difficult, because of the presence of transitional forms, but not impossible. At

present, our knowledge of the total range of variation of the inner whorls of

C. (C.) australis is restricted to the holotype. However, the possible dimorphic

association of finely ribbed Acrioceras (A.) zulu with Crioceratites (C.) australis

may justify formal separation of these from coarsely ribbed Acrioceras (A.)

zulu. Another alternative might be to link C. (C.) australis with Acrioceras

sp.—an association substantiated by the co-occurrence of only two specimens—

the holotype of C. (C.) australis and Acrioceras sp. in the basal bed (Bed 1) at

locality 170.

The association of, and the morphological similarity between, the early

whorls of Zululand representatives of Acrioceras and Crioceratites (‘Cryptocrio-

CRETACEOUS FAUNAS FROM SOUTH AFRICA ii

ceras’) suggest that it is possible, or probable, that they constitute a dimorphic

pair. This association is analogous to that of Acrioceras—‘Emericiceras’ in the

Tethyan Realm, and Acrioceras—‘Paracrioceras’ in the Boreal Realm.

A full discussion on dimorphism in the subfamilies Crioceratitinae and

Ancyloceratinae should be based on precisely located material from the

classic European localities in the Boreal and Tethyan realms, respectively.

We here limit ourselves to a discussion of some obvious taxonomic impli-

cations of the recognition of possible dimorphism in the Crioceratitinae and

Ancyloceratinae.

Taxonomic implications

If the association of Acrioceras—‘Emericiceras’ in the Tethyan Realm,

Acrioceras—‘Paracrioceras’ in the Boreal Realm, and Acrioceras—‘Crypto-

crioceras’ in Zululand, respectively, is due to dimorphism, it would suggest a

rather complicated social life for Acrioceras with three different, geographically

separated sexual counterparts. Either Acrioceras has to be split into three differ-

ent supra-specific taxa, one for each biogeographically distinct dimorphic

partner, or ‘Emericiceras’, ‘Paracrioceras’ and ‘Cryptocrioceras’ have to be

regarded as synonyms.

There do appear to be differences between Tethyan and Boreal Acrioceras.

Subsurface material from Salzgitter, Federal German Republic, described by

Immel (1978, pl. 6 (fig. 5)) as Acrioceras gr. ex. A. tabarelli (see also Immel

1979a: 136), differs from the typical Tethyan species, but these differences are

not sufficient for generic separation. It rather seems to suggest that the differ-

ences between Boreal and Tethyan Acrioceras might be analogous, or of the

same order as those between ‘Paracrioceras’ and ‘Emericiceras’.

From a phylogenetic point of view, there are no serious objections to

uniting (Boreal and Tethyan) ‘Paracrioceras’ and ‘Emericiceras’, and regarding

them as synonyms or, at most, as a subgenus of Crioceratites. Kemper et al.

(1981), in fact, refuted Immel’s (1978) views that the north-west European and

Tethyan crioceratid faunas are specifically distinct.

The association of ‘Cryptocrioceras’ with Acrioceras in Zululand lends

support to Thomel’s (1964: 419) objection that ‘Sornayites’ (of which ‘Crypto-

crioceras’ is a possible junior objective synonym) need not be separated formally

from Crioceratites s.s. If that is the case, ‘Paracrioceras’, ‘Emericiceras’ and

‘Sornayites’ (‘Cryptocrioceras’) can all be regarded as synonyms. If separation of

these (predominantly strongly tuberculate on the inner whorls) Barremian

species from Crioceratites at subgeneric level is required, the oldest name, Para-

crioceras Spath, 1924, has priority.

If Acrioceras s.1. and the ‘Paracrioceras’—‘Emericiceras’—‘Cryptocrioceras’

part of Crioceratites are indeed dimorphs, it would be untenable to place Acrio-

ceras s.1. and Crioceratites into different subfamilies (Crioceratitinae, Ancylo-

ceratinae, Helicancylinae) as is current practice (cf. Wright 1957; Aguirre Urreta

1986) and taken to extremes by Dimitrova (1970).

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

From a taxonomic point of view, the recognition of Acrioceras s.1. and the

‘Paracrioceras—Emericiceras—Cryptocrioceras’ lineage of Crioceratites as a dimor-

phic pair would certainly make sense. To prove it is difficult on the basis of the

present material; we need more stratigraphically substantiated records on the

co-occurrence of crioceratitid and ancyloceratid forms with similar early whorls.

We doubt if the name Acrioceras will ever disappear from common use—but

the confirmation that it is the dimorphic partner of Crioceratites will confirm the

futility of referring them to two different subfamilies.

CONCLUSION

From the above discussion it appears likely, or even probable, that small

forms with acrioceratid/aspinoceratid coiling (Acrioceras) and larger forms with

crioceratitid coiling (Crioceratites) might constitute a dimorphic pair in the

family Ancyloceratidae. From a systematic point of view, this hypothesis is very

attractive, because it would unambiguously clear the way for uniting the sub-

families Crioceratitinae and Ancyloceratinae, and would further simplify the

systematics of the ‘Paracrioceras—Emericiceras—Cryptocrioceras’ plexus by

removing the traditional Boreal, Tethyan and Gondwanid geographic taxonomic

barriers.

However, because there is an element of uncertainty, we retain the names

Acrioceras and Crioceratites in the systematic description. But, being aware of

their possible dimorphic association with Acrioceras, there is no justification for

maintaining Paracrioceras, Emericiceras or Cryptocrioceras separate from each

other or from CrTioceratites s.s.; neither is separation into two subfamilies,

Ancyloceratinae and Crioceratitinae (as here reluctantly employed for taxo-

nomic ‘tidiness’) justifiable.

ACKNOWLEDGEMENTS

Financial aid from the Foundation for Research Development (South

Africa) (FRD) to Klinger, and the National Environment Research Council

(NERC) to Kennedy, is gratefully acknowledged. For discussions, advice and

literature we thank Drs M. B. Aguirre Urreta (Buenos Aires), H. Immel

(Munich), M. V. Kakabadze (Tbilisi), and G. Thomel (Saint-Martin-du-var),

but we accept responsibility for the opinions expressed here. Thanks are due to

Jacqueline Blaeske, Sally Dove and Madel Joubert (South African Museum) for

technical assistance.

REFERENCES

AGUIRRE UrreTa, M. B. 1981. In: AGUIRRE UrrETA, M. B. & Ramos, V. A. Estratigrafia y

paleontologia de la Alta Cuenca del Rio Roble Cordillera Patago6nica— Provincia de Santa

Cruz. Actas. VII Congreso Geoldgico Argentino, 20-26 septiembre 1981, San Luis 3:

101-148.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 133

AGuirrE UrretaA, M. B. 1986. Aptian ammonites from the Argentinian Austral Basin. The

subfamily Helicancylinae Hyatt, 1894. Annals of the South African Museum 96 (7):

271-314.

AGutrrE Urreta, M. B. & KLINGER, H. C. 1986. Upper Barremian Heteroceratinae (Cephalo-

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

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

ANDERSON, F. M. 1938. Lower Cretaceous deposits in California and Oregon. Special Papers

of the Geological Society of America 16: 1-339.

AstigR, J. E. 1851. Catalog descriptif des Ancyloceras appartenant 4 |’étage néocomien

d’Escragnolles et des Basses-Alpes. Annales des sciences physiques et Naturelles d’Agricul-

ture et d’Industrie 3: 435-456.

BatsEvicH, L. F. & Simonovicu, S. E. 1873. Geologicheskoe opisanie chasti kutaiiskogo

uezda, Kutaisskoi gubernii, izvestnoi pod imenem Okriba. Materialy dlya geologii

Kavkaza.

Brasco, G., NuLLo, F. E. & PLoszkiEwicz, J. 1980. El género Colchidites Djanélidzé, 1926 y

la posicion estratigrafica del género Hatchericeras Stanton, 1901 en la Estancia Tucu-Tucu,

Provincia de Santa Cruz. Revista de la Asociaci6n geologica argentina 35 (1): 41-58.

Boepanova, T. N. 1971. New Barremian ammonites from Western Turkmenia. Paleontologi-

cal Journal 3: 334-344.

BREISTROFFER, M. 1947. Notes de Nomenclature paleozoologique. Procés-verbaux de la

Société dauphinoise d’études biologiques et de protection de la nature 26: 1-S.

BREISTROFFER, M. 1952. Sur la position systématique du genre Ptychoceras D’Orb. Travaux du

Laboratoire de géologie de la Faculté des sciences de l Université de Grenoble 29: 47-54.

BRESKOVSKI, S. 1966. Biostratigrafija na barema juzhno ot selo Brestok, Varnensko. Trudove

vurkhu geologiyata na Bulgariya. Sofia (Palaeontologiya) 8: 71-121.

BusNnarbo, R. 1970. Les Pseudothurmannia (Ammonoidea) de |’Hauterivien Supérieur de la

Montagne de Lure. Documents du Laboratoire de géologique de la Faculté des sciences de

Lyon 37: 133-145.

Camacuo, H. & Riccarpi, A. C. 1978. Invertebrados. Megafauna. VII Congreso Geol6gico

Argentino. 9-15 April 1978, Neuquén: 137-144.

Casey, R. C. 1960. The Ammonoidea of the Lower Greensand. Part I. Monographs.

Palaeontographical Society: 1-44.

CasEy, R. C. 1961. The Ammonoidea of the Lower Greensand. Part 2. Monographs.

Palaeontographical Society 114: 45-118.

CoLLIGNON, M. 1949. Faune Néocomienne des couches a criocéres de Belohasifaka (Cercle de

Sitampiky) (Madagascar). Annales géologiques du Service des mines, Madagascar 15:

CoLLIGNoN, M. 1962. Atlas des fossiles caractéristiques de Madagascar (Ammonites). VIII.

(Berriasien, Valanginien, Hauterivien, Barremien). Tananarive: Service Géologique.

CoTrrEAu, J. 1937. Types du Prodrome de Paléontologie stratigraphique universelle de

d’Orbigny, B.—Néocomien supérieur ou Urgonien. Annales de paléontologie 26 (1):

17-48.

Dimitrova, N. 1967. Les fossiles de Bulgarie. IV. Crétacé inférieur. Cephalopoda (Nauti-

loidea et Ammonoidea). Izvestiya na Geologicheskiya Institut, Bulgarska Akademiya na

Naukite, Sofiya (Palaeontologiya) 4: 1-424.

Dimitrova, N. 1970. Phylogénése des ammonites hétéromorphes du Crétacé inférieur. Jzves-

tiya na _ Geologicheskiya Institut, Bulgarska Akademiya na _ Naukite, Sofiya

(Palaeontologiya) 19: 71-110.

DouviLL£, H. 1916. Les terrains secondaires dans le massif du Moghara a l’est de l’isthme de

Suez. Mémoires de l’ Académie des sciences de l'Institut de France 54: 1-184.

DrusuHcuits, V. V. & KupryavtsEv, M. Ts. 1960. Ammonitiy. Jn: Atlas niznemelovoj fauny

Severnogo Kavkazai Kryma. Trudy Vsesojuz institut gazowoj 1960: 249-355.

EGoran, V. L. 1974. A new family of loosely coiled ammonites from the Aptian deposits in the

Western Caucasus. Doklady Akademii nauk SSSR (Earth Science Sections) 217 (1-6):

223-226.

Etayo SERNA, F. 1968. Apuntaciones acerca de algunas amonitas interesantes del Hauteriviano

y del Barremiano del region de Villa de Leiva (Boyaca, Colombia, S.A.). Boletin de

geologia, Facultad de petroleas, Universidad industrial de Santander 24: 51-70.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

FOrsTER, R. 1975. Die geologische Entwicklung von Siid-Mozambique seit der Unterkreide

und die Ammoniten-Fauna von Unterkreide und Cenoman. Geologisches Jahrbuch (B)

12: 3-324.

GERHARDT, K. 1897. Beitrag zur Geologie und Palaeontologie von Sidamerika. V. Beitrag zur

Kenntnis der Kreideformation in Venezuela und Peru. Neues Jahrbuch ftir Mineralogie,

Geologie und Paldontologie (Beilageband) 11: 5—208.

GerTH, H. 1925. La fauna Neocomiana de la Cordillera Argentina en la parte meridional de la

provincia de Mendoza. Actas de la Academia nacional de ciencias en Cordoba 9 (2):

57-132.

Git, T. 1871. Arrangement of the families of Mollusks. Smithsonian Miscellaneous Collec-

tions 227: 1-49.

GroviINnE, A. T. Y. 1950. Algunos cefal6podos del Hauterivense de Neuquén. Revista de la

Asociacion geoldgica argentina 5 (2): 35-76.

GrovinE, A. T. Y. 1952. Sobre una nueva especie de Crioceras. Revista de la Asociacion geo-

logica argentina 7 (1): 71-75.

HONNORAT-BASTIDE, E. F. 1889. Formes nouvelles d’ Ammonites de Belemnites et de Crio-

ceras. Compte rendu de Il’Association francaise pour l'avancement des sciences 2: 462—466.

Hyatt, A. 1894. Phylogeny of an acquired characteristic. Proceedings of the American Philo-

sophical Society 32: 349-647.

Hyatr, A. 1900. Cephalopoda. Jn: Eastman, C. R. transl. Text-book of palaeontology, by

Karl A. von Zittel 1: 502-604. London: Macmillan & Co.

ImMtay, R. W. 1938. Ammonites of the Taraises Formation of northern Mexico. Bulletin of the

Geological Society of America 49: 539-602.

IMMEL, H. 1978. Crioceratiten (Ancyloceratina, Ammonoidea) des mediterranen und borealen

Hauterive-Barreme (Unterkreide). Palaeontographica (A) 163 (1-3): 1-85.

IMmeL, H. 1979a. Die Ammonitengliederung des mediterranen und borealen Hauterive und

Barreme unter besonderer Bericksichtigung heteromorpher Ammoniten der Gattung

Crioceratites Leveillé. Newsletters on Stratigraphy 7 (3): 121-141.

ImmeL, H. 1979b. Uber den Ursprung der borealen Crioceratiten und zur Phylogenie der Gat-

tung Crioceratites Leveillé (Ammonoidea, Kreide). Aspekte der Kreide Europas. IUGS

(Series A) 6: 129-140.

IMmMeEL, H. 1987. Die Kreideammoniten der nérdlichen Kalkalpen. Zitteliana 15: 3-163.

JELETZKY, J. A. 1964. Illustrations of Canadian fossils. Lower Cretaceous marine index fossils

of the sedimentary basins of western and arctic Canada. Geological Survey Papers. Mines

and Geology Branch, Canada 64 (11): 1-100.

KAKABADZE, M. V. 1978. On the systematic and stratigraphical significance of the genera Pseu-

docrioceras Spath, Adouliceras Thomel and Kutatissites Kakabadze. Annales du Musée

W histoire naturelle de Nice 1976 (4): XXXIV.1-—XXXIV.5.

KaKABADZE, M. V. 1981. [The ancyloceratids of the south of the USSR and their stratigraphic

significance]. Trudy Geologicheskogo instituta. Akademiya nauk Gruzinskoi SSR (n.s.)

71: 1-196. (In Russian.)

KEMPER, E., RAwson, P. F. & THrEuLoy, J. P. 1981. Ammonites of Tethyan ancestry in the

early Lower Cretaceous of north-west Europe. Palaeontology 24 (2): 251-311.

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

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

(Geology) 25 (4): 263-315.

KENNEDY, W. J. & KLINGER, H. C. 1979. Cretaceous faunas from Zululand and Natal, South

Africa. The ammonite superfamily Haplocerataceae Zittel, 1884. Annals of the South

African Museum 77 (6): 85-121.

KENNEDY, W. J. & KLINGER, H. C. 1990. Cretaceous faunas from Zululand and Natal, South

Africa. Hatchericeras Stanton, 1901 (Cephalopoda, Ammonoidea), from the Bar-

remian of Zululand. Annals of the South African Museum 99 (8): 231-243.

Kitian, W. 1895. Notice stratigraphique sur les environs de Sisteron et contribution a la con-

naissance des terrains secondaires du Sud-Est de la France. Bulletin de la Société

géologique de France (3e sér) 23: 659-803.

Kuncer, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Memoirs. Geologi-

cal Survey. Republic of South Africa 69: 1-142.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 135

KuinGer, H. C. 1990. Upper Barremian Heteroceratinae (Ammonoidea) from Caucasus,

Zululand and Patagonia: palaeobiogeographic significance. Cretaceous Research 11 (3):

321-329.

KLINGER, H. C., KAKABADZE, M. V. & KENNEDY, W. J. 1984. Upper Barremian (Cretaceous)

heteroceratid ammonites from South Africa and the Caucasus and their palaeo-

biogeographic significance. Journal of Molluscan Studies 50 (1): 43-60.

KLINGER, H. C. & KENNEDY, W. J. 1977. Cretaceous faunas from Zululand, South Africa, and

southern Mozambique. The Aptian Ancyloceratidae (Ammonoidea). Annals of the South

African Museum 73 (9): 215-359.

KoENEN, A. VON. 1902. Die Ammonitiden des Norddeutschen Neocom (Valanginien; Haute-

rivien; Barrémien und Aptien). Abhandlungen der Ko6niglich preussischen geologischen

Landesanstalt und Bergakademie (n.f.) 24: 1-451. Berlin.

KoretisHviLi, E. V. 1970. Stratigrafia i fauna kolchiditogo i smeshnix gorizontov zapadnoi

gruzi. Trudy Geologicheskogo instituta. Akademiya nauk Gruzinskoi SSR (n.s.) 25: 5-116.

KULLMANN, J. & WIEDMANN, J. 1970. Significance of sutures in phylogeny of Ammonoidea.

Paleontological Contributions. University Kansas 47: 1-32.

LapeyreE, J. F. 1974. Révision de l’ammonite-index Pseudothurmannia angulicostata d’Orbigny.

Annales du Musée dHistoire naturelle de Nice 2: 81-87.

LEANZA, A. F. 1970. Ammonites nuevos 0 poco conocidos del Aptiano, Albiano y Cenoma-

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

Asociaci6n geoldgica argentina 25 (2): 197-261.

LEVEILLE, C. 1837. Description de quelques nouvelles coquilles fossiles du département des

Basses-Alpes. Mémoires de la Société géologique de France 2: 313-315.

Manotov, J. R. 1962. New ammonites from the Barremian of North Bulgaria. Palaeontology

5i(3)2 927-939.

MEEK, F. B. 1876. A report of the invertebrate Cretaceous and Tertiary fossils of the Upper

Missouri country. Jn: HAYDEN, F. V. Report of the United States Geological Survey of the

Territories 9: |xiv, 1-629.

MULLER, G. 1892. Uber das Vorkommen von Ancyloceras gigas-Schichten bei Mellendorf

nordlich von Hannover. Jahrbuch der Koniglich preussischen geologischen Landesanstalt

13: 16-22.

Murpny, M. A. 1975. Paleontology and stratigraphy of the Lower Chickabally Mudstone (Bar-

remian—Aptian) in the Ono Quadrangle, Northern California. University of California

Publications in Geological Sciences 113: 1-52.

MyczynskIi, R. 1977. Lower Cretaceous ammonites from Sierra del Rosario (Western Cuba).

Acta palaeontologica polonica 22 (2): 139-173.

Neumayr, M. & UHLIG, V. 1881. Ueber Ammonitiden aus den Hilfsbildungen Deutschlands.

Palaeontographica 27: 129-203.

Notan, H. 1894. Sur les Crioceras du groupe du Crioceras Duvali. Bulletin de la Société géo-

logique de France (3e sér) 22: 183-196.

Opata, I., HAwicarA, S. & KAmiko, S. 1975. Geological age of the Cretaceous Chosi Group.

Bulletin of the National Science Museum (Series C, Geology and Paleontology) 1 (1):

17-36. [In Japanese. |

Opata, I. & Matsukawa, M. 1984. A Barremian occurrence of an olcostephanid, a peri-

sphinctacean ammonite, from the Chosi Group, Japan. Bulletin of the National Science

Museum (Series C, Geology & Paleontology) 10 (4): 169-180.

Opata, I., MATSUKAWA, M., TsupDaA, H., FuTAKAMI, M. & OGawa, Y. 1976. Geological age of

the Cretaceous Ishido Formation, Japan. Bulletin of the National Science Museum (Series

C, Geology & Paleontology) 2 (3): 21-138.

OpatTA, I., MayiA, S., INoUE, Y. & Matsukawa, M. 1982. Integrated mega- and micro-fossil

biostratigraphy of the Lower Cretaceous Chosi Group, Japan. Bulletin of the National

Science Museum (Series C, Geology & Paleontology) 8 (4): 145-179.

Osata, I. & Ocawa, Y. 1976. Ammonite biostratigraphy of the Cretaceous Arida Formation,

Wakayama Prefecture. Bulletin of the National Science Museum (Series C, Geology &

Paleontology) 2 (2): 93-110.

OosterR, W. A. 1860. Catalogue des Céphalopodes fossiles des Alpes suisses, 5 €me partie.

Nouveaux mémoires de la Société helvétique des sciences naturelles 18: 1-100.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

OrsBiGny, A. D’ 1840-1842. Paléontologie francaise: Terrains crétacés. 1. Céphalopodes: 1-120

(1840); 121-430 (1841); 431-662 (1842). Paris: Masson.

OrsiGny, A. D’ 1850. Prodrome de Paléontologie stratigraphique universelle des animaux mol-

lusques et rayonnés 2: 1—428. Paris: Masson.

PARONA, C. F. 1898. Descrizione di alcune Ammoniti del Neocomiano veneto. Palaeonto-

graphia italica 3: 137-144.

PuiLuirs, J. 1829. Illustrations of the geology of Yorkshire; or, a description of the strata and

organic remains of the Yorkshire Coast. York.

PicreT, J. F. & LortoL, P. DE 1858-59. Description des fossiles contenus dans le terrain néo-

comien des Voirons. Mat. Paléontologique Suisse (2): 1-64.

Rawson, P. F. 1975. The interpretation of the Lower Cretaceous heteromorph ammonite

genera Paracrioceras and Hoplocrioceras Spath, 1924. Palaeontology 18 (2): 275-283.

Rawson, P. F. 1983. The Valanginian to Aptian stages—current definitions and outstanding

problems. Zitteliana 10: 493-500.

Rawson, P. F. & Mutrer.ose, J. 1983. Stratigraphy of the Lower B and basal Cement Beds

(Barremian) of the Speeton Clay, Yorkshire, England. Proceedings of the Geologists’

Association 94 (2): 133-146.

RENNGARTEN, V. 1926. La faune des dépots crétacés de la région d’ Assakambileevka, Caucase

du Nord. Trudy Geologicheskago komiteta. St. Petersburg 147: 1-132.

Riccarpi, A. C. 1984a. Las Asociaciones de Amonitas del Jurasico y Cretacico de la Argen-

tina. Actas. Noveno Congreso Geologico Argentino 4: 559-595.

Riccarpbi, A. C. 1984b. Las Zonas de Amonitas del Cretacico de la Patagonia (Argentina y

Chile). Memoria. III Congreso Latinoamericano de Paleontologia, México, 1984:

396-405.

Riccarpi, A. C. 1988. The Cretaceous system of southern South America. Memoirs. Geologi-

cal Society of America 168: 1-161.

Riccarpi, A. C. & AGUIRRE UrRRreTA, M. B. 1989. Hemihoplitid ammonoids from the Lower

Cretaceous of Southern Patagonia. Palaeontology 32 (2): 447-462.

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

tina) from the Hauterivian—Albian of Southern Patagonia. Palaeontographica (A) 196

(4-6): 105-185.

Rocu, E. 1930. Etudes géologiques dans la région méridionale du Maroc occidental. Thése,

Paris.

Roemer, F. A. 1841. Die Versteinerungen des norddeutschen Kreidegebirges. Hannover.

Roman, F. 1938. Les ammonites Jurassiques et Crétacées. Paris: Masson.

ROUCHADZE, J. 1933. Les ammonites aptiennes de la Géorgie occidentale. Bulletin de l'Institut

géologique de Géorgie 1: 165-273.

ROUCHADZE, J. 1938. Céphalopodes nouveaux ou peu connus de |’Aptien de la Géorgie. Bul-

letin de l'Institut géologique de Géorgie 3: 129-189.

Royo y GéMEz, J. 1945. Fosiles del Barremiense Colombiano. Compilacion de los estudios

geologicos oficiales en Colombia 6: 459-494.

SARASIN, C. & SCHONDELMAYER, C. 1901-2. Etude monographique des Ammonites du Crétacé

inférieur de Chatel au Saint-Denis. Mémoire de la Société paléontologique suisse 28-29:

1-195.

SARKAR, S. S. 1954. Some new genera of uncoiled ammonites from Lower Cretaceous. Science

and Culture 19: 618-620.

SARKAR, S. S. 1955. Révision des Ammonites déroulées du Crétacé inférieur du Sud-Est de la

France. Mémoires de la Société géologique de France (n.s.) 34: 1-176.

SARKAR, S. S. 1977. Sur Binelliceras genre nouveau d’Heteromorpha et sur la validité du genre

Emericiceras Sarkar, 1954. Paldontologische Zeitschrift 51 (3-4): 258-261.

Smmionescu, I. M. 1900. Note sur quelques ammonites du néocomien frangais. Travaux du

Laboratoire géologie de la Faculté des sciences de l Université Grenoble 5: 1-17.

Skwarko, S. K. & TurEuLoy, J. P. 1989. Early Barremian (Early Cretaceous) mollusca from

western Irian Jaya, Indonesia. Publication of the Geological Research and Development

Centre, Republic of Indonesia (Palaeontology) 6: 26-42.

Sornay, J. 1968. Sur des ammonites du Barrémien et de |l’Aptien d’Ibizia (Baléares). Annales

scientifiques de l'Université de Besancon (Géologie) 5: 3-8.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 137

SpaTH, L. F. 1924. On the ammonites of the Speeton Clay and the subdivisions of the Neo-

comian. Geological Magazine 61: 73-89.

SpaTH, L. F. 1930. On the Cephalopoda of the Uitenhage Beds. Annals of the South African

Museum 28 (2): 131-157.

STANTON, T. W. 1901. The marine Cretaceous invertebrates. Report of the Princeton Univer-

sity Expeditions to Patagonia 1: 1-43.

TAVANI, G. 1942. Paleontologia della Somalia. III. Fossili del Cretaceo. Molluschi del Creta-

ceo della Somalia. Palaeontographia italica 32 (supl. 4): 7-47 (93-133).

TuiEuLoy, J. P. 1964. Un Céphalopode remarquable de |’Hauterivien basal de la Dréme:

Himantoceras nov. gen. Bulletin de la Société géologique de France (6) 7: 205-213.

THIEULOY, J. P. 1966. Leptocéres berriasiens du massif de la Grande-Chartreuse. Travaux du

Laboratoire de géologie de la Faculté des sciences de Il Université de Grenoble 42: 281-295.

THIEULOY, J. P. 1979. Matheronites limentinus n. sp. (Ammonoidea). Espéce-type d’un horizon-

repére Barrémien Supérieur du Vercors Méridional (Massif Subalpin Frangais). Géobios,

Mémoire spécial 3: 305-317.

TureEuLoy, J. P. & THOMEL, G. 1964. Sur lutilisation éventuelle des Ammonites déroulées

dans la chronologie du Crétacé inférieur. Travaux du Laboratoire de géologie de la

Faculté des sciences de l Université de Grenoble 40: 121-126.

THOMEL, G. 1964. Contribution a4 la conaissance des céphalopodes Crétacés du sud-est de la

France. Note sur les ammonites dérouleés du crétacé inférieur. Mémoires de la Société

géologique de France (n.s.) 43 (2): 1-80.

THOMEL, G. 1965. Contribution a la conaissance des céphalopodes Crétacés du sud-est de la

France. Sur deux especes du Crétacé inférieur des Basses-Alpes. Bulletin de la Société

géologique de France (7) 7: 417-420.

THOMEL, G. 1981. Ammonites. Saint-Laurent-du-Var: Editions Serre.

HHOMEL, G., DELANOY, G., CouLLET, P., Davoux, J., Fuur, F., Fuur, M., Gaze, L.,

GiIverso, J., JULLIAN, H., JULLIAN, V., LEVAN, G., MASCARELLI, E., MAUREL, L., PouPpon,

A. & Satomon, M. 1990. Le genre Acrioceras Hyatt, 1900 dans le Barrémien du sud-est

de la France. Annales du Musée dhistoire naturelle de Nice 7: 20-110.

THOMEL, G., DELANoy, G. & AUTRAN, G. 1987. Valeur taxonomique, position straigraphique

et relations phylétiques des genres d Ammonoidea: Acrioceras Hyatt, 1900 et Aspinoceras

Anderson, 1938 au cours des temps hauteriviens, ainsi que de leurs dérivés. Compte rendu

de l Académie des sciences, Paris (2) 305: 215-219.

TuHomson, M. R. A. 1971. Ammonite faunas of south-eastern Alexander Island and their strati-

graphical significance. In: ApiE, R. J. ed. Antarctic geology and geophysics: 155-160.

Oslo: Universitetsforlaget.

THomson, M. R. A. 1974. Ammonite faunas of the Lower Cretaceous of South-eastern Alex-

ander Island. Scientific Reports. British Antarctic Survey 80: 1-44.

THomson, M. R. A. 1980. Mesozoic ammonite faunas of Antarctica and the break-up of

Gondwana. Fifth International Gondwana Symposium, Wellington, New Zealand, 11-16

February 1980: 269-275.

THomson, M. R. A. 1983. ‘European’ ammonites in the Lower Cretaceous of Antarctica. Zit-

teliana 10: 407-412.

TORCAPEL, A. 1884. Quelques fossiles nouveaux de l’Urgonien du Languedoc. Bulletin de la

Société des sciences naturelles, Nimes 9-11: 1-12.

Unc, V. 1883. Die Cephalopoden der Wernsdorfer-Schichten. Denckschrift der K. Akade-

mie der Wissenschaften. Wien. (Mathematisch-Naturwissenschaften Klasse) 46: 127-290.

VaSsICEK, Z. & MICHALiK, J. 1988. Some heteromorphic ammonites from Polomec (Hauteri-

vian—Barremian, Central Western Carpathians, Czechoslovakia). Geologicky sbornik 39

(6): 655-674.

Weaver, C. 1931. Paleontology of the Jurassic and Cretaceous of West Central Argentina.

Memoirs of the University of Washington 1: 1-469.

WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentralblatt fiir Mineralogie,

Geologie und Paldontologie (B) 8: 185-195.

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

Aptychi. Abhandlungen der Mathematisch-naturwissenschaftliche Klasse. Akademie der

Wissenschaften und der Literatur 1962 (1): 1-148.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

WIEDMANN, J. 1966. Stammesgeschichte und System der Posttriadischen Ammonoideen.

Neues Jahrbuch fiir Geologie und Paldontologie. Abhandlungen 127: 13-81.

WIEDMANN, J. 1973. Ancyloceratina (Ammonoidea) at the Jurassic/Cretaceous Boundary. In:

Hatiam, A. ed. Atlas of paleobiogeography: 309-316. Amsterdam, London, New York:

Elsevier.

Wricut, C. W. 1952. A classification of the Cretaceous ammonites. Journal of Paleontology

26 (2): 213-222.

WricuT, C. W. 1957. Cephalopoda Ammonoidea. In: Moore, R. C. ed. Treatise on Inverte-

brate Paleontology. Part L, Mollusca 4: xxii, L1-L490. New York: Geological Society of

America; Lawrence, Kansas: University of Kansas Press.

Wricut, C. W. 1981. Cretaceous Ammonoidea.: Jn: House, M. R. & SENIoR, J. R. eds. The

Ammonoidea. Systematics Association. Special Volume 18: 157-174.

YENNE, K. 1949. Pedioceras, a synonym of Crioceras (Pseudocrioceras). Journal of Paleon-

tology 23: 623-624.

i ¢ i ir yee i

nen b 7 Lee

‘ee | : ei “i t

a ve s 7 ee teil

nae 0 £ <

: 7 " : i :

y ar ‘ *

ot t . : os Mek =

; ree A ¥ 3 .

i , f if

; t i

o ar We

can

1 _

. 4 7

| 7 ‘

;

fey)

: ie

fe Ch

‘ ;

= j

{

w his

‘ 4 : rm

se oie

: ea

5 Dae {

+ , 7 F)

0 eon . ee i |

I i a i

i r i

a i "i ( a, 3

oe f

ra > -

ue i : Pe over

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

Ee. 4. 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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

BARREMIAN REPRESENTATIVES OF THE

AMMONITE FAMILY ANCYLOCERATIDAE

GILL, 1871

r VOLUME 101 PART 6 MARCH 1992 ISSN 0303-2515

’

~ANNALS

"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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes... .’

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 +#£4Band

March 1992 Maart

Part 6 Deel

2 S

Sourn now XS

FOUR NEW SPECIES OF THE GENUS

PARAMELITA

(AMPHIPODA, CRANGONYCTOIDEA)

FROM SOUTH AFRICA.

BARBARA A. STEWART

CHARLES L. GRIFFITHS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

AGS, 5-2), HOD, JES, 8, tos), SUES, 5, 7-9).

Als Hos) ED), ©, OGE2, 1) MOGED), WN =y, 3,7, ii.)

WD), 1 CS), BAO. 5), 7, SGES), SAG), 2,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 129 6

Printed in South Africa by In Suid-Afrika gedruk deur

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

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D1054

FOUR NEW SPECIES OF THE GENUS PARAMELITA

(AMPHIPODA, CRANGONYCTOIDEA)

FROM SOUTH AFRICA

BARBARA A. STEWART* & CHARLES L. GRIFFITHS

Zoology Department, University of Cape Town, Rondebosch, South Africa

(With 8 figures)

[MS accepted 11 December 1990]

ABSTRACT

Four new species of the endemic South African freshwater amphipod genus Paramelita are

described from material collected in the south-western Cape Province. Males of all four species

exhibit enlargement and thickening of the second antenna and various modifications of

pereopod 3, with two of the four having this limb fully subchelate. Morphological similarities

between the four new species (P. pinnicornis, P. magnicornis, P. andronyx, and P. platypus)

and the 12 previously known species of Paramelita are discussed.

CONTENTS

PAGE

LITTASCLCTION oad cho due Nee SSN SEO BOT OC See TIE He Ta Erne ae yes Nea de a Me, xa nee ee 139

SSN SUMMA COMME pne Pert Cd facia cs ok ceca itciet actuntis us cltcinale es e cincleeieulates emleoe Selene een 140

IFBAIS ISLES SEO MIRP PPE Oe eno icles trict eke erie waitin Upc cta a eee ama A AE Sy

PRGKM ONCE PS CM CTES ere at year che alate wets cats Ae nietats bik ale Sale Pore ce (nctae Sle ule ames 157

LRXGHEIASTIGES cnn doco OSH RY OE AE oe CC REIT RS ne eI nL Oe ON Pe AR oP Do ei te 158

INTRODUCTION

The first records of the freshwater amphipod genus Paramelita were those

of Barnard (1916), who described four species collected from streams on Table

Mountain, originally placing them in the genus Gammarus. Barnard (1927)

added another six species to this list and, in 1937, Schellenberg transferred all of

these species to the genus Paramelita. Two other species have subsequently been

recognized by Thurston (1973), who described specimens collected from a cave

on the Cape Peninsula, and Griffiths (1981), who described a new species from

the Palmiet River near Grabouw.

In 1989, a research project was initiated to investigate the distribution

patterns and phylogenetic relationships of the Paramelita species. Accordingly,

an extensive sampling programme was undertaken to record distributions and to

collect samples for the purposes of constructing a phylogenetic tree based on

morphological and isozyme variation. This sampling programme has resulted in

range extensions for many of the known species, and has also revealed several

* Present address: Department of Marine Biology, South African Museum, Cape Town.

139

Ann. S. Afr. Mus. 101 (6), 1992: 139-158, 8 figs.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

new undescribed forms. Some of these could be linked to existing species

complexes, and will be discussed elsewhere. However, the relationships of four

of the newly discovered taxa were not immediately evident, and are described

below.

SYSTEMATICS

Superfamily CRANGONYCTOIDEA Bousfield, 1973

Family Paramelitidae Bousfield, 1977

Paramelita Schellenberg, 1926

Paramelita pinnicornis sp. nov.

Figs 1-2

Material examined

Holotype. SAM-—A40004, male, 13,5 mm, from a tributary of the

Burgersbos River (34°01'S 18°25'E) crossing Rhodes Drive, Constantia, on the

Cape Peninsula. Collected by B. A. Stewart and Y. Dempster on 9 August

1989.

Paratypes. SAM-—A40005, 14 males, 20 ovigerous females, from the same

sample as the type specimen.

Other material. SAM—A40006, 52 specimens, collected 12 August 1990, and

SAM-A40007, 5 specimens, collected 12 August 1990, from two adjacent

streams flowing into Koeélbaai in the Cape Hangklip area on the east coast of

False Bay. SAM—A40008, 83 specimens (13 July 1988), from Kenilworth Race

Course on the Cape Peninsula.

Etymology

From the Latin pinna (feather or plume) and cornis (horn), an allusion to

the fin, or wing-like projections that are present on article 5 of the second

antenna.

Description (of holotype, male, 13,5 mm)

Body colour when alive grey tinged with pink. Head slightly shorter than

pereon segments 1 and 2 combined, anteroventral margin excavate to

accommodate inflated article 1 of antenna 2, eyes glistening white when alive,

difficult to discern in preserved material. Antenna 1 0,6 times length of body,

setation sparse, articles 1 and 2 of peduncle subequal, each twice length of

article 3, flagellum twice length of peduncle, 31-articulate, accessory flagellum

6-articulate, reaching past article 4 of primary flagellum. Antenna 2 approxi-

mately the same length as antenna 1, but considerably stouter, peduncle sparsely

to moderately setose, article 4 and 5 each about 2,7 times length of article 3,

outer margin and tip of article 5 extended into an elongate triangular flange,

flagellum 0,6 times length of enlarged peduncle, 16-articulate, sparsely setose.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 141

S—_—_—_..-_——_

Jas on oe — —

Fig. 1. Paramelita pinnicornis sp. nov., holotype, male, 13,5 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Pereopod 3. E. Coxa 4 and pereopod 4. F. Pereopod 7.

Scales = 1 mm.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Paramelita pinnicornis sp. nov., holotype, male, 13,5 mm. A. Upper lip. B. Left

mandible. C. Lower lip. D. Maxilla 1. E. Maxilla 2. F. Maxilliped. G. Gnathopod 1.

H. Gnathopod 2. I. Pleopod 3. J. Uropod 1. K. Uropod 2. L. Uropod 3. M. Telson.

Scale = 1 mm.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 143

Left mandible with incisor bluntly 4-toothed, lacinia mobilis with four blunt

teeth, six spinose accessory blades, molar strongly triturative, 3-articulate palp

longer than body of mandible, article 1 as long as wide, article 2 3,5 times length

of 1, with approximately 16 setae anteriorly, article 3 1,2 times length of 2, distal

half lined with many short setae, six long apical setae present, tuft of four setae

approximately 0,7 along length. Right mandible, incisor 3-toothed, lacinia

mobilis bifurcate, three accessory blades. Maxilla 1, inner plate with five

pectinate setae, inner margin pubescent, outer plate bearing two terminal rows

each of about five stout serrated spines, palp exceeding outer plate, with eight

apical spines. Maxilla 2, inner plate a little shorter and narrower than outer

plate, proximally sparsely pubescent, both plates strongly setose terminally.

Maxilliped, inner plate with many curved spinose setae, outer plate with

approximately 10 stout, blunt spine-teeth on inner margin and 10 terminal

curved spinose setae, palp article 2 the longest, 2 and 3 densely setose medially,

4 with four short setae on margin.

Pereon segments with very few dorsal setae, coxae 1-3 slightly deeper than

corresponding segments, quadrate, sparsely setose ventrally, coxa 4 posteriorly

excavate, slightly deeper than long, sparsely setose on ventral margin, coxae 5

and 6 longer than deep, bilobed, few setae ventrally, coxa 7 semicircular,

smooth, segments 2—7 bearing one pair of coxal gills each, segment 2 with

one, segments 3, 4, 5 and 7 with two, and segment 6 with four sausage-shaped

sternal gills. Gnathopod 1 subchelate, articles 5 and 6 together slightly longer

than 2, article 6 approximately twice as long as 5, longer than wide, palm

relatively straight, oblique, palmar angle with two long and three short spines,

dactyl as long as palm. Gnathopod 2 similar in structure to, but 1,4 times length

and sturdier than 1, inner margin of article 2 bearing seven groups of strong

spines, articles 5 and 6 combined longer than article 2, article 6 approximately

twice as long as 5, slightly longer than wide, palm slightly convex, oblique,

defined by four stout spines, dactyl as long as palm. Pereopod 3 1,4 times length

of 4, articles 5 and 6 highly modified, 5 being posteriorly lobed, the lobe armed

with five long and four shorter spines, article 6 folded back against lobed

posterior margin of 5, bearing five short stout spines, dactyl stout, with eight

short spinules. Pereopod 4 unmodified, article 5 with three posterior spines,

article 6 with five pairs of posterior spines, dactyl with seven spinules. Pereopod

5 basis posteriorly expanded, article 4 0,8 length of 5, 5 and 6 subequal in

length, article 5 with three pairs of spines, article 6 with five pairs of spines,

dactyl with 10 spinules. Pereopods 6 and 7 similar in structure, bases expanded

posteriorly, article 6 with six pairs of spines, dactyls each with 14 spinules

anteriorly.

Pleon segments 1-3 sparsely setose dorsally, first pleonal epimeron

rounded-quadrate, 2 and 3 quadrate, setose on posterior margin. Pleon

segments 4—6 sparsely setose dorsally. Uropod 1 extending slightly beyond 2, 1,1

length of uropod 3, rami equal, 0,7 length of peduncle, each ending in five

spines. Uropod 2 shorter, stouter than 1, rami subequal, each with five apical

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

spines. Uropod 3 exceeding 2 by 0,9 length of outer ramus, peduncle longer

than broad, inner ramus reduced, 0,4 times length of peduncle, terminating in

two spines and a few long setae, outer ramus three times length of peduncle, six

groups of setae on inner and eight on outer margin, small second article ending

in two spines. Telson as broad as long, deeply cleft, each lobe with one large

subapical spine and several apical setae.

Remarks

Paramelita pinnicornis sp. nov. adult males are clearly distinguished from

other Paramelita species by the fin-like projections on the peduncle of antenna 2

and the claw-like structure of the distal end of pereopod 3. Antenna 2 in females

is slender and shorter than 1, and pereopod 3, like 4, is not modified. In most

other respects, the females resemble the males. Although the fin-like projections

of the second antenna in males are unique, a ‘claw-like’ pereopod 3 is also found

in P. auricularius (Barnard, 1916), from Table Mountain, and in P. andronyx

sp. nov. from Kasteelsberg. Despite their superficial similarity, however, these

structures are not homologous, and therefore not evidence of close affinities

between these three species. In P. pinnicornis sp. nov. the ‘claw’ is achieved by

the folding back of article 6 against the lobed spiny posterior margin of article 5.

In P. andronyx sp. nov., however, it is article 4 that is strongly protruded,

with the right angle joint between articles 5 and 6 completing the claw. In

P. auricularius, an elongated article 6 folds back against the lobed, swollen

posterior margin of article 5, but this is of a quite different shape to the structure

in P. pinnicornis sp. nov. Coxa 4 in the latter two species is either quadrangular,

or gently concave posteriorly, whereas in P. pinnicornis sp. nov. it is distinctly

excavate posteriorly.

Paramelita magnicornis sp. nov.

Figs 3-4

Material examined

Holotype. SAM-A40009, male, 15,0 mm, from a stream draining the

Swartkop Mountains (34°14’S 18°29’E) near Millers Point on the southern Cape

Peninsula. Collected by B. A. Stewart and C. L. Griffiths on 30 November 1989.

Paratypes. SAM-—A40010, 13 males and eight females, from the same

sample as the holotype.

Other material. SAM—A40013, numerous specimens, from the same stream

as the holotype (date unrecorded). SAM-—A40011, numerous specimens,

collected 9 August 1989, and A40015, 3 specimens, collected 16 August 1990,

a stream draining Chapman’s Peak, Cape Peninsula. SAM-—A40012, 10 speci-

mens (date unrecorded), a stream draining the Kalk Bay Mountains near

Clovelly, Cape Peninsula. SAM—A40014, 39 specimens, 30 November 1989, and

A40016, 10 specimens (date unknown), Peck’s Valley stream on Boyes Drive,

Cape Peninsula.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 145

Etymology

From the Latin magnus (large), alluding to the swollen and elongated

second antenna.

Description (of holotype, male, 15,0 mm)

Body colour when alive off white. Head shorter than pereon segments 1

and 2 together, anteroventral margin excavate to accommodate inflated article 1

of antenna 2, eyes glistening white when alive, difficult to discern when

preserved. Antenna 1 relatively short, 0,4 length of body, setation sparse,

articles 1 and 2 subequal, each twice length of 3, flagellum 1,7 times length of

peduncle, 30-articulate, accessory flagellum 6-articulate, reaching to end of

article 4 of flagellum. Antenna 2 1,2 times length of 1 and considerably stouter,

peduncle moderately setose posteriorly, article 4 1,9 length of article 3, distally

inflated, article 5 slightly shorter than article 4, flagellum 0,8 times length of

peduncle, 16-articulate, moderately setose posteriorly. Left mandible, incisor,

bluntly 5-toothed, lacinia mobilis with 4 blunt teeth, four accessory blades,

molar strongly triturative, palp longer than body of mandible, article 1 as long as

wide, article 2 five times length of article 1, with 10 strong setae anteriorly,

article 3 slightly shorter than 2, distal half with comb of short setae, six long

apical setae, tuft of setae half-way along length. Right mandible, incisor

4-toothed, lacinia mobilis bifurcate, two accessory blades. Maxilla 1, inner

plate with 7 setae, inner margin pubescent, outer plate terminating in nine

stout serrated spines, palp exceeding outer plate, with eight stout apical setae.

Maxilla 2, inner plate a little shorter and narrower than outer plate, proximally

pubescent, both plates strongly setose terminally. Maxilliped, inner plate with

many curved spinose setae, outer plate with approximately seven stout blunt

Spine-teeth on inner margin and eight terminal curved setae, palp article 2 the

longest, inner margin with row of strong curved setae, article 3 densely setose.

Pereon segments dorsally smooth, coxae 1-3 deeper than corresponding

segments, quadrate, moderately setose ventrally, coxa 4 excavate posteriorly,

approximately as deep as long, moderately setose ventrally, coxa 5 and 6 longer

than deep, bilobed, coxa 5 moderately setose ventrally, coxa 6 with a few short

setae and spinules, coxa 7 semicircular, setose ventrally, segments 2—7 bearing

one pair of coxal gills each, segments 2 with one, segments 3, 4, 5 and 7 with two,

and segment 6 with four sausage-shaped sternal gills. Gnathopod 1 subchelate,

article 2 bearing plumose setae on both anterior and posterior margins and two

groups of spines on inner surface, articles 5 and 6 together longer than 2, article 6

1,7 times length of 5, longer than wide, palm gently convex, oblique, with five

palmar spines, dactyl as long as palm. Gnathopod 2 similar to, but 1,2 length and

sturdier than 1, article 2 with two groups of spines on inner margin and a few

plumose setae on anterior margin, articles 5 and 6 together longer than article jhe

article 6 1,7 times the length of 5, longer than wide, palm convex, distinctly

oblique, defined by four stout spines, dactyl as long as palm. Pereopod 3

approximately the same length as 4, article 4 anteriorly lobed over 5 and

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Paramelita magnicornis sp. nov., holotype, male, 15,0 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Pereopod 3. E. Pereopod 4. F. Pereopod 5. G. Pereopod 7.

Scales = 1 mm.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 147

Fig. 4. Paramelita magnicornis sp. nov., holotype, male, 15,0 mm. A. Upper lip. B. Left

mandible. C. Lower lip. D. Maxilla 1. E. Maxilla 2. F. Maxilliped. G. Gnathopod 1.

H. Gnathopod 2. I. Pleopod 1. J. Uropod 1. K. Uropod 2. L. Uropod 3. M. Telson.

Scale = 1 mm.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

posterodistally protruded into a triangular tooth, articles 4, 5 and 6 densely setose

posteriorly, dactyl with six spinules. Pereopod 4 similar in structure to 3, article 4

posterodistally protruded into a distinct triangular tooth, dactyl with four

spinules. Pereopods 5, 6 and 7, bases moderately expanded posteriorly, with

some simple and plumose setae anteriorly and posteriorly, article 4 shorter than 5

and 6, articles 5 and 6 approximately equal in length, article 5 with two groups,

and 6 with four to five groups of spines posteriorly, articles 4, 5 and 6 moderately

to densely setose anteriorly, dactyls of pereopods 5 and 7 with six spinules, and of

pereopod 6 with seven spinules.

Pleon segments 1-3 with some dorsal setae, epimeral plates rounded to

quadrate, ventrally setose. Pleon segments 4-6 moderately setose dorsally.

Uropod 1 extending a little beyond uropod 2, 1,5 length of uropod 3, rami

subequal, 0,6 times length of peduncle, each ending in four spines. Uropod 2

shorter than 1, inner ramus marginally longer than outer, each with four apical

spines. Uropod 3 relatively short, exceeding 2 by 0,7 length of outer ramus,

peduncle longer than broad, inner ramus short, 0,6 length of peduncle and 0,3

times length of outer ramus, with four apical spines and one seta, outer ramus

2,4 times length of peduncle, three groups of spines and setae on inner and two

on outer margin, second segment reduced, only 5 per cent of length of first

segment. Telson broader than long, deeply cleft, each lobe bearing two stout

subapical spines, seven apical setae and two setae arising from the dorsal surface

about half way along the length.

Remarks

Paramelita magnicornis sp. nov. is most similar to the common and widely

distributed P. capensis (Barnard, 1916), with which it lives sympatrically in at

least two known localities on the Cape Peninsula. Adult males of this newly

described species are distinguished from P. capensis primarily by the swollen and

elongate peduncle of antenna 2 and the ‘spur-like’ projections of the posterodistal

apices of article 4 of the first and second pereopods. The thickening and

elongation of articles 4 and 5 of antenna 2 are also characteristic, these articles

being noticeably more swollen distally than proximally. In females, antenna 2 is

relatively slender and shorter than 1; similarly, pereopods 3 and 4 are unmodified.

Coxa 4 in P. magnicornis sp. nov., as in P. capensis, is distinctly excavate.

Paramelita andronyx sp. nov.

Figs 5-6

Material examined

Holotype. SAM—A40017, male, 16,1 mm, from a tributary of the Riebeek’s

River (33°22’S 18°50’E), above the farm Waterval, on the slopes of

Kasteelsberg, in the Malmesbury district. Collected by B. A. Stewart and

P. A. Cook in September 1989.

—————

FOUR NEW SPECIES OF THE GENUS PARAMELITA 149

Paratypes. SAM—A40018, 10 males, three females, from the same locality

as the type specimen.

Other material. SAM—A40019, 12 specimens, collected 24 September 1989,

from a nearby farm, Wynkeldersberg. This is the only known other record of

this species to date.

Etymology

From the Greek aner (man) and onux (claw), alluding to the claw-like

structure of pereopod 3 in adult males.

Description (of holotype, male, 16,1 mm)

Body colour when alive whitish, tinged with pink. Head shorter than

pereon segments 1 and 2 together, margin between eye lobe and post-antennal

angle gently excavate to accommodate inflated article 1 of antenna 2, eyes

glistening white when alive, invisible when preserved. Antenna | relatively long,

0,6 length of body, sparsely setose, flagellum 1,5 times length of peduncle,

29-articulate, accessory flagellum 3-articulate, reaching to article 3 of flagellum.

Antenna 2 a little stouter and 0,8 times length of antenna 1, peduncle

moderately setose, article 3 bearing a semicircular lobe posteriorly, article 4

three times length of 3, laterally swollen, article 5 0,8 times length of 4,

flagellum 1,2 times length of peduncle, 17-articulate, moderately setose. Left

mandible, incisor bluntly 5-toothed, lacinia mobilis with four blunt teeth, two

bifurcate, one simple and one pectinate accessory blade, molar strongly tritu-

rative, palp longer than body of mandible, article 1 longer than wide, article 2

2,6 times length of 1, with nine setae anteriorly, article 3 approximately the

same length as 2, distal half lined with short setae, nine long apical setae

present, two tufts of setae about half-way along length. Right mandible, incisor

4-toothed, lacinia mobilis bifurcate, four accessory blades. Maxilla 1, inner plate

setose terminally, outer plate bearing eight serrate spines, palp exceeding outer

plate, with six apical spines and three apical setae. Maxilla 2, inner plate shorter

than outer, proximally pubescent, both plates strongly setose terminally.

Maxilliped, inner plate with three spines and five curved setae, outer plate

with eight stout spines on inner margin and seven terminal curved setae, palp

article 3 as long as article 2, both articles strongly setose medially.

Pereon segments dorsally smooth, coxae 1-3 deeper than corresponding

segments, quadrate, setose ventrally, coxa 4 only very slightly concave, deeper

than long, setose on ventral margin, coxae 5 and 6 longer than deep, bilobed,

bearing setae and a few spinules ventrally, coxa 7 semicircular, bearing short

stout setae ventrally, segments 2—7 bearing one pair of coxal gills each, segment 2

with one, segments 3, 4, 5 and 7 with two, and segment 6 with four sausage-

shaped sternal gills. Gnathopod 1 subchelate, articles 5 and 6 together longer

than article 2, inner posterior margin of article 2 with five stout spines, article 6

1,4 times length of article 5, longer than wide, palm slightly convex, gently

oblique, with four palmar spines, dactyl as long as palm. Gnathopod 2 similar

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Paramelita andronyx sp. nov., holotype, male, 16,1 mm. A. Lateral aspect. B. Antenna 1.

C. Antenna 2. D. Pereopod 3. E. Pereopod 4. F. Pereopod 5. G. Pereopod 6. H. Pereopod 7.

Scales = 1 mm.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 151

Fig. 6. Paramelita andronyx sp. nov., holotype, male, 16,1 mm. A. Upper lip. B. Left

mandible. C. Lower lip. D. Maxilla 1. E. Maxilla 2. F. Maxilliped. G. Gnathopod 1.

H. Gnathopod 2. I. Pleopod 3. J. Uropod 1. K. Uropod 2. L. Uropod 3. M. Telson.

Scale = 1 mm.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

to, but larger than, 1, articles 5 and 6 together longer than article 2, inner

posterior margin of article 2 bearing five pairs of stout spines, article 6 1,6 times

the length of 5, longer than wide, palm convex, slightly oblique, defined by four

stout spines, dactyl as long as palm. Pereopod 3 highly modified and 1,3 times

length of 4, inner posterior margin of article 2 bearing five pairs of stout spines,

articles 4, 5 and 6 modified to form a claw-like structure, article 4 posterodistally

strongly projected, moderately setose, article 5 short and stout, bearing short

spine-like setae posteriorly, article 6 bent at right angles to article 5, bearing a

few short stout setae posteriorly, forming a claw with projection of article 4,

dactyl with a single spinule. Pereopod 4 unmodified, articles 4, 5 and 6

moderately setose and bearing some spines, dactyl with a single spinule.

Pereopods 5, 6 and 7, bases slightly expanded posteriorly, bearing spinules and

setae anteriorly and setae posteriorly, articles 4 and 5 moderately setose and

bearing some groups of spines, article 6 with five or six clusters of spines

anteriorly, dactyls always with only a single spinule.

Pleon segments with a few setae along posterodorsal margins, first epimeral

plate rounded-quadrate, 2 and 3 quadrate, setose ventrally. Pleon segments 4—6

more heavily setose dorsally. Uropod 1 extending to tip of uropod 2, 1,5 length

of uropod 3, rami subequal, 0,8 times length of peduncle, both rami with some

setae and spines along lateral margins, each ramus terminating in four spines.

Uropod 2 shorter than 1, inner ramus slightly longer than outer, 1,2 times length

of peduncle, both rami with setae and spines laterally, each ending in five

terminal spines. Uropod 3 relatively short, exceeding uropod 2 by 0,6 length of

outer ramus, peduncle longer than broad, inner ramus short, 0,6 length of

peduncle and 0,3 length of outer ramus, terminating in two spines and a single

seta, outer ramus approximately twice the length of peduncle, two groups of

spines and setae on inner and three on outer margin, second article absent.

Telson broader than long, deeply cleft, each lobe with one large subapical spine

and four to five apical setae.

Remarks

In addition to their uniquely subchelate first pereopods, P. andronyx sp.

nov. males from Kasteelsberg are easily identified by the large semicircular lobe

on the posterior margin of article 3 of antenna 2. In adult females, antenna 2 is

more slender and shorter than 1, article 3 is not lobed, and an unmodified

pereopod 3 resembles pereopod 4 in structure. In other respects, females are

similar to males. Paramelita andronyx sp. nov. males share a lobed article 3

(along with a distinctly swollen article 4) of antenna 2, with both P. flexa

Griffiths, 1981, and P. auricularius, which also possesses a modified pereopod 3.

The claw on pereopod 3 in P. auricularius is, however, formed from articles 5

and 6 only, not article 4. Paramelita flexa is clearly distinguished from

P. andronyx sp. nov. both by the shape of antenna 2 and by its unmodified

pereopod 3. In addition to the swelling of article 4 of the second antenna, other

FOUR NEW SPECIES OF THE GENUS PARAMELITA 153

features, such as the possession of only a single spinule on each dactyl, the

poorly excavate coxa 4, and the loss of a second segment on the outer ramus of

the third uropod, suggest that P. andronyx sp. nov. might have affinities with

P. crassicornis (Barnard, 1916), and P. tulbaghensis (Barnard, 1927).

Paramelita platypus sp. nov.

Figs 7-8

Material examined

Holotype. SAM-—A40020, male, 12,8 mm, from Fisherman’s Kloof, a

tributary of the Fernkloof River flowing through the Fernkloof Nature Reserve

(34°24’S 19°14’E) near Hermanus, Cape Province. Collected in September 1989

by B. A. Stewart and P. A. Cook.

Paratypes. SAM-—A40021, 8 males, 12 females, also from Fisherman’s

Kloof.

Other material. SAM-—A40022, numerous specimens, collected 22 July 1990,

from a stream near Stanford, Cape Province.

Etymology

From the Greek platus (broad) and pous (foot), alluding to the widened

article 4 of pereopods 3 and 4.

Description (of holotype, male, 12,8 mm)

Body colour when alive orange to pink, eyes white when alive, invisible

when preserved. Head considerably shorter than pereon segments 1 and 2

together, ventral margin excavate to accommodate inflated article 1 of antenna 2.

Antenna 1 relatively long, 0,7 times length of body, setation sparse, flagellum

2,2 length of peduncle, 41-articulate, accessory flagellum S-articulate, reaching

to article 5 of primary flagellum. Antenna 2 approximately the same length as,

but considerably stouter than, antenna 1, peduncle elongate, moderately setose,

article 4 three times length of unmodified article 3, articles 4 and 5 equally long

and relatively slender, lacking projections, flagellum 1,1 times length of enlarged

peduncle, 22-articulate, sparsely setose. Left mandible, incisor with two blunt

teeth, lacinia mobilis with four blunt teeth, three simple, and one bifurcate

accessory blade, molar strongly triturative, palp longer than body of mandible,

article 1 as long as wide, article 2 six times length of article 1, with

approximately four groups of setae and one spine on anterior margin, article 3

1,3 times length of 2, distally lined with short setae and bearing six long apical

setae, tuft of about four setae half way along length. Right mandible, incisor

3-toothed, lacinia mobilis bifurcate, three flattened spinose accessory blades.

Maxilla 1, inner plate terminally setose, inner margin pubescent, outer plate

terminating in about nine stout serrated spines, palp exceeding outer plate, with

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

eight stout apical setae. Maxilla 2, inner plate shorter and narrower than outer

plate, proximally pubescent, both plates strongly setose terminally. Maxilliped,

inner plate with many curved spinose setae, outer plate with about nine stout

spine-teeth on inner margin and six terminal spinose setae, palp article 2 the

longest, articles 2 and 3 densely setose medially.

Pereon segments with a few setae dorsally, coxae 1-3 slightly deeper than

corresponding segments, quadrate, setose ventrally, coxa 4 virtually quadrate,

only very slightly concave posteriorly, height and length subequal, setose

ventrally, coxa 5 and 6 longer than deep, bilobed, setose ventrally, coxa 7

semicircular, setose ventrally, segments 2—7 bearing one pair of coxal gills each,

segments 4, 5 and 7 with two, and segment 6 with four sternal gills. Gnathopod

1 subchelate, articles 5 and 6 together longer than 2, article 6 1,6 times length of

5, longer than wide, palm slightly convex, palmar angle with two long and three

short spines, dactyl as long as palm. Gnathopod 2 similar in structure but larger

than 1, articles 5 and 6 combined a little longer than 2, two pairs of short spines

on inside of article 2, article 6 1,7 times length of 5, longer than wide, palm

strongly convex, transverse, defining angle rectangular, bearing four strong

spines, dactyl as long as palm. Pereopod 3 enlarged, 1,2 times length of 4, article 2

with seven spinules on anterior, and eight spinules on posterior margin, article 4

greatly expanded laterally and lobed posteriorly, three spinules on anterior

margin, articles 4, 5 and 6 densely setose posteriorly, dactyl with five spinules.

Pereopod 4 article 2 with nine anterior and five posterior marginal spinules,

article 4 laterally expanded, although not quite as pronounced as in pereopod 3,

with two small spinules on anterior margin, articles 4, 5 and 6 again densely setose

posteriorly, dactyl bearing five spinules. Pereopods 5, 6 and 7, article 2

moderately expanded posteriorly, with spinules and some setae anteriorly, setose

posteriorly, article 4 shorter than 5 and 6, bearing three groups of spines

posteriorly, articles 5 and 6 subequal in length, article 5 with three groups of

spines and article 6 with five groups of spines posteriorly, both 4 and 5 densely

setose anteriorly, 6 moderately setose, dactyl of pereopod 5 with seven spinules,

those of pereopods 6 and 7 with 10 spinules each.

Pleon segments 1-3 sparsely setose dorsally, epimeral plates rounded-

quadrate, ventrally setose. Pleon segments 4—6 moderately setose dorsally.

Uropod 1 extending slightly beyond 2, 0,9 length of uropod 3, rami subequal, 0,8

times length of peduncle, each ending in four spines. Uropod 2 shorter than 1,

inner ramus slightly longer than outer, each with one dorsal and four apical

spines. Uropod 3 elongate, exceeding uropod 2 by 0,9 length of outer ramus,

peduncle longer than broad, inner ramus short, 0,6 length of peduncle and only

0,2 length of outer ramus, with 3 apical spines, outer ramus about four times

length of peduncle, six groups of spines and setae on each margin, second

segment very reduced and only 4 per cent of length of first segment. Telson

broader than long, deeply cleft, each lobe bearing one stout subapical spine, four

apical setae, two subapical setae, and two small plumose setae about one third the

way along the outer margin.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 155

Fig. 7. Paramelita platypus sp. nov., holotype, male, 12,8 mm. A. Lateral aspect. B. Antenna 1.

C. Antenna 2. D. Coxa 3 and pereopod 3. E. Coxa 4 and pereopod 4. F. Pereopod 6.

G. Pereopod 7. Scales = 1 mm.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Paramelita platypus sp. nov., holotype, male, 12,8 mm. A. Upper lip. B. Left mandible.

C. Lower lip. D. Maxilla 1. E. Maxilla 2. F. Maxilliped. G. Gnathopod 1. H. Gnathopod 2.

I. Pleopod 2. J. Uropod 1. K. Uropod 2. L. Uropod 3. M. Telson. Scale = 1 mm.

FOUR NEW SPECIES OF THE GENUS PARAMELITA 157

Remarks

Paramelita platypus sp. nov. males are unusual in two respects—the

possession of extremely elongate and sturdy second antennae, and the wide,

flattened articles 4 of pereopod 3, and to a lesser extent, pereopod 4. In

females, antenna 2 is slender and shorter than antenna 1, articles 4 of

pereopods 3 and 4 are not flattened, and the antennae and pereopods are only

moderately setose. In other respects, the females resemble the males. In

addition, all of the pereopods are markedly setose. Although several Paramelita

species have elongated second antennae, none of the known species have males

with the first two pereopods modified as in P. platypus sp. nov. Coxa 4 in this

species is only very slightly concave posteriorly, a condition found in several of

the other Paramelita species, such as P. aurantius (Barnard, 1927), P. granuli-

cornis (Barnard, 1927), P. crassicornis, P. auricularius and P. andronyx sp. nov.

DISCUSSION

In his account of the ten Paramelita species known at that time, Barnard

(1927) commented on three ‘evolutionary tendencies’ in the genus: the

thickening of the second antennae, modifications of pereopod 3, and variations

in the shape of coxa 4. All four species described here show unusual

modifications of these features. Of the 12 previously known and the additional

four species described here, at least 11 show some degree of enlargement, or

‘pediformity’, of the second antennae. This development is most marked in large

adult males. The Paramelita species share this phenomenon with the Australian

paramelitid genus Uroctena (Williams & Barnard 1988). A_ ‘claw-like’

pereopod 3, found in three of the Paramelita species, has not been recorded in

other paramelitids. This modification appears to have evolved more than once,

and is probably a clasping organ used in reproduction. The shape of coxa 4 in

Paramelita species varies from being strongly excavate posteriorly, such as in

P. capensis, to being quadrate, as in the case in P. granulicornis. Although this

coxal plate is deeply excavate in the most primitive Australian genus

Austrogammarus, it is only ‘weakly’ emarginate in Uroctena (Williams &

Barnard 1988). The evolutionary trends within the Paramelita species, as well as

the relationship of this genus with the Australian paramelitid genera will be the

subject of a later study.

ACKNOWLEDGEMENTS

Financial support for this project was provided by the Foundation for

Research Development. We are most grateful to our willing and tireless families

for accompanying us on our numerous collecting trips and tolerating frequent

roadside amphipod stops. Richard Brooke kindly helped with the naming of the

species.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

BARNARD, K. H. 1916. Contributions to the crustacean fauna of South Africa. 5. The

Amphipoda. Annals of the South African Museum 15 (3): 105-302.

BARNARD, K. H. 1927. A study of the freshwater isopodan and amphipodan Crustacea of South

Africa. Transactions of the Royal Society of South Africa 14 (2): 139-215.

GriFFiTHS, C. L. 1981. The freshwater Amphipoda (Crustacea) of South and South West

Africa. Annals of the South African Museum 83 (5): 79-97.

SCHELLENBERG, A. 1937. Kritische Bemerkungen zur Systematik der Stsswassergammariden.

Zoologische Jahrbucher. Abteilung fiir Systematik, Geographie und Biologie der Thiere 69:

469-516.

Tuurston, M. H. 1973. A new species of Paramelita (Crustacea: Amphipoda) from South

Africa. Annals of the South African Museum 62 (5): 159-168.

WiLuiAMs, W. D. & Barnarp, J. L. 1988. The taxonomy of crangonyctoid Amphipoda

(Crustacea) from Australian fresh waters: foundation studies. Records of the Australian

Museum. Supplement 10: 1-180.

at T i "

i ay Oi

, i fi

1 J i

“all

{sine :

{ i ; i :

i in

{ Site

= ‘

d ) /

; 3 ;

een

/ he

i .

nas

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7) SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

Cee tue Figure depicting C: namacolus.. ..’: “. . . mC. 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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

BARBARA A. STEWART

CHARLES L. GRIFFITHS

FOUR NEW SPECIES OF THE

GENUS PARAMELITA

(AMPHIPODA, CRANGONYCTOIDEA)

FROM SOUTH AFRICA

Y VOLUME 101 PART 7 —- JUNE 1992 ISSN 0303-2515

\ ‘ :

‘ ANNALS

—— aD

‘OF THE SOUTH AFRICAN

CAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.

2. LAYOUT should be as follows:

(a) Centred masthead to consist of

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes...’

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 + °+#£4Band

June 1992 Junie

Part 7 Deel

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES.

PART 18. HOLOTHUROIDEA

AHMED S. THANDAR

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

i, DUS, GO, ED, 4-5, Cotto, %,), SUES, 5, JO),

Aly Mens), TCD). @, O22, Dy MOGES), TNT, 5, 7, tsjo.ti),

14(1-2), 15(4-5), 24(2, 5), 27, 31(1-3), 32(5), 33,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 130 X

Printed in South Africa by In Suid-Afrika gedruk deur

ihe Rustica (Press; Pty... Ltde; Die Rustica-pers, Edms., Bpk.,

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D1318

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES.

PART 18. HOLOTHUROIDEA

AHMED S. THANDAR

Department of Zoology, University of Durban-Westville, Durban

(With 7 figures)

[MS accepted 12 September 1991]

ABSTRACT

Eighteen specimens representing five species of Holothuroidea collected off the east coast

of South Africa in waters between 280 m and 1 050 m are recorded. Three species, Pseudo-

stichopus (P.) echinatus, Orphnurgus aspersignis, and Orphnurgus natalasper are new. The

remaining two, Mesothuria parva (Théel) and Orphnurgus insignis Fisher, are new to the fauna

of southern Africa, south of the Tropic of Capricorn. The material indicates a general paucity

of species of deep-sea benthic holothuroids on the continental shelf and slope off the east

coast of South Africa.

CONTENTS

MntROGUCHONMB ame cite sian “cee ett soa nee ee ae 159

IAI EWINOXOIS as 4eiey SPR A RE haa sce ce, ce Pe aD er RR Ee 160

SPECICSHIS tar rsmrciern Toene eras or sa Pha geste EN ene ntas 160

SVWStemlaticACCOUNE ak cee le enc aes Steere oe 160

Order Aspidochirotida Grube, 1840 ............. 160

Orndemelasipodidamlihcel SS eas eee 167

PNCKMOWICAUSEMENES nb ee oS 6 cee cet Sele ie ee eet el 179

INCTOREM CES meinen mets ra cro tiscelaas Mllomes o oew we ee uretren 179

INTRODUCTION

Eighteen specimens of deep-sea holothuroids taken by the South African

Museum’s R.V. Meiring Naude cruises in 1975 and 1976 are recorded. The

specimens represent three genera and five species, of which three species are

new and two are new records for the southern African region. The material

originates from localities on the continental shelf and slope from just south

of the Mozambique—Natal border, from depths ranging between 280 m and

1050m. The other echinoderms collected by the same cruises were

recorded by A. M. Clark (1977). Full station data and a map of the survey

area, as well as background information concerning the cruises, can be found

in Louw (1977).

The material is deposited in the South African Museum under SAM-A

catalogue numbers. The five species and the stations are recorded below.

Ann. S. Afr. Mus. 101 (7), 1992: 159-180, 7 figs.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

METHODS

The specimens were studied according to conventional methods outlined by

other writers such as Fisher (1907), Deichmann (1948), and Rowe & Doty

(1977). The spicules were removed in antiformin (see Mahoney 1966), washed in

two changes of distilled water and illustrated with a camera lucida. For scanning

electron microscopy the spicules were transferred directly from distilled water to

absolute alcohol and mounted together with a little alcohol on to a specimen

stub to which they normally stick once the alcohol evaporates. They were then

sputter-coated with gold for 5 minutes at 30-40 mAmp, and photographed

using a Philips SEM 500.

SPECIES LIST

SM station No. of

no. specimens

ORDER ASPIDOCHIROTIDA

Family Synallactidae

Mesothuria parva (Théel, 1886) 5

Pseudostichopus (P.) echinatus sp. nov. 38 it

ORDER ELASIPODIDA

Family Deimatidae

Orphnurgus aspersignis sp. nov. WZ 5

UY 1

Orphnurgus insignis Fisher, 1907 2 2

74 6

Orphnurgus natalasper sp. nov. a 1

SYSTEMATIC ACCOUNT

Order ASPIDOCHIROTIDA Grube, 1840

Family Synallactidae Ludwig, 1894

Diagnosis (after Ludwig 1894: 8, 26)

Aspidochirotid holothurians with a flattened, rarely cylindrical body with

pedicels in three ventral ambulacra and papillae dorsally. Tentacular ampullae

or Cuvierian organs absent. Stone canal usually connected to body wall. Longi-

tudinal muscles generally undivided. Respiratory trees usually not in association

with rete mirabile. Spicules of body-wall tables, occasionally C-shaped bodies,

rarely buttons, or absent.

Remarks

This is a cosmopolitan family comprising mainly deep-sea forms. Approxi-

mately 15 genera are currently recognized (Pawson 1965) and of these only the

—————

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 161

genus Synallactes, represented by two species, has hitherto been recorded from

southern Africa. The material at hand contains two further species that belong

in the genera Mesothuria and Pseudostichopus.

Genus Mesothuria Ludwig, 1894

Mesites Ludwig, 1893: 79. Type species M. multipes Ludwig (nomen nudum).

Mesothuria Ludwig, 1894: 31. Fisher, 1907: 679. Deichmann, 1930: 91.

Zygothuria Perrier, 1898: 126.

Allantis Hérouard, 1902: 18.

Diagnosis (partly after Fisher 1907: 679)

Cylindrical to subcylindrical synallactids with a slightly flattened ventral

surface. Mouth and anus usually terminal. Brim or anal furrow absent. Ten-

tacles 12-20. Pedicels on ventrolateral ambulacra usually well developed, on

mid-ventral ambulacrum small or absent, small, scattered and papilliform dor-

sally. Gonad in a single tuft. Tables with a circular disc with large perforations

and a spire terminating in one or more toothed or smooth apices.

Type species. Mesothuria multipes Ludwig, 1894 (by monotypy).

Remarks

The genus Mesothuria at present comprises about 25 nominal species of

which only M. parva (Théel) is here recorded for the first time from southern

Africa. This is also the first record of the occurrence of this species from the

Western Indian Ocean. Mesothuria lactea, described by Heding (1940) from ‘off

the west coast of South Africa’ at 30°34,9’S 6°E, is from too far offshore to be

regarded as strictly southern African.

Mesothuria parva (Théel, 1886)

Figs 1, 6A

Holothuria murrayi var. parva Théel, 1886: 186, pl. 9 (fig. 2), pl. 16 (figs 4, 5).

Mesothuria parva Fisher, 1907: 686, pl. 71 (fig. 2, 2a—c).

Diagnosis (after Fisher 1907: 686)

Medium-sized, cylindrical species, up to 100 mm long. Colour, in alcohol,

yellowish white, greyish brown or purplish black. Pedicels of variable size, scat-

tered all round, largest in ventrolateral ambulacra, few and small in mid-ventral

ambulacrum, numerous, scattered and papilliform dorsally. Table disc large,

with a large central hole and several marginal holes; spire of three non-diverging

pillars with a compact, toothed, often tripartite apex and a single cross bar.

Pedicels with small tables, large end plates but no supporting rods.

Previous southern African record

None.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material examined

SAM-A23434, off Sodwana Bay, R.V. Meiring Naude, SM 15, 27°31,5'S

32°45,6'E, 25 May 1975, 280-454 m, beam trawl, 2 specimens.

Description

Lengths 75 mm and 37 mm. Colour, in alcohol, dull greyish brown, darker

ventrally. Podia more sparsely distributed mid-ventrally, absent just behind

collar. Polian vesicles paired. Body-wall spicules in the form of tables of two

distinct sizes. Discs of large tables (Figs 1A, C, 6A) (0,100—0,135 mm) overlap-

ping, marginal holes 6—24; spire (0,078—0,113 mm) of three (occasionally fused)

pillars, a single cross bar situated near disc, and a compact, tripartite or irregular

apex, occasionally pierced by a minute hole. Smaller tables (Fig. 1B) less

common, discs 0,055—0,095 mm, marginal holes 6-18. Anal region with large

periproctal plates (Fig. 1D). Pedicels with small tables and large (0,27-

0,30 mm) end plates. Tentacles with smooth to spinulated rods, occasionally

perforated at extremity (Fig. 1E).

Fig. 1. Mesothuria parva (Théel). SAM-—A23434. Spicules and other internal structures.

A. Large tables from dorsal body wall. B. Small tables from ventral body wall. C. Large table

from side. D. Periproctal plate. E. Tentacle rods. F. Gonadal tubule. G. Madreporite.

H. Part of calcareous ring. Abbreviation: r—radial plate.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 163

Remarks

The present material differs from that described by both Théel (1886) and

Fisher (1907) in the absence of a marked difference in size between the ventro-

lateral pedicels and those from elsewhere on the body. However, its spicules are

identical to those illustrated by the above authors. Tables with perforated apices

were also observed by Fisher. According to Théel, the smaller tables are com-

moner in the body wall, whereas Fisher stated that they are confined to the

pedicels. In the present material both small and large tables occur in the body

wall but the former are not at all common, whereas the pedicels only contain

small tables. Although the maximum size of the table discs (0,135 mm) in the

present material corresponds roughly to that of Fisher’s material (0,12 mm), the

spires are much taller (max. size 0,113 mm) compared to 0,085 mm recorded by

him. Unless Fisher recorded the mean, this difference might suggest geographi-

cal variation. The mean for the present material is 0,096 mm.

The species is closely related to M. murrayi (Théel); Fisher (1907) pointed

out that the differences in the calcareous deposits between the two are possibly

attributable to the substrate, M. murrayi living in softer bottoms than M. parva.

However, he mentioned that, due to the absence of intermediate forms and the

lack of difficulty in separating them, they should be regarded as separate

species.

Distribution

Off Sodwana Bay, Natal, 280-454 m. Also recorded from Hawaii, Admi-

ralty Island, and south-east Africa.

Habitat

Globigerina ooze, sand grains and broken shells.

Genus Pseudostichopus Théel, 1886

Pseudostichopus Théel, 1886: 169. Fisher, 1907: 691. Heding, 1940: 356.

Diagnosis (after Fisher 1907: 691)

Body form cylindrical, subcylindrical or flattened. Podia of diverse size,

usually large in the ventrolateral ambulacra, absent or rudimentary in the odd

ambulacrum. Anus situated in a perpendicular furrow at posterior end; anal

teeth absent. Body wall usually encrusted with foreign material such as foramini-

ferans, pteropod shells, other shell fragments, sponge spicules and sand grains.

Type species. Pseudostichopus mollis Théel, 1886 (by subsequent desig-

nation of Fisher 1907: 691).

Remarks

Heding (1940) referred this genus, together with several others previously

occupying isolated positions in the Synallactidae, to the family Gephyrothuri-

idae, diagnosed by Koehler & Vaney (1905) for their monotypic Gephyrothuria

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

alcocki. A similar form from the Eastern Pacific was described as Himasthle-

phora glauca by H. L. Clark (1907). Gephyrothuria alcocki was stated to be

molpadid-like, whereas H. glauca, which possessed a tail-like appendage, was

described as a molpadiid. According to Clark, both H. glauca and G. alcocki

may be congeneric, whereas Rowe (pers. comm.) considers that they may even

be conspecific. If this is true, Pseudostichopus cannot be placed in the Gephyro-

thuriidae and should therefore, at least tentatively, be transferred back to the

Synallactidae.

Pseudostichopus is a fairly large cosmopolitan genus subdivided by Heding

(1940) into the subgenera Pseudostichopus and Trachostichopus, each containing

six nominal species. Pseudostichopus (P.) echinatus, here described as new and

represented in the collection by a single specimen, is the only representative of

the genus in southern Africa.

Subgenus Pseudostichopus Théel, 1886

Diagnosis (after Heding 1940: 356)

Cylindrical species with no or little distinction between the ventrolateral

and other podia. Spicules absent from body wall, including anal region.

Type species. Pseudostichopus mollis Théel, 1886 (by original designation of

Heding, 1940: 356).

Remarks

This subgenus includes all cylindrical species without body-wall spicules and

with little or no distinction between the ventrolateral and other podia. When

proposed by Heding (1940), the subgenus contained six nominal and one

unnamed species. The single specimen in the SAM collection from off Natal is

identical to Heding’s unnamed species from off the East African coast. It is here

described as P. (P.) echinatus sp. nov.

Pseudostichopus (Pseudostichopus) echinatus sp. nov.

Fig. 2

Pseudostichopus (Pseudostichopus) sp. Heding, 1940: 360, text-fig. 16.

Diagnosis

A medium-sized species up to 65 mm long. Body wall encrusted with shells,

foraminiferans, sand grains, etc. Tentacles ?17. Podia scattered, only slightly

longer in ventrolateral radii. Stone canal rudimentary or absent. Spicules

restricted to tentacles, in the form of large (up to 0,3 mm), irregularly knobbed

rods. Body wall, podia, respiratory trees and gonad without spicules.

Etymology

The specific name is derived from echinos (Greek): sea urchin, with

reference to the echinoid appearance of the holotype.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 165

Material examined

Holotype, SAM-—A23435, SAM, R.V. Meiring Naude, SM 38, off Natal

coast, 28°21,9’S 32°34,6’E, 28 May 1975, 775-825 m, beam trawl.

Description

Specimen (Fig. 2A) partially eviscerated, most of alimentary canal lost.

Length 55 mm, breadth in mid-body 12 mm. Colour, in alcohol, a uniform pale

greyish brown. Body almost cylindrical, dorsal surface arched, ventral less so.

Mouth small, ventral. Tentacles white, peltate, about 17, all except three

retracted (exact number not determined for fear of damage to holotype). Anus

subventral, situated in a conspicuous vertical furrow; no anal papillae. Body wall

encrusted with Globigerina ooze, shell fragments, coral debris, sand grains and

ha Le anes

0,05 mm

2mm

Fig. 2. Pseudostichopus (P.) echinatus sp. nov. SAM—A23435. Holotype.

A. Entire animal. B. Part of calcareous ring. C. Tentacle rods. Abbreviation:

mdir—mid-dorsal interradial plate.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

pteropod (?Creseis) shells, the latter projecting perpendicularly from surface

giving specimen a prickly appearance, not unlike that of heart urchins. Body

wall remarkably thin, especially ventrally.

Podia thin, non-retractile, barely visible to unaided eye, scattered but more

numerous ana slightly longer in the ventrolateral ambulacra, shorter dorsally, few

and small in mid-ventral ambulacrum, where they are difficult to distinguish from

encrustations; discs small, brownish; end plates and supporting rods absent.

Calcareous ring (Fig. 2B) delicate, radial plates dissimilar, usually squarish,

concave posteriorly but either scalloped anteriorly or with a single median

notch; interradial plates more uniform, four times as wide as long, with a short

anterior triangular median projection and a concave posterior margin. No

tentacular or podial ampullae. Polian vesicle single, mid-ventral, sacciform.

Stone canal not observed (?rudimentary or absent). Longitudinal muscles thick,

unpaired. Respiratory trees short, well branched and with common origin, left

tree shorter than right.

Gonad (?testis) attached in anterior third of body, developed as two tufts of

unbranched tubules, each constricted serially along entire length. Cloaca wide,

suspensors well developed.

Spicules. Spicules (Fig. 2C) confined to tentacles, in the form of numerous

slightly curved, often branched rods of varying size (up to 0,3 mm long) and

thickness, each ornamented with numerous knobs of varying size and density;

rarely rods thin and smooth but occasionally with holes resulting from fusion of

once parallel branches.

Remarks

There are no differences between the present specimen and Heding’s (1940)

Pseudostichopus (P.) sp. Their size, form, encrustations, distribution of podia

and the presence of identical spicules (confined to the tentacles) indicate that

they belong together.

A species that closely resembles P. (P.) echinatus is P. (Trachostichopus)

trachus, originally described by Sluiter (1901) from the West Pacific but also

recorded by Mitsukuri (1912), Mortensen (1917), and Heding (1940). According

to Heding, both Mitsukuri’s and Mortensen’s descriptions of P. (T.) trachus

refer to another species as they do not correspond with that of the type.

Pseudostichopus (P.) echinatus resembles Sluiter’s and Heding’s descriptions of

P. (T.) trachus in the restriction of the spicules to the tentacles but differs in its

cylindrical form, absence of thickened flanks, lack of a sharp distinction between

the ventral podia, and different form and size of the tentacle spicules. In P. (T.)

trachus the spicules have fewer knobs, lack perforations and measure 0,13 mm

in length, unlike those of P. (P.) echinatus which are heavily knobbed,

occasionally perforated and have an upper size limit of 0,3 mm.

Heding’s P. (T.) trachus was taken off the central East African coast at

638-977 m and his Pseudostichopus (P.) sp. also came from more or less the

same locality at 693m. Despite this, Heding not only emphasized the

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 167

differences between the two species but went so far as to refer them to different

subgenera. Therefore it does not seem likely that the two forms are conspecific.

Pseudostichopus (P.) echinatus differs from the six species included in the

subgenus Pseudostichopus by Heding as follows: the type species P. (P.) mollis

Théel, 1886, has at least the ventrolateral pedicels arranged in rows and, in

addition to the tentacles, has spicules also in the gonad and pedicels; P. (P.)

pustulosus Sluiter, 1901, also has the ventrolateral pedicels arranged in rows but

the stone canal is distinct and the genital tubules branched; P. (P.) unguiculatus

Ohshima, 1915, has deposits in the tentacles, gonad and pedicels; P. (P.)

lapidus Hérouard, 1923, has very few pedicels, which are united by some sort of

webbing posteriorly; P. (P.) globigerinae Hérouard, 1923, has no spicules,

whereas P. (P.) marenzelleri Hérouard, 1923, has podia apparently arranged in

bands and spicules in only the genital tubules.

Distribution

Type locality and East African coast, south of the equator.

Order ELASIPODIDA Théel, 1881

Remarks

This is an exclusively deep-sea order of usually large, often bizarre

holothurians with peltate tentacles and no respiratory trees. In an excellent

revision of the order, Hansen (1975) assembled the approximately 110 species

recognized by him into the suborders Deimatina and Psychropotina. Both

suborders are represented in southern Africa by a total of 15 species; however,

only the former is present in the collection at hand.

Suborder DEIMATINA Hansen, 1975

Diagnosis (after Hansen 1975: 14)

Spicules perforated plates (or derivatives of these), spatulated crosses, rods

or wheels; no primary crosses with arrested development of dichotomous div-

isions. Papillae usually numerous and large.

Remarks

This suborder comprises elasipodids with numerous, usually large papillae.

It includes the families Deimatidae and Laetmogonidae, both with represen-

tatives in southern Africa. However, only the former is represented in the

present collection.

Family Deimatidae Ekman, 1926

Diagnosis (after Hansen 1975: 15)

Spicules varying from perforated plates and spatulated primary crosses, to

spatulated, spinous or reduced and deformed bodies; wheels absent. Gonad

consisting of few, unbranched, sacciform tubules.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This family, characterized by the absence of wheels, includes the genera

Deima, Oneirophanta and Orphnurgus, collectively containing eight species. All

three genera are represented in southern Africa, the former two each by a single

species recorded by Hansen (1975), but not present in the collection here

examined. The genus Orphnurgus, first recorded from southern Africa by A. M.

Clark (1977), is represented by three species, of which two are new.

Genus Orphnurgus Théel, 1879

Orphnurgus Théel, 1879: 8; 1882: 82. Hansen, 1975: 38 (synonymy).

Diagnosis (after Hansen 1975, modified herein)

Tentacles 15-20, non-retractile, discs with ramified processes. Circumoral

papillae absent. Spicules spatulated crosses and/or rods of greatly varying shape,

either spatulate, spindle-shaped, smooth with dichotomous ramifications, or

spinous, often a combination of two types.

Type species. Orphnurgus asper Théel, 1879 (by monotypy).

Remarks

This genus was erected by Théel (1879) for the West Indian species

O. asper, characterized by only spinous rods in the body wall. Since then the

following eight Indo-West Pacific species, all referable to Orphnurgus, have

been described: Orphnurgus glaber Walsh, 1891 (Koehler & Vaney, 1905): with

smooth elongate rods with short, sometimes spinous, terminal ramifications;

Orphnurgus invalidus Koehler & Vaney, 1905: with mainly spatulate rods;

Orphnurgus insignis Fisher, 1907: with crosses and dichotomously ramified rods;

Orphnurgus vitreus Fisher, 1907: with spatulate rods and sometimes spatulate

crosses; Orphnurgus rigidus Ohshima, 1915: with spatulate rods only; Orph-

nurgus bacillus Cherbonnier & Feral, 1981: with rods with spinulated ends;

Scotodeima protectum Sluiter, 1901: with spatulate crosses only; and Amphi-

deima investigatoris Koehler & Vaney, 1905: with spatulate rods only.

These species were further distinguished by the number and distribution of

pedicels and papillae in either single or double rows and by the presence or

absence of any transformation of the spicules ventrally.

In his revision, Hansen (1975) recognized only O. asper, O. glaber,

O. vitreus and O. protectus, and relegated the remaining four species to the

synonymy of QO. glaber because of the ‘presence of many intergradations’

(Hansen 1975: 40). This action resulted in O. glaber being represented by

several morphological types showing numerous geographical variations, not only

in the spicules but also in the number and distribution of pedicels and papillae.

A study of the present material reveals the presence of O. insignis (sensu

Fisher 1907) and two further morphological types, also referable to Orphnurgus.

Of the latter, one is intermediate between O. asper and O. insignis, having both

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 169

O. asper and O. insignis type of rods, whereas the other, which has double rows

of pedicels and dorsal papillae, is characterized by only O. asper type of rods in

the dorsal body wall and a mixture of O. asper and O. insignis type in the

ventral body wall. However, all three types show similar transformation of

deposits ventrally. Because of these intergradations, the writer was at first

inclined to refer all three types to O. asper and to relegate O. glaber (sensu

Hansen) and its synonyms to the synonymy of this species. Such a step would

have not only affected the status of O. glaber but also of O. vitreus and

O. protectus, characterized by spatulate rods and/or crosses, since spatulate rods

are the basic type of deposit in O. glaber (sensu Hansen) and spatulate crosses

also occur sporadically in some forms of the latter species. Rowe (pers. comm.),

who examined my slides, is of the opinion that the three types represent

different species and is inclined to consider Hansen’s decision as too sweeping.

This problem would be further aggravated if all nominal species of Orphnurgus

and the southern African material are lumped under O. asper, which would then

assume the status of a superspecies.

Since the southern African forms differ not only in their deposits but also in

some other significant features, all three types are here described as distinct

species, namely, O. insignis Fisher and two others, O. aspersignis sp. nov. and

O. natalasper sp. nov.

KEY TO THE SOUTHERN AFRICAN SPECIES OF ORPHNURGUS

1. Pedicels and dorsal papillae in double rows on each side of body; dorsal

spicules exclusively short, spinous, often perforated, rods ...............

53055555 0 ee ee Orphnurgus natalasper sp. nov.

— Pedicels and dorsal papillae in single rows on each side of body; dorsal

spicules either exclusively smooth rods with terminal dichotomous

ramifications or accompanied by short, spinous, non-perforated rods .... 2

2. Dorsal spicules exclusively smooth rods with open, often spiny, dichot-

omous ramifications; no short spinous rods in integument; papillae elongate,

OurcomenMOnMM... Ls. ses eee. Ses Orphnurgus insignis Fisher, 1907

— Dorsal spicules with short, spinous, non-perforated rods in combination

with dichotomously ramified rods; papillae short, stout, often rudimentary

100 60.0.9 6S Sie eee ee Orphnurgus aspersignis sp. nov.

Orphnurgus aspersignis sp. nov.

Figs 3, 6B

Orphnurgus glaber (non Walsh, 1891) A. M. Clark, 1977: 146.

Diagnosis

A medium-sized species up to about 100 mm long. Tentacles 15-20 with

bright yellow discs. Ventrolateral pedicels in single rows of 15-20 on each side,

mid-ventral pedicels absent. Ventrolateral papillae 6-10 on each side, usually

larger than dorsal papillae, the latter in a single row of 7-20 on each side of mid-

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Orphnurgus aspersignis sp. nov. SAM—A23436. Holotype. Spicules. A. Anterodorsal

body wall. B.Mid-dorsal body wall. C. Posterodorsal body wall. D. Anteroventral body

wall. E. Mid-ventral body wall. F. Posteroventral body wall. G. Papillae rods. H. Pedicel

rods. I. Rods from tentacle tips. J. Cross from body wall. All drawn to same scale.

dorsal line; papillae generally short, often stout or rudimentary. Spicules

numerous, short, non-perforated, spinous rods and a few elongate smooth rods

with open, often spinous, dichotomous ramifications; ventral spicules always

sturdier and transformed into dumbbell-shaped, rounded, ellipsoidal or

amorphous bodies. Smooth primary crosses with arrested dichotomous divisions

occasionally present.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES yi

Etymology

The specific name is derived from a combination of asper and insignis

because of the presence of spicules similar to those of O. asper Théel and

O. insignis Fisher.

Previous record

Off Natal coast, 1 000-1 200 m, as O. glaber A. M. Clark, 1977.

Material examined

Holotype, SAM—A23436, off Sibaya, Natal, R.V. Meiring Naude, SM 72,

27°17,8'S 33°04,5'E, beam trawl, 20 May 1976, 1050 m. Paratypes, SAM-

A23437, same data as holotype, 4 specimens. Paratype, SAM—A23438, off

Sodwana Bay, Natal, R.V. Meiring Naude, SM 77, 27°31,6'S 32°50'E, heavy

dredge, 21 May 1976, 780 m, 1 specimen.

Description

Holotype. Length 90 mm, breadth in mid-body 20 mm. Anterior and

posterior ends of more or less equal width, only slightly less than that of mid-

body. Colour yellowish white with bright yellow to orange tips to tentacles,

pedicels and larger papillae. Form subcylindrical, ventral surface flattened,

dorsal somewhat arched. Odd ambulacrum marked by absence of podia but

presence of a faint longitudinal line, indicating position of mid-ventral

longitudinal muscle. Skin opaque to somewhat translucent, but not as much as

in O. insignis. Body wall thin and rough to the touch.

Mouth terminal but ventrally directed, collar absent. Tentacles 20, non-

retractile, stalks thin, white, discs with contracted ramifications and hence

distinctly peltate or subglobose in appearance. Anus terminal but slightly dorsal,

anal papillae absent.

Ventrolateral pedicels in single rows, 15 on the left and 18 on the right,

conical, non-retractile, projecting horizontally from body, longest anteriorly

(8 mm), decreasing in size posteriorly. Ventrolateral papillae 10 on each side,

placed in a single row, short (max. length 6 mm), stout, often quite rudi-

mentary, never filiform. Dorsal papillae 16 on the left and 13 on the right, one

row in each side of dorsal mid-line, papillae short (max. length 8 mm), often

stout or quite rudimentary.

Calcareous ring delicate, poorly calcified. Tentacular ampullae absent.

Water ring well developed, proximal ends of radial canals broad. Polian vesicle

single, elongate (14 mm), reaching level of gonad, originating from water ring

slightly to left of left dorsal radius. Madreporite sieve-like, external, situated

between the most anterodorsal papillae. Longitudinal muscles as thin double

strands. Gonad in two clusters of small globular sacs of varying size, each cluster

resembling a bunch of grapes.

Spicules. Dorsally simple non-perforated short spinous rods (0,06—

0,15 mm) of the O. asper type but with spines usually developed only at

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

extremities, and elongate rods (0,29-0,47 mm) of the O. insignis type with

terminal, often spiny, open dichotomous ramifications (Figs 3A—C, 6B);

ventrally sturdier rods (0,20—-0,63 mm), transformed into huge dumbbell-

shaped, ellipsoidal, rounded or amorphous bodies with spinous extremities, the

spines representing pointed ends of otherwise fused ramifications (Figs 3D, F,

6B). Simple primary crosses (0,285 mm long) with arrested dichotomous

ramifications also present (Fig. 3K). Pedicels with elongate, slender to stout,

smooth rods with short spinous terminal ramifications (Fig. 3H). Papillae with

small slender rods with dichotomous, usually non-spinous ramifications

(Fig. 3G). Tentacles with rods similar to those of papillae but often with spinous

ramifications (Fig. 3J), those of tentacular stalks occasionally Y-shaped and up

to 0,45 mm long (Fig. 31).

Paratypes. Size range 65-95 mm. Colour and body wall as in holotype.

Tentacles 15-19 (two specimens each with 15, one with 18, and two each

with 19). Ventrolateral pedicels 17—20 on each side (max. length 15 mm).

Ventrolateral papillae 6—9 on each side (max. length 10 mm); dorsal papillae

always in single rows, 8-16 on each side (max. length 8 mm). Spicules as in

holotype.

Remarks

This species is well characterized by the presence of both spinous rods and

elongate rods with dichotomous ramifications. The simultaneous presence of

both these deposits renders the new species intermediate between the West

Atlantic O. asper Théel and the Indo-West Pacific O. insignis Fisher. No species

with a similar combination of spicules has yet been described from other parts of

the world. Orphnurgus aspersignis is, however, closer to the sympatric

O. insignis since it shares with it a low number of dorsal papillae arranged in

single rows, identical rods with dichotomous ramifications and a_ similar

transformation of the ventral deposits. Orphnurgus asper, on the other hand,

has a high number of dorsal papillae arranged in double rows, different type of

spinous rods, lacks ramified rods and shows no transformation of the deposits.

As in the other species of the genus, the number of tentacles in the new

species varies from 15 to 20 and there is no apparent correlation between

tentacle number and size of the specimen since the two largest specimens have

15 and 20 tentacles.

No general increase in the number of ventrolateral pedicels is apparent with

increase in the size of the specimens, perhaps because the sample size

(6 specimens) is too small or the size range of the specimens too narrow

(65-95 mm) for any effective analysis. However, such a correlation, with

notable exceptions, was reported for O. insignis by Fisher (1907) and for

O. glaber (s.1.) by Hansen (1975).

As in O. glaber (s.1.), there is a general increase in the number of both

dorsal and ventrolateral papillae with increased size of the specimens, the larger

specimens having 10-12 pairs of each type, whereas the smaller ones have

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES jes)

7-8 pairs, with a 75 mm specimen being an exception with 8 pairs of ventro-

lateral and up to 11 pairs of dorsal papillae.

There is also, in each specimen, a close correlation between the number of

ventrolateral and dorsal papillae, with one or two exceptions. Generally a high

number of ventrolateral papillae is accompanied by a more or less similar

number of dorsal papillae. However, no correlation exists in each specimen

between the number of papillae and pedicels.

The short spinous rods are derived from tiny bow-shaped deposits that

come to bear a cluster of spines, usually at each of the four extremities or, more

exceptionally, all round. As in O. glaber, the basic spicules are found dorsally

but both the spinous and dichotomously ramified rods become robust postero-

dorsally. The ventral spicules are always robust but tend to increase in sturdiness

posteriorly while becoming progressively deformed. Such transformation is not

dependent on age since all specimens, without exception, show similar

transformation. The type of transformation, as in O. glaber (sensu Hansen),

does not involve the extremities of the spicules, as these are nearly always

preserved.

A juvenile specimen, referred to O. glaber by A. M. Clark (1977), was

described as having spicules similar to those of O. asper and O. insignis.

Regrettably this specimen could not be located by the South African Museum in

the material returned by Clark. However, judging from the data given by Clark,

the specimen in question was part of the present collection and hence O. glaber

A. M. Clark (non Walsh) is here relegated, without doubt, to the synonymy of

O. aspersignis.

Distribution

Off Natal coast between Richards Bay and Kosi Bay, 780-1 200 m.

Orphnurgus insignis Fisher, 1907

Figs 4, 7A

Orphnurgus insignis Fisher, 1907: 702, pl. 73 (fig. 1), pl. 77 (figs 1-3). Ohshima, 1915: 234;

1916-1919, 3 figs. Cherbonnier & Feral, 1981: 361, fig. 2A, K.

Orphnurgus glaber Hansen, 1975: 39 (part.), text-fig. 13 (28-33, 40-45).

non Orphnurgus glaber Walsh, 1891. A. M. Clark, 1977: 146 (= O. aspersignis sp. nov.).

Diagnosis (from Fisher 1907, modified herein)

Small to medium-sized species, reaching a length of 160 mm. Tentacles

15-20. Ventrolateral pedicels 14-24 on each side in single rows, rarely about 30

in double rows on each side; mid-ventral pedicels rarely present. Ventrolateral

papillae 4—24 on each side. Dorsal papillae 4-36 on each side in single or double

rows per dorsal ambulacrum. Papillae elongate and filiform, never stout or

rudimentary. Spicules dorsally crosses and rods with dichotomously ramified

ends and small rods with rudimentary ramifications; ventrally robust rods with

enlarged spiny ends and large ellipsoidal to rounded bodies plus all

intermediates between these two types.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material examined

SAM-A23441, R.V. Meiring Naude, SM 72, 27°17,8'S 33°04,5'E, 20 May

1976, 1 050 m, beam trawl, 2 specimens. SAM—A23439, SM 74, 21 May 1976,

860 m, beam trawl, 6 specimens.

Description

SAM-—A23439. Specimens flattened. Maximum length 60 mm. Colour white

with yellowish tentacles and pale yellow tips to pedicels. Tentacles 19-20.

Ventrolateral pedicels 14—20, in single rows (max. length 8 mm). Ventrolateral

papillae approximately 4-10 on each side (max. length 12 mm). Dorsal papillae

2-10 in single rows per dorsal radius (max. length 10 mm).

Body wall thin, translucent. Dorsal deposits a few crosses and dichot-

omously ramified rods (0,27—-0,88 mm), sturdier posteriorly but with spiny

extremities (Figs 4A, 7A). Ventral deposits (0,81-1,08 mm), resembling those

of O. aspersignis (Figs 4B—C, 7A).

Fig. 4. Orphnurgus insignis Fisher. SAM-A23439. Body wall spicules. A. Rods from

mid-dorsal body wall. B. Rods from anteroventral body wall. C. Rods from

posteroventral body wall. All drawn to same scale.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 73

SAM-—A23441. Maximum length 70 mm, form subcylindrical. Tentacles 20

in one, 13 (?15) in the other. Ventrolateral pedicels 18-21 (max. length 8 mm).

Ventrolateral papillae 7-10 (max. length 14 mm). Dorsal papillae 7—20 in single

rows, One pair radius (max. length 16 mm). Spicules as above.

Remarks

In possessing only crosses and dichotomously ramified rods and their

derivatives, the southern African material is identical to Fisher’s (1907) species

from Hawaii and Ohshima’s (1915) from Japan. However, it shows some

geographic variations, differing from the Hawaiian form in its smaller size and

lower number of papillae, and from the Japanese form in having only one row of

dorsal papillae on each side. Hence the diagnosis of the species is here modified

to take in the new form. The aberrant Hawaiian specimens of Fisher were

extraordinary in possessing mid-ventral pedicels not encountered elsewhere in

the genus. Further, they have a higher number of ventrolateral pedicels in one

or two rows and double rows of dorsal papillae.

As in O. aspersignis and in other species of the genus, there is no

correlation between the size of the specimens and the number of tentacles. The

largest individual (70 mm) has only 13 (?15) tentacles, whereas the majority

(including the smallest) have 20. There is also no correlation between the size of

the specimens and the number of pedicels, which varies from 15 to 21. Although

the highest pedicel number (21 pairs) is found in the largest individual (70 mm),

the two smallest individuals (48 mm) both have 20 pairs each. Perhaps, here

again, the sample is too small (8 specimens) and the size range too narrow

(48-70 mm) for any effective analysis.

As in O. aspersignis, the number of ventrolateral and dorsal papillae

generally increases with the increase in size of the specimens. However, the

smallest individual has a relatively high number of dorsal and ventrolateral

papillae—9 and 10 pairs respectively. There is also a close correlation, in each

specimen, between the number of ventrolateral and dorsal papillae, except in

the largest specimen, which has twice as many dorsal as ventrolateral papillae.

In O. insignis, as in other species of the genus, the basic type of deposit is

found in the dorsal body wall and sturdier deposits, in various stages of

transformation, occur ventrally. As in O. .aspersignis and other related species,

the transformation does not affect the extremities of the deposits.

Orphnurgus insignis most closely resembles O. glaber from the Bay of

Bengal, which has rods with few terminal ramifications and rather sturdy rods

with enlarged spinous extremities. In O. glaber (s. s.), however, there are no

ellipsoidal or dumbbell-shaped deposits. It is likely that the condition in

O. insignis is a modification of that present in O. glaber.

Distribution

Off Natal, between Cape St Lucia and Kosi Bay, 860-1 050 m, south-east

Africa, Japan, Hawaii, and Philippines.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Orphnurgus natalasper sp. nov.

Figs 5, 7B

Diagnosis

Tentacles 19. Ventrolateral pedicels about 40 on each side in alternating

double or triple rows. Mid-ventral pedicels absent. Ventrolateral papillae about

20 on each side in a single row. Dorsal papillae about 43-44 in alternating

double rows in each dorsal radius, decreasing to single rows posteriorly. Dorsal

deposits very spinous, often perforated rods with spines frequently present

throughout length of rod; ventral deposits sturdier rods with spiny ends and

huge dumbbell-shaped deposits with spiny extremities.

Etymology

The specific name is derived from a combination of Natal, the type locality,

and asper because of rods of the O. asper Théel type.

Fig. 5. Orphnurgus natalasper sp. nov. SAM-—A23440. Holotype. Spicules.

A. Rods from mid-dorsal body wall. B. Rods from anteroventral body wall.

C. Rods from mid-ventral body wall. D. Rods from posteroventral body wall.

All drawn to same scale.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 197,

Material examined

Holotype, SAM-—A23440, off Lake St Lucia, Natal, R.V. Meiring Naude,

SM 74, 27°38,6'S 32°52,6'E, beam trawl, 2 April 1976, 860 m.

Description

Length 82 mm, breath in mid-body 16 mm. Form subcylindrical. Tentacles

19, discs with extremely contracted ramifications giving the tentacles a distinctly

peltate appearance. Ventrolateral pedicels short (max. length 10 mm), stout,

about 40 on each side, arranged alternately in double or triple rows. Ventro-

lateral papillae about 20 on each side with some indication of a zigzag arrange-

ment, size equal to that of pedicels. Dorsal papillae about 43-44 on each side in

Fig. 6. SEM micrographs of body-wall spicules. A-B. Mesothuria parva (Théel). A. Large

tables. B. Small tables. C-—D. Orphnurgus aspersignis sp. nov. C. Dorsal body wall.

D. Ventral body wall. Scale bars = 50 um.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

alternating double rows per ambulacrum, varying from short, rudimentary to

long and filiform (max. length 10 mm). Anterior papillae with light yellow tips.

Body wall relatively thick, opaque.

Spicules. Dorsally short spinous rods (0,14—0,30 mm long), spines long,

usually undivided but occasionally bifid or even trifid at apex. Rods frequently

Fig. 7. SEM micrographs of body-wall spicules. A-B. Orphnurgus insignis Fisher.

A. Dorsal body wall. B. Ventral body wall. C-D. Orphnurgus natalasper sp. nov. C. Dorsal

body wall. D. Ventral body wall. (X—scale Y; others—scale Z). Scale bars = 50 wm.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 179

perforated, the holes resulting from fusions of distal ends of some long spines

(Figs 5A, 7B). Ventrally sturdier rods (Figs SB—D) with few spiny projections,

slender rods with spinous terminal ramifications, and huge elongate to

dumbbell-shaped bodies, also with spinous extremities. Length of ventral

spicules 0,22—0,58 mm. Spatulate rods, crosses, ellipsoidal and amorphous

deposits absent. Pedicels with stout, elongate or tripartite rods with spinous

extremities. Tentacular discs and papillae with slender rods (0,22—0,35 mm

long) with spinous dichotomous ramifications.

Remarks

In the presence of double rows of ventrolateral pedicels and dorsal papillae

and the occurrence of different form of spinous rods, this species is distinct from

the other southern African species of the genus. Since double rows of pedicels

and papillae do occur sporadically in members of a single species within the

genus, it may be argued that O. natalasper is a variant of O. aspersignis.

However, the nature of the spinous rods and the absence of any dichotomously

ramified deposits support the taxonomic status of O. natalasper.

Whereas the spinous rods, with their frequent perforations, are reminiscent of

those of O. asper, the ventral spicules are somewhat like those of O. aspersignis;

hence O. natalasper appears intermediate between the two species. It differs from

O. asper not only in the nature of the ventral deposits (no transformation of

deposits occur in O. asper) but also in the high number of double-rowed pedi-

cels and the low number of dorsal papillae. The similarity of the spinous rods in

both species might suggest that O. asper probably developed from a form

resembling O. natalasper in which there was a loss of transformation of the

ventral deposits. It is possible that further intermediates, more closely approach-

ing the O. asper condition, may occur in waters of the Eastern Atlantic Ocean.

Distribution

Known only from the holotype, off Lake St Lucia, Natal coast.

ACKNOWLEDGEMENTS

I am indebted to Dr Naomi Millard and the late Dr T. H. Barry, formerly

of the South African Museum, for the opportunity to study this interesting

material. Dr Rowe, formerly of the Australian Museum in Sydney, is thanked

for his expert advice and guidance. Financial assistance from the University of

Durban-Westville is gratefully acknowledged.

REFERENCES

CHERBONNIER, G. & FERAL, J. P. 1981. Echinodermes: Holothuries. Résultats des Campagnes

Musorstom. I. Philippines 17: 357-412.

Ciark, A. M. 1977. The South African Museum’s Meiring Naude cruises. 4. Echinoderms.

Annals of the South African Museum 73 (6): 133-147.

Crark, H. L. 1907. The apodous holothurians. Smithsonian Contributions to Knowledge 35:

1-206.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

DEICHMANN, E. 1930. The holothurians of the western part of the Atlantic Ocean. Bulletin of

the Museum of Comparative Zoology at Harvard College 71 (3): 41-226.

DEICHMANN, E. 1948. The holothurian fauna of South Africa. Annals of the Natal Museum 11

(2): 325-375.

ExkMAN, S. 1926. Systematisch-phylogenetische Studien tiber Elasipoden und Aspidochiroten.

Zoologische Jahrbticher (Anatomie) 47: 429-540.

FISHER, K. W. 1907. The holothurians of the Hawaiian Islands. Proceedings of the United

States National Museum 32: 637-744.

HANSEN, B. 1975. Systematics and biology of the deep sea holothurians. 1. Elasipoda. Gala-

thea Report 13: 1-262.

HepinG, S. G. 1940. Die Holothurien der deutschen Tiefsee-Expedition. II. Aspidochirote

und Elasipode Formen. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf

dem Dampfer ‘Valdivia’ 1898-1899 24 (3): 317-375.

HEROUARD, E. 1902. Holothuries provenant des campagnes de la ‘Princesse-Alice’ (1892—

1897). Résultats des campagnes scientifiques accomplies par le Prince Albert I 1 (21): 1-61.

KOEHLER, R & Vaney, C. 1905. Holothuroidea. An account of the deep-sea Holotyuroidea

collected by the Royal Indian Marine Survey Ship ‘Investigator’. Jn: Echinoderma of the

Indian Museum. Calcutta: Trustees of the Indian Museum.

Louw, E. 1977. The South African Museum’s Meiring Naude cruises. 1. Station data 1975,

1976. Annals of the South African Museum 72 (8): 147-159.

Lupwic, H. L. 1893. Vorlaufigen Bericht uber die auf den Tiefsee Fahrten des ‘Albatross’

(Frukling 1891) im Ostlichen Stillen Ocean erbeuteten Holothurien. Zoologischer Anzei-

ger 16: 177-186.

Lupwic, H. 1894. Reports on an exploration off the west coasts of Mexico, Central and South

America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish

Commission Steamer ‘Albatross’ during 1891. 12. The Holothurioidea. Memoirs of the

Museum of Comparative Zoology at Harvard College 17 (3): 1-183.

MAHONEY, R. 1966. Laboratory techniques in zoology. London: Butterworth.

Mitsukur!, K. 1912. Studies on actinopodous Holothuroidea. Journal of the College of

Science, Imperial University of Tokyo 29 (2): 1-284.

Mortensen, T. 1917. Papers Dr Th. Mortensen’s Pacific Expedition (1914-16). I. Obser-

vations on protective adaptations and habitats, mainly in marine animals. Videnskabelige

Meddelsen fra Dansk naturhistorisk Forening i Kjobenhavn 69: 57-96.

OusHIMA, H. 1915. Report on the holothurians collected by the United States Fisheries

steamer ‘Albatross’ in the Northwestern Pacific during the summer of 1906. Proceedings of

the United States National Museum 48 (2073): 213-291.

OHSHIMA, H. 1916-1919. Northwestern Pacific holothurians collected by the ‘Albatross’. (An

expanded edition with numerous text-figures and 7 plates was published during these years)

Zool. Mag. 28-31: 230 pp (irregular pagination).

Pawson, D. L. 1965. The bathyal holothurians of the New Zealand region. Zoology Publica-

tions from Victoria University College 39: 1-33.

PERRIER, R. 1898. Sur les Holothuries recueillies par le “Travailleur’ et le ‘Talisman’. Compte

rendu hebdomadaire des séances de l Académie des sciences (3) 126: 1664-1666.

Rowe, F. W. E. & Doty, J. E. 1977. The shallow-water holothurians of Guam. Micronesica

1S ep AWVeZI0.

SLUITER, C. PH. 1901. Neue Holothurien aus der Tief-See des Indischen Archipels. Gesam-

melt durch die ‘Siboga’ Expedition. Tijdschrift der Nederlandsche dierkundige vereeniging

2 (7): 1-28.

THEEL, H. 1879. Preliminary report on the Holothuridae of the exploring voyage of H.M.S.

‘Challenger’ I. Bihang till K. Svenska vetenskapsakademiens handlingar 5 (19): 1—20.

Tuf&eL, H. 1882. Report on the Holothurioidea dredged by H.M.S. Challenger during the

years 1873-1876. Part I. Report on the scientific Results of the Voyage of H.M.S. ‘Chal-

lenger’, Zoology 4 (13): 1-176.

Tué£eL_, H. 1886. Report on the Holothurioidea dredged by H.M.S. Challenger during the

years 1873-1876. Part II]. Report on the scientific Results of the Voyage of H.M.S. ‘Chal-

lenger’, Zoology 14 (39): 1-176.

Watsu, J. H. T. 1891. Natural history notes from H. M. Indian Marine survey steamer ‘Inves-

tigator’. No. 24. List of deep-sea holothurians, collected during seasons 1887—91, with

descriptions of new species. Journal of the Asiatic Society of Bengal 60: 197-204.

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

ope eeethe Fieure 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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

AHMED S. THANDAR

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES. PART 18.

HOLOTHUROIDEA

VOLUME 101 PART 8 JUEY 1992 ISSN 0303-2515

OF THE SOUTH AF RICAN

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes...”

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 °+#£4Band

July 1992 Julie

Part 8 Deel

THE ALCYONACEA OF SOUTHERN AFRICA.

GORGONIAN OCTOCORALS

(COELENTERATA, ANTHOZOA)

GARY C. WILLIAMS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

iL, AC-8, 58), HCD, 445, B teas), SS, 5, 7-9),

6G, tapi), TCO), 8, S02, 7), WOES), 1G, 5, 7, tao.t.).

ID), ISS), WO, 5), Vi, INGE), 325), 38,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 131 8

Printed in South Africa by In Suid-Afrika gedruk deur

ihe Rustica Press; Pty-, Wide Die Rustica-pers, Edms., Bpk.,

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D1411

————

a =

oo es

pos

Fig. 1. Living gorgonians of southern Africa. A-B. Acabaria rubra (Esper, 1798) (Melithae-

idae). C. Leptogorgia palma (Pallas, 1766) (Gorgoniidae). D. Eunicella albicans (Kolliker,

1865) (Gorgontidae). E. Eunicella papillosa (Esper, 1797) (Gorgoniidae). F. Eunicella tri-

coronata Velimirov, 1971 (Gorgontidae). G. Narella gilchristi (Thomson, 1911) (Primnoidae).

H. Thouarella hicksoni Thomson, 1911 (Primnoidae).

THE ALCYONACEA OF SOUTHERN AFRICA.

GORGONIAN OCTOCORALS

(COELENTERATA, ANTHOZOA)

GARY C. WILLIAMS*

Department of Marine Biology, South African Museum, Cape Town

and Department of Zoology, University of Cape Town, Rondebosch

(With 71 figures)

[MS accepted 28 September 1989]

ABSTRACT

Gorgonian octocorallian coelenterates representing the families Anthothelidae, Melithae-

idae, Keroeididae, Acanthogorgiidae, Plexauridae, Gorgoniidae, Ellisellidae, Chrysogorgi-

idae, Primnoidae, and Isididae are described mainly from the shallow African sublittoral

(<200 m depth), south of 20°S. Twenty-two species are illustrated and described in detail from

material in the marine invertebrate collection of the South African Museum, Cape Town. Most

of this material has been collected recently by means of SCUBA and dredge. Eight additional

species are mentioned from literature sources but, since satisfactory material is not available for

examination, these are not illustrated or described in detail. Nine genera are represented that

do not occur in <200 m depth; these are included in the text and key but most are not iden-

tified to species. There are at least seven genera in which species identification is uncertain or

not possible at present. Several species are considered problematical. These belong mainly to

the families Melithaeidae and Ellisellidae, which are in need of thorough revision before most

material can be properly assigned even to genus. Two gorgonian genera are here considered

southern African endemics: Homophyton and Ideogorgia.

Scanning electron micrographs of sclerite form complement all detailed species descrip-

tions. New information has allowed for the revision of taxonomic status and extent of

intraspecific variation of several species. A key is provided to all identified species and genera

that are considered valid and known to occur in southern Africa.

This regional account of the gorgonian fauna includes ten families, 30 genera, and 24 iden-

tified species that are here considered valid. This includes six new combinations and 17 genera

recorded from southern Africa for the first time. At least two species are known that are pres-

ently unidentified but probably represent undescribed species; these are not included in the

present account.

CONTENTS

PAGE

JTntiTPOGICTOMD ss 55a sea Bal See ete eve ee eae rg 182

ittermalsran@omethOdS: 045. 6s. 02.6255 0 66 escent ee eee ee 183

SvSISMAMIC ACCOMME 548s aac ane o aes ob omen cs Geen ooo ao oe 183

Key (© tine TeNMNES won eoce sos escce secs seo ane uaeuoo crocs 183

RamilyeANmthothelidae e295. 6-.6-2 32622: -))- s 184

HrarmilvaNWMclithtaeidaesaaarasij-sc40¢- 22 97) 25e 6 oe emne 197

amulyaKer@cididae Yay- ee. ee os 2 203

Family Acanthogorgiidae ...........--....--------+:- 207

* Present address: Department of Invertebrate Zoology, California Academy of Sciences,

Golden Gate Park, San Francisco, California 94118, U.S.A.

181

Ann. S. Afr. Mus. 101 (8), 1992: 181-296, 71 figs.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

amily: BiexaunGd ac nth... et wares aoe teat aes 209

FamilysGorgonidace. oie cet ee er ere A ee 218

Family Mis etic iy wat neve siee sede ot ries eects tore eee 2A

BamulyiChrysocorsiidace.-- 155 piv at er oe nee Psy

Family Primnoidae 21a iene ghee: ste See ee eee 270

Ramah Vs SAT AG cts Sr Pele «ex eek rept ©. ee 278

Key to the gorgonian octocorals of southern Africa.......... 286

List of gorgonian octocorals recorded from southern Africa... 290

Summary andiconclusion.s. 2.02 4.2 se ose5 seas 5 eee 292

Acknowledgements cee nde 6 ane Sects ae ene eee 292

IRGLCRENCES ye tee oc See ta ee ae oa Nee ee eT ie ee jas)

INTRODUCTION

The present survey represents the third paper in a series that attempts to

describe the octocorals of southern Africa that have been identified and

recorded to date. Williams (1990) described the pennatulacean octocorals

(families Veretillidae, Echinoptilidae, Kophobelemnidae, Anthoptilidae, Funi-

culinidae, Protoptilidae, Scleroptilidae, Chunellidae, Umbellulidae, Virgulari-

idae, Pennatulidae and Pteroididae). Williams (1992) addressed the stoloni-

ferous octocorals and soft corals (families Clavulariidae, Tubiporidae, Coelogor-

giidae, Alcyoniidae, Nidaliidae, Nephtheidae and Xentidae). Gorgonian octo-

corals (families Anthothelidae, Melithaeidae, Keroeididae, Acanthogorgiidae,

Plexauridae, Gorgoniidae, Ellisellidae, Chrysogorgiidae, Primnoidae and Isidi-

dae) are dealt with in the present work.

The gorgonians, as defined here, consists of those octocoral cnidarians that

are attached to hard substrata by a proximal holdfast or embedded in soft sub-

strata by root-like projections of the holdfast, and have an internal medulla or

axis distinct from the cortex or outer coenenchyme. The composition and struc-

ture of this axial component varies considerably. In the Anthothelidae the

medulla is uniform and composed of loosely bound sclerites. In the family

Melithaeidae the axis is comprised of alternating internodes of fused sclerites

and soft nodes of horny material (the protein gorgonin, which contains

sclerites). The horny central chord in the family Keroeididae is chambered and

surrounded by a cortex of unfused sclerites. In the family Acanthogorgtidae the

axis is composed purely of gorgonin with a chambered central chord. The family

Plexauridae has horny axes with a chambered central chord and a cortex usually

containing calcareous material not in the form of sclerites. The family Gorgoni-

idae has purely horny axes with a chambered central chord. The families

Ellisellidae, Chrysogorgiidae, and family Primnoidae have solid, non-spicular

calcified axes of concentric layers or lamellae. Finally, the axis in the family

Isididae is composed of alternating nodes of pure horn and internodes of

calcareous material not in the form of sclerites.

The most important general studies of gorgonians include Kukenthal

(1924), which remains the most comprehensive review even though dated, and

Kikenthal (1919), which provides detailed descriptions of many species. Bayer

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 183

(1956) provided brief diagnoses of all families and genera recognized at the

time. Major sources concerning the southern African regional gorgonian fauna,

include Hickson (1900, 1904), Simpson (1910a, 1910b), Kiikenthal (1919),

Thomson (1911, 1917, 1923), Molander (1929), Broch (1939), Stiasny (1940),

Tixier-Durivault (1954), and Velimirov (1971).

The octocoral classification system used here follows that of Bayer (1981),

in which the gorgonians are allocated to 16 families within the order Alcyonacea

(which also includes the soft corals); the traditionally accepted order

Gorgonacea and the suborders Scleraxonia (gorgonians with free axial sclerites)

and Holaxonia (gorgonians without free axial sclerites) are abandoned. These

traditional divisions cannot be precisely defined due to the presence of inter-

mediate forms, and thus no distinct boundaries between Alcyonacea and Gorgon-

acea, or Scleraxonia and Holaxonia can be maintained. Terminology in the

present work conforms to Bayer et al. (1983). The notation (Lit.!) pertains to a

reference containing a detailed synonymy.

The present work is concerned primarily with the gorgonians that occur in

the coastal waters from the littoral to 200 m in depth. Most deeper water forms

(those found between 200 m and 1 200 m in depth) are identified to genus only;

this material is included in the text but the representative material is not

described in detail or illustrated, although the genera are included in the key

and species list. These include the genera Acanthogorgia Gray, 1857 (family

Acanthogorgiidae); Chrysogorgia Duchassaing & Michelotti, 1864, Radicipes

Stearns, 1883, and Xenogorgia Bayer & Muzik, 1976 (family Chrysogorgiidae);

Callogorgia Gray, 1858, Calyptrophora Gray, 1866, Primnoeides Wright &

Studer, 1887, and Primnoella Gray, 1858 (family Primnoidae); and Acanella

Gray, 1870, and Keratoisis Wright & Studer, 1869 (family Isididae).

MATERIALS AND METHODS

Recently acquired material was collected with the aid of SCUBA or dredge.

Colonies were preserved in 70 per cent ethanol. Sclerites were disassociated

from tissues with sodium hypochlorite. A Cambridge S200 scanning electron

microscope was used to make micrographs of sclerites.

SYSTEMATIC ACCOUNT

KEY TO THE FAMILIES

1A. Colony composed of an outer cortex and an inner medulla. Medulla

composed entirely of free but densely set sclerites.......... Anthothelidae

1B. Colony with axial support composed of sclerites united by horny material

and/or CaCOs, or entirely proteinous and permeated by varying amounts of

-non-spicular CaCOs, or totally calcified without sclerites or horn........ 2

2A. Axis jointed, composed of distinct and alternating nodes and internodes . 3

2B. Axis continuous and uniform, without alternating nodes and internodes . 4

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

3A. Nodes are short and proteinous, internodes are longer and composed of

non-spicular Cal Ose sass eh a iS elke Rea Bears ara gee a Isididae

3B. Axis composed of sclerites united by horny material and/or CaCO;

sida a Beate Rig antes h ol asthe toa UNS aN eae eg OS a CIR AEA Melithaeidae

4A. Axis with a central core that is hollow and cross-chambered ............ 5)

4B. Axis solidsthroughout .2..4e¢ oe se a ee 8

SA. Chambered core of axis surrounded by smooth sclerites bound together by

Sheathsiofhorn-hikeymatenaliten ae an ee Keroeididae

5B. Chambered core of axis surrounded by horn-like layers that may contain

non-spicular'CaCO3.. co eee ges es ae 4 6

6A. Calyces permanent, with slender sclerites arranged into eight chevroned

POLES Gee heals Lito St Ni Ons sar hee rae cc cae Acanthogorgiidae

6B. Calyces without eight chevroned points; or permanent calyces absent.... 7

_ 7A. Axis core narrow; axis cortex dense and hard with loculation generally

NAOKI see sae avers Real Remit Te os ON ik acy ed ae el Gorgoniidae

7B. Axis core wide; axis cortex soft or wood-like with abundant loculation

OG sn ee ee a AC Re tne RETIN Sho 6 0 0 Plexauridae

SAS Sclenites ineludemanydoubleiheadsiaa) 454-4) a5 see Ellisellidae

8B. No double heads are present 20.00.25 0. .) o e 5

VAS) Suntace olvaxis lonertudinallyseroovedie en ae eee nee Primnoidae

OB: Surface-Of axis smooth te. See tok ce ree Chrysogorgiidae

Family Anthothelidae Broch, 1916

Medulla and cortex separated by a boundary of longitudinal canals. Solenia

do not extensively penetrate medulla except sometimes distally. Axis composed

of a medulla of loosely-bound calcite sclerites. Polyps monomorphic. Antho-

codiae retractile. Polyp calyces usually present. Sclerites are needles, tuber-

culate spindles, radiates, or rods.

Eleven genera in three subfamilies from the Atlantic, southern Africa, and

Indo-Pacific. Three genera in southern Africa.

Subfamily Anthothelinae Broch, 1916

Anthocodiae retractile into permanent cylindrical calyces. Cortical sclerites

are mostly elongate and tuberculate spindles. Medullar sclerites of slender

spindles.

Genus Anthothela Verrill, 1879

Anthothela Verrill, 1879: 199. Kiikenthal, 1919: 43; 1924: 14 (Lit.!).

Diagnosis

Colonies composed of few slender, often tangled or sinuous branches, with

or without a main basal stem. Polyps widely spaced (5S—10 mm), scattered on all

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 185

sides of round branches, not in definite rows. Conical to cylindrical permanent

calyces with anthocodiae usually preserved partly exserted. Stems round; a thin

cortex surrounds a thick medulla.

Two or three species of the northern Atlantic, South Africa, and possibly

the Pacific coast of central America.

Type species. Briareum grandiflorum Sars, 1856; north Atlantic.

Anthothela parviflora Thomson, 1917

Figs 2-3, 4A—D

Anthothela parviflora Thomson, 1917: 3, pl. 2 (fig. 5), pl. 5 (fig. 4).

Material

SAM-—H1153, herein designated lectotype, off Cape Recife, eastern Cape

Province (34°27’S 25°42'45"E), 468 m, 14 November 1898, several fragmented

colonies; dredge, S.S. Pieter Faure survey, PF 655. SAM-—H3653, off Sandy

Point, Transkei (32°41,2’S 28°43,9'E), 480-490 m, 14 July 1984, 4 whole

colonies, dredge, G. C. Williams, R.V. Meiring Naude.

Description

Colonies examined are 50-90 mm in length. Most colonies have two or

three branches arising from a short (<20 mm) main stem. The main branches

have few or no further lateral branches, and are sinuous or tortuous in appear-

ance. The main stem and lowermost portions of the branches are devoid of

polyps. Polyps restricted to distal half of branches, widely spaced (5-8 mm

apart), arranged on all sides of branches. Calyces elongate conical to cylindrical,

with more-or-less truncate apices (<5 mm long). Eight longitudinal grooves

conspicuous on external surface of calyces. Anthocodia often preserved partially

exserted; with numerous needle-like spindles or a few stout rods (0,04—-0,35 mm

long), arrangement not determined. Cortex with oval to elongate spindles

(0,11-0,35 mm in length), sparsely to densely tuberculate, some spindles club-

shaped. Medulla with slender spindles (0,17—0,43 mm long), sparsely tuberculate

to somewhat thorny or weakly branched.

Distribution

Known only from the south coast of South Africa—Cape Recife (Algoa Bay)

and central Transkei; 183-490 m in depth; apparently endemic to southern

Africa (Williams in press a). Type locality is the Algoa Bay region.

Remarks

Thomson (1917: 4-5) differentiated in detail between A. grandiflora and

A. parviflora. According to him, A. grandiflora is luxuriantly bushy with anasto-

mosing branches but without a principal stem. In A. parviflora, the branching ts

very sparse, anastomosis does not take place, and the colony arises from a single

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Anthothela parviflora Thomson, 1917. A. Entire colony on a dead coral fragment, total

length of colony 50 mm. B. Detail of a branch in transverse section, with two polyps; length of

figure 10 mm. Abbreviations: a—anthocodia, c—cortex, ca—calyx, m—medulla.

basal stem. The characters that distinguish A. grandiflora, as well as details of its

distribution, can be found in the following works: Sars (1856), Kukenthal (1919:

43, fig. 17), Verrill (1922), Thomson (1927: 16), and Grasshoff (1982a, map 1;

1982b: 942, fig. 4).

Subfamily Spongiodermatinae Aurivillius, 1931

Calyces present or absent. Cortical sclerites with capstans or radiates.

Medullar sclerites are rods that are often branched or spiny.

Genus Homophyton Gray, 1866

Solanderia Mobius, 1861: 3 (preoccupied by Solanderia Duchassaing & Michelin, 1846: 218).

Homophyton Gray, 1866: 27.

Spongioderma Kolliker, 1870: 14. Kuikenthal, 1919: 89; 1924: 34 (Lit!).

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 187

Diagnosis

Colonies elongate, unbranched or with few dichotomous branches arising

from a single basal stem. Calyces prominent or absent. Single ring of longitudi-

nal canals between cortex and medulla. Cortex in one layer, composed primarily

of globose radiates. Medulla composed of stout rods that are somewhat

branched or spiny.

A monotypic genus restricted to southern Africa.

Fig. 3. Anthothela parviflora Thomson, 1917. A. Half transverse section through base of

colony showing internal structure; total diameter 3 mm. Abbreviations: c—cortex, |—longi-

tudinal canal, m—medulla. B. Anthocodial sclerites. C. Sclerites from the calyx and cortex.

D. Sclerites from the medulla. Scale bar =0,4 mm.

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Scanning electron micrographs. A-—D. Anthothela parviflora Thomson, 1917, sclerites

of calyx and cortex. A.0,130mm. B.0,210mm. C. Two sclerites 0,080—0,145 mm.

D. 0,125 mm. E-H. Homophyton verrucosum (Mobius, 1861), coenenchymal sclerites.

E. 0,075 mm. F. 0,055 mm. G. 0,100 mm. H. Three sclerites, 0,040—0,055 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 189

Type species. Solanderia verrucosa Mobius, 1861; South Africa.

Homophyton verrucosum (Mobius, 1861)

Figs 4E—-H, 5-6

Solanderia verrucosa Mobius, 1861: 3. Kolliker, 1865: 141.

Homophyton Gattyiae Gray, 1866: 27; 1869: 96.

Spongioderma verrucosa Kolliker, 1871: 11. Kiikenthal, 1919: 91, figs 39-43, pl. 3 (fig. 8):

1924: 34, figs 26-28. Molander, 1929:5. Stiasny, 1940: 13, pl. 1 (figs 1-7). Tixier-

Durivault, 1954: 529. Day et al., 1970: 16.

Spongioderma verrucosum: Hickson, 1900: 78. Thomson, 1911: 874.

Spongioderma chuni Kikenthal, 1908: 18; 1919: 94, pl. 3 (fig. 9); 1924: 36. Molander, 1929: 6.

Stiasny, 1940: 16. Tixier-Durivault, 1954: 529; 1960: 362.

Homophyton: Bayer, 1981: 910.

Homophyton verrucosum Williams, 1989: 142; in press c.

Material

SAM-—H3645, off East London (33°02,3’S 27°55,9'E), 30-35 m, 16 July

1984, 2 colonies, dredge, G. C. Williams, R.V. Meiring Naude. SAM—H3341,

Algoa Bay (33°50’S 25°40'E), 12 m, 16 May 1984, 2 colonies, SCUBA, G. C.

Williams.

Description

Colonies examined are up to 420 mm in length and 5-15 mm in width.

Colonies rarely unbranched and whip-like; commonly dichotomously branched

from a single basal stem that is 60-120 mm in length. Base of colony up to

15 mm in diameter. Amount of branching variable, sparse to profuse. Branches

elongate, finger-like tapering distally. Polyps cover entire colony except the

basal holdfast region. Calyces prominent and conical-truncate to cylindrical (up

to 3 mm long), or low and rounded, or absent (being retracted into branches).

Cortex composed of densely set globular to ovoid radiates, small triradiates, and

tuberculate spheroids, 0,07—0,22 mm long. Medulla composed of densely-set

rods 0,13—0,18 mm long, which are somewhat branched or sparsely spiny.

Colour yellow, pinkish-white, red-orange, or deep wine-red; calyces whitish,

yellow, or wine-red. Colonies commonly bicoloured, i.e. red-orange with yellow

calyces; or monochromatic, i.e. entirely whitish, yellow, or wine-red.

Distribution

Apparently restricted to the south and east coasts of southern Africa,

known definitely from Cape Agulhas to Sodwana Bay, Natal; intertidal to depth

of 168 m (Williams in press a). The species was also listed without descriptions

or figures from False Bay, western Cape Province by Day et al. (1970: 16), and

Inhaca Island, southern Mozambique by Tixier-Durivault (1960: 362). This is a

variable and locally common species, often found on or adjacent to rocky reefs

in depressions or gullies filled with sand. Specific type locality not known.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Homophyton verrucosum (Mobius, 1861). Three entire colonies. A.A colony from

100 m depth off East London, length 200 mm. B. A colony from 30 m depth off East London,

length 230 mm. C. A colony from 12 m depth in Algoa Bay, length 460 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 19]

Remarks

Kikenthal (1908) described Spongioderma chuni from uniformly wine-red

colonies from Algoa Bay. The species was retained by Molander (1929), Stiasny

(1940), and Tixier-Durivault (1960). Comparative examination of many colonies

of this species from throughout its range has shown the only consistent differ-

ence between Kukenthal’s species and H. verrucosum to be colour. No other

morphological features are distinctive, including sclerite characteristics. I there-

fore consider Kukenthal’s species to be conspecific with H. verrucosum.

Solanderia of Mobius, 1861, is preoccupied by Solanderia Duchassaing &

Michelin, 1846. Since the two are synonymous, the name Homophyton Gray.

1866, has priority over Spongioderma KoOlliker, 1871.

Homophyton verrucosum is a highly variable species in regards to colour,

growth form, calyx development, and sclerite size and shape. The monochroma-

tic wine-red form seems to be restricted to the Algoa Bay region. In the East

London area, an orange form with yellow calyces is commonly encountered.

Whitish, pinkish, or yellowish forms with prominent conical to cylindrical, non-

retractile calyces (up to 3 mm) are infrequently observed from East London to

Natal. The Algoa Bay and East London forms have retractile calyces that are

most often preserved as low rounded or conical-truncate protuberances

(<2 mm) on the colony surface, although a few calyces may be totally retracted

and are preserved flush with the colony surface. The cortical sclerites vary in

shape and size from colony to colony. Some colonies have narrower elliptic

radiates up to 0,22 mm in length, whereas others have more spheroid radiates

that do not exceed 0,18 mm in diameter.

Most specimens of Homophyton verrucosum possess varying amounts of

encrusting sponge (an unidentified species of Demospongiae). The sponge is

thin, greyish-white, and may completely cover the external surface of the octo-

coral, except for the conical calyces that remain exposed. The sponge spicules

are numerous styles (long needle-like forms) with a few anisochelas (small hook-

like forms).

In addition to the conspicuous ring of longitudinal canals that separate the

cortex and medulla, several other canals may be dispersed throughout the

cortex. In some specimens, these additional canals may appear to form a second

concentric ring between the cortex and medulla, as described by Kukenthal

(1924: 34). These rings are exclusive of the ring of small canals that is sometimes

apparent directly beneath the external surface (Fig. 5A).

Homophyton verrucosum is distinguished by the presence of globular

radiates and tuberculate spheroids (0,07—0,22 mm in diameter) in the calyces

and coenenchyme.

The superficial resemblances between Homophyton verrucosum and Ideo-

gorgia capensis (family Keroeididae) deserve mention. The two species are

presently placed in different families (and would even be placed in separate

orders if the formerly accepted classification system (Bayer 1956) was used). The

sclerites of both the cortex and medulla are remarkably similar and the growth

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Homophyton verrucosum (Mobius, 1861). A. Half transverse section through the basal

portion of a colony, showing internal structure; total diameter 12 mm. Abbreviations: c—

cortex, |—longitudinal canal, m—medulla. B. Sclerites from cortex. C. Sclerites from

medulla. Scale bar =0,2 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 193

forms and colony colour are also somewhat similar. However, the axis of /deo-

gorgia possesses a chambered central chord surrounded by a layer of smooth

sclerites, which is bound together by horn-like sheaths. Homophyton, on the

other hand, lacks a central chord in the axis but has a thick medulla of uniform

consistency. Examination of many specimens of Homophyton at a variety of

levels within the colonies has revealed no trace of a central chord. Because of

this, the two species are retained provisionally in separate families despite their

striking superficial similarities.

Genus Diodogorzia Kiikenthal, 1919

Diodogorgia Kukenthal, 1919: 96; 1924: 36.

Diagnosis

Colonies branched and tree-like with capitate branch tips, or unbranched

and capitate. Cortex organized into a thin outer layer and a thick, non-vesicular

inner layer separated by a plexus of narrow canals. Medulla separated from

cortex by a layer of wide canals. Outer cortex with small capstans and tri-

radiates, inner cortex with stout tuberculated spindles, medulla with sparsely

thorny spindles and rods that are sometimes branched. Colony colour reddish,

incorporated in sclerites of the cortex.

Three species from the Gulf of Mexico, West Indies, and South Africa.

Type species. Diodogorgia ceratosa Kikenthal, 1919: 97; Gulf of Mexico.

Diodogorgia capensis (Thomson, 1911) comb. nov.

Figs 7-8, 9A—D

Suberia capensis Thomson, 1911: 871, pl. 43 (fig. 4), pl. 45 (fig. 4a—c). Kukenthal, 1919: 89;

1924: 34. Stiasny, 1940: 12.

Material

SAM-—H584, off Hood Point near East London (33°05'S 27°50'E), 90 m,

15 July 1901, 2 partial colonies, dredge, S.S. Pieter Faure survey, PF 13056.

SAM-—H3590, off Port Grosvenor, Transkei (31°25'6"S 29°56'0"E), 80-84 m,

August 1981, 2 colonies, dredge, R. N. Kilburn, R.V. Meiring Naude.

Description

Colonies examined are 25-30 mm in length. Colonies are unbranched,

slightly clavate or markedly capitate. The polyp-bearing capitulum may arise

from a narrower basal stalk, or polyps may be evenly distributed throughout

most of digitiform colonies. Polyps generally with low rounded or truncate

calyces, 2mm in diameter. Calyces may retract completely into polyparium.

Sclerites of outer cortex are triradiates and capstan radiates, 0,05—0,15 mm in

length. Inner cortex sclerites are robust tuberculate spindles, 0,30—0,50 mm in

length. The medulla possesses spindles and rods, 0,15—0,50 mm in length, which

are weakly branched or sparsely thorny. The outer cortex is a thin dense layer,

0,25 mm in thickness. The inner cortex is less dense, but without large vesicles.

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Diodogorgia capensis (Thomson, 1911). Two entire colonies, 25 mm and 20 mm in

length.

A plexus of narrow canals separates the inner and outer layers of the cortex.

The longitudinal canals that separate the cortex and medulla are conspicuous,

0,05 mm in diameter. Colony colour uniform deep rust-orange to wine-red.

Distribution

Known only from the southern coast of South Africa from East London to

northern Transkei; 80—90 m in depth; apparently endemic to southern Africa.

Type locality is Cape Morgan, border of Cape Province and Transkei.

Remarks

Bayer (1955: 208) showed that the genus Suberia Studer, 1879, is a junior

synonym of Semperina Kolliker, 1870. Diodogorgia has radiates in the outer

cortex, whereas Semperina does not contain radiates in the cortex. Diodogorgia

is therefore the correct generic designation.

The southern African species can be placed in the genus Diodogorgia on

the basis of capitate terminal tips of the colony, organization of the cortex into a

thin outer layer, a thick inner layer without large vesicles and separated by a

plexus of thin canals, radiates in the outer cortex, stout tuberculated spindles in

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 195

Fig. 8. Diodogorgia capensis (Thomson, 1911). A. Half transverse section through basal

region showing internal structure, total diameter 4mm. Abbreviations: ic—inner cortex,

1—longitudinal canal, m—medulla, oc—outer cortex. B. Sclerites from the outer cortex.

C. Sclerites from the inner cortex. D. Sclerites from the medulla. Scale bar =0,2 mm.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Scanning electron micrographs. A-D. Diodogorgia capensis (Thomson, 1911).

A. Sclerite from the outer cortex 0,088 mm. B. Sclerite from the inner cortex 0,350 mm.

C. Two sclerites from the inner cortex 0,30-0,40 mm. D. Sclerite from the medulla, 0,30 mm.

E-G. Ideogorgia capensis (Simpson, 1910), coenenchymal sclerites. EE. Five sclerites,

0,035-—0,040 mm. F. 0,040 mm. G. 0,060 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 197

the inner cortex, sparsely thorny spindles and rods in the medulla, and reddish

colony due to permanent coloration of sclerites of the outer cortex.

Diodogorgia capensis is distinguished from the other two species of the

genus, D. ceratosa Kukenthal, 1919, and D. cervicornis Kikenthal, 1919, by an

unbranched growth form, the latter two species being bushy or arborescent.

The present work represents a new record for the genus Diodogorgia in

southern Africa.

Family Melithaeidae Gray, 1870

Axis composed of internodes of fused sclerites and nodes of horny material

with sclerites. Branches arise from the nodes. Polyps monomorphic. Coenenchy-

mal sclerites diverse, conspicuously sculptured, spindles, clubs, foliates, and

spheroids. Axial sclerites are smooth rods often with median protuberances

forming a transverse ring.

Five genera of Indo-Pacific distribution. Only one of these (Acabaria Gray,

1859) is considered valid, whereas the remaining four, Melithaea Milne Edwards

& Haime, 1857 (= Melitella, Melitodes, Birotulata), Mopsella Gray, 1857, Clath-

raria Gray, 1859, and Wrightella Gray, 1870, are considered nominal (and

probably do not even merit subgeneric status since they all have more-or-less the

same sclerite forms) by Bayer (1981: 917) and Van Ofwegen (1987: 6). Classi-

fication of the family is in great need of revision.

Genus Acabaria Gray, 1859

Acabaria Gray, 1859: 484; 1868: 445. Ridley, 1884: 361, 363. Kikenthal, 1919: 174; 1924: 73

(Lit!). Bayer, 1956: F200. Van Ofwegen, 1987: 5.

Anicella Gray, 1868: 445.

Psilacabaria Ridley, 1884: 361, 363.

Melitodes (part.) Wright & Studer, 1889: 171.

Diagnosis

Sclerites of the coenenchyme are large thorny spindles, rods, and some

capstans, with thorn clubs or leaf clubs also present.

An Indo-Pacific genus of at least 20 species.

Type species. Acabaria divaricata Gray, 1859; Australia.

Acabaria rubra (Esper, 1798) comb. nov.

Figs 1A-B, 10-13

Isis dichotoma cortice rubro Esper, 1798: 6, pl. 1 (figs 4-5).

Melitodes dichotoma (non Pallas, 1766) Hickson, 1900: 80, pls 1-2, 6 (fig. B). Thomson, 1911:

SH.

Melitodes africana Kiikenthal, 1908: 194; 1919: 146, text-figs 69-74, pl. 31 (fig. 10), pl. 36

(fig. 30); 1924: 60, figs 45, 46. Molander, 1929: 6. Broch, 1939: 15, figs 10-15. Day et al.,

1970216:

Melitodes esperi (non Wright & Studer, 1889) Thomson, 1911: 874, fig. 167.

Melitodes nodosa (non Wright & Studer, 1889) Thomson, 1911: 876.

Mopsella faurii (non Thomson, 1917) Broch, 1939: 20, figs 16-21.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mopsella dichotoma (non Pallas, 1766) Day et al., 1970: 16.

Wrightella coccinea (non Ellis & Solander, 1786) Day et al., 1970: 16. Day, 1974: 34.

Wrightella: Branch & Branch, 1981: 151, fig. 191.

Material

SAM-—H3327, Algoa Bay, eastern Cape Province (33°50’S 25°40'E), 15 m,

17 May 1984, 1 colony, SCUBA, G. C. Williams. SAM—H3718, wreck of the

Transvaal, False Bay, south-western Cape Province (34°15'S 18°35’E), 30 m,

1983, 3 colonies, SCUBA, P. Zoutendyk. SAM—H3874, Hout Bay, Cape of

Good Hope Peninsula (34°04'S 18°20’E), 30 m, 27 April 1985, several colonies;

SCUBA, G. C. Williams.

terror

joer

{ocneepo

a wr OD

ASST

2 a)

OV0S OOS RET URE

od ce a0)

pee

oi taee

Oise

= Sa,

pres)

Fig. 10. Acabaria rubra (Esper, 1798). An entire colony, 200 mm in length,

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 199

Description

The colonies examined vary from 25 mm to 200 mm in length. Growth form

bushy, planar or multiplanar. Branching dichotomous, intricate. Anastomosis

frequent. Internodes mostly 8-20 mm in length. Colonies with relatively brittle

branches. Polyps arranged on all sides of branches in most colonies. Some

colonies may have branches with opposite bare tracts perpendicular to the plane

of the colony. In most colonies, the polyps are crowded and cover almost the

entire surface of the colony, virtually extending to the base near the holdfast.

Retracted polyps form conspicuous calyces that are either conical or hemi-

spherical, usually 1,0—1,5 mm in diameter. Anthocodia with crown and points of

mostly strongly tuberculated spindles and clubbed spindles, 0,08—0,2 mm long.

The calyx is densely set with spindles clubs and leaf clubs, 0,12—0,26 mm in

length. These have prominent tubercles. The coenenchyme covering the

branches is densely set with spindles, clubs, leaf clubs, and unilaterally foliate

spheroids, 0,12-—0,25 mm long. These too are conspicuously tuberculated.

Colony colour variable: red, orange, yellow, or white; orange with yellow

calyces or anthocodiae also common.

Distribution

The species is known from Melkbos on the Cape west coast to East

London, eastern Cape Province; intertidal to 159m in depth (Williams in

press a). Type locality Cape of Good Hope.

Remarks

This species is herein allocated to the genus Acabaria because it has

sclerites in the coenenchyme that are mainly long spindles, with thorn clubs and

leaf clubs also present. Dr M. Grasshoff (pers. comm.) has recently examined

the type of this species and has observed the sclerites to be as described here.

Acabaria rubra is a highly variable species regarding growth form as well as

colour. Low-growing (<150 mm high) colonies that are multiplanar and densely

bushy occur in protected localities on vertical walls. Tall colonies (up to 200 mm

or more in length) that are planar with relatively less branching occur in open

areas such as on horizontal reefs and the tops of boulders and shipwrecks. These

colonies are aligned perpendicular to prevailing currents. Anastomosis occurs

frequently in most colonies, both in planar and multiplanar growth forms.

Polyps may occur on only two sides of the branches (as in planar colonies in

which the polyps are aligned parallel to the plane of branching), or may occur

crowded together on all sides of the branches (as in most multiplanar and many

planar colonies). Some colonies may grow in tangled mats together with large

assemblages of sponges, bryozoans, or tunicates. Colonies may be entirely red,

orange, yellow or white, or may be various shades of red-orange with yellow

calyces or anthocodiae. Anthocodiae may also be white. Thus, depending on the

amount of expansion or retraction of the anthocodiae, colonies may be either

bicoloured or monochromatic.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

ASS %

iN Ko

%, cf Pi

oss, ON Ly

a <2 3g

Be Sereaer

Fig. 11. Acabaria rubra (Esper, 1798). A. An entire colony, 125 mm long, and five sclerites

from the axis. B. A single polyp showing placement of sclerites in the partially retracted antho-

codia and basal calyx; length of polyp 1,2 mm. C. Anthocodial sclerites. D. Calyx sclerites.

E. Coenenchymal sclerites. Scale bar =0,1 mm for all sclerites figured.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 201

Fig. 12. Scanning electron micrographs. Acabaria rubra (Esper, 1798); sclerites of the calyx

and coenenchyme. A. 0,080 mm. B. 0,100 mm. C. 0,170 mm. D. 0,240 mm. E. 0,125 mm.

F. 0,120 mm.

DW ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Scanning electron micrographs. Acabaria rubra (Esper, 1798); sclerites of the calyx

and coenenchyme. A. 0,21 mm. B.0,225 mm. C. 0,215 mm. D.0,20 mm. E. 0,125 mm.

F. 0,120 mm. G. Two sclerites, 0,185 mm and 0,210 mm. H. Two sclerites, 0,080 mm and

0,240 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 203

The relative proportions of the various types of sclerites (spindles, clubs,

leaf clubs, and unilaterally foliate spheroids) is also quite variable.

Molander (1929: 7) recognized the remarkable variation within this species.

The great variability of many species in the family has also led to much confu-

sion in the literature as several names have often been applied to the same

species (as exemplified by Acabaria rubra).

Other species of Melithaeidae

Acabaria rubra is by far the commonest melithaeid in southern African

coastal waters, but at least twelve other melithaeids have been recorded or

described from southern Africa. Because of the present state of the literature, it

is not possible to identify most melithaeids with much degree of confidence

(PeAldersiade and Dr F. M. Bayer, pers. comm.). I believe that a generic

revision, comparison of holotypes, and a study of intraspecific variation, are

necessary before the regional species (other than A. rubra) can be identified

with assurance.

Those that appear in the literature (excluding the names here considered to

be synonyms of Acabaria rubra) are:

Acabaria capensis (Studer, 1879) (recorded by Tixier-Durivault (1954: 530) as a

melithaeid; = Isidella capensis Studer, 1879 [incertae sedis}]).

Acabaria rosea Tixier-Durivault, 1954: 531.

Acabaria sp. Thomson, 1917: 12.

Acabaria valdiviae Kikenthal, 1908: 198.

Melitodes faurti Thomson, 1917: 6.

Melitodes grandis Thomson, 1917: 8.

Mopsella singularis Thomson, 1917: 10.

Wrightella coccinea Gray, 1870; recorded by Hickson, 1904: 219 and Tixier-

Durivault, 1954: 530.

Wrightella fragilis Thomson, 1917: 15.

Wrightella furcata Thomson, 1917: 17.

Wrightella sp. Thomson, 1917: 19.

Wrightella trilineata Thomson, 1917: 13.

An unidentified, cherry-red species tentatively referable to the genus

Wrightella occurs in Algoa Bay (10 m depth). A delicate deep-red species of

Acabaria has been found on the Kosi Bay reefs, 18-24 m in depth (Williams

1989; in press c).

Family Keroeididae Kinoshita, 1910

Axis composed of a central core of horn that is chambered with a surround-

ing cortex of smooth rod-like unfused sclerites in sheaths of horn. Axis

surrounded by an axial sheath. Coenenchymal sclerites are spindles, plates, or

triradiates. Polyps monomorphic.

Three Indo-Pacific genera; one genus in southern Africa.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus /deogorgia Bayer, 1981

non Juncella (misspelling of Junceella Valenciennes, 1855) Hickson, 1900: 85.

Dendrogorgia Simpson, 1910: 62; preoccupied by Dendrogorgia Duchassaing, 1870 (see Bayer

1981: 902).

Ideogorgia Bayer, 1981: 919.

Diagnosis

Polyps retractile, not forming calyces. Coenenchymal sclerites are primarily

triradiates.

A monotypic genus endemic to South Africa.

Type species. Dendrogorgia capensis Simpson, 1910; South Africa.

Ideogorgia capensis (Simpson, 1910)

Figs 9E-G, 14-16

Juncella elongata (non Pallas, 1766) Hickson, 1900: 85, pl. 5 (fig. D).

Juncella elongata (non Pallas, 1766) var. capensis Hickson, 1904: 233.

Dendrogorgia capensis Simpson, 1910: 62, figs 1-4.

Spongioderma capensis Kikenthal, 1919: 95; 1924: 36.

Ideogorgia capensis Bayer, 1981: 919, fig. 31.

Material

SAM-—H3870, off Cape St Francis, eastern Cape Province (34°16'S 24°50'E),

30 m, 20 October 1985, 1 colony, SCUBA, W. R. Liltved. SAM-—3875, off Cape

St Francis (34°16’S 24°50’E), 37 m, 14 January 1986, 1 colony, SCUBA, W. R.

Liltved. SAM-3592, Algoa Bay (33°50'S 25°45’E), depth unrecorded, 13 dry

colonies, bottom trawl, Ocean Pearl. SAM-—H1129, off Algoa Bay (33°58’S

25°51'30"E), 46 m, 1 November 1898, 1 colony fragment, dredge, S.S. Pieter

Faure survey, PF 504.

Description

The colonies examined range in length from 180 mm to 750 mm. Colonies

elongate, upright, more-or-less planar. Branching dichotomous, sparse. Anas-

tomosis absent. Branching begins from the top of the single basal stem, which

does not exceed 70 mm in length. Base of colony up to 30 mm in diameter.

Branches 40-120 mm long. Terminal branches slender and whip-like, tapering

gradually toward the terminus. Retracted polyps do not form calyces and are

capable of totally withdrawing into the coenenchyme. Surface of branches

smooth; or with minute, low, rounded protuberances corresponding to polyps;

or weakly longitudinally grooved (particularly near the base). Sclerites from the

coenenchyme are primarily triradiates with some double heads, 0,03—0,08 mm

in length. Sclerites from the axis cortex are irregularly-shaped flattened rod-like

forms 0,07—0,17 mm in length. Anthocodial sclerites absent. Colony colour

uniform brick-red in alcohol, pink when dry. Anthocodiae white.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 205

Distribution

The species is known only from Cape St Francis and Algoa Bay, eastern

Cape Province; 30—73 m in depth; presumably endemic to southern Africa. It is

frequently encountered in large quantities by trawlers. Type locality is Algoa

Bay.

Remarks

Ideogorgia capensis is distinguished by the presence of sclerites that are

mainly small compact triradiates (0,03—0,08 mm in diameter) and a horn-like

central chord in the axis.

Fig. 14. Ideogorgia capensis (Simpson, 1910). An entire colony, 200 mm in length, low growth

form.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. Ideogorgia capensis (Simpson, 1910). An entire colony, 280 mm in length, tall growth

form.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

SS)

Fig. 16. Ideogorgia capensis (Simpson, 1910). A. Half transverse section through a distal

branch showing internal structure; total diameter 5mm. Abbreviations: a—axis, ac—axis

cortex, as—axial sheath, c—coenchyme, cc—central chord, I1—longitudinal canals, p—

polyp. B. Sclerites from the coenenchyme. C. Sclerites from the axis cortex.

Scale bar = 0,1 mm.

Family Acanthogorgiidae Gray, 1859

Axis composed solely of horn with a chambered central core. Coenenchyme

thin. Polyps monomorphic, non-retractile, cylindrical or clavate, heavily armed

with spindles.

Seven genera of cosmopolitan distribution. Two genera in southern Africa.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Acanthogorgia Gray, 1857

Acanthogorgia Gray, 1857: 128. Bayer, 1956: 203. Kiikenthal, 1919: 298; 1924: 239. Stiasny,

OAS SATE

Blepharogorgia (part.) Duchassaing & Michelotti, 1864: 15.

Boarella Gray, 1870: 406.

Paracanthogorgia Stiasny, 1947: 53.

Diagnosis

Calyces tall cylindrical. Calyx walls covered with bent spindles arranged in

eight en chevron longitudinal rows. Distal terminus of calyx armed with eight

groups of projecting crown spines.

Approximately 30 species from all seas in deeper water (50 m to over

1 200 m).

Type species. Acanthogorgia hirsuta (? = A. aspera Pourtales, 1867); see

Bayer, 1956: 203; northern Atlantic.

Remarks

At least two unidentified species of Acanthogorgia are known from south-

ern Africa. One species is present from off the Cape Peninsula and southern

Transkei, 303-585 m in depth. Hickson (1904: 225) identified this species as

A. armata Verrill, 1878, originally described from the Atlantic coast of North

America. Thomson (1911: 880; 1916: 21; 1923: 75) followed Hickson’s idensifi-

cation for additional material. Thomson (1917: 21) also described a fragmentary

and unidentified specimen of Acanthogorgia from Cape Vidal, northern Natal,

146-183 m in depth. Stiasny (1940: 21) questioned Hickson’s identification. A

species different to Hickson’s and Thomson’s material has recently been found

off Durban, Natal, at 133 min depth. I consider all these specimens to be unidenti-

fiable at present. A comparison with the original descriptions and type material

of the many recognized species is necessary for valid species identifications.

Genus Anthogorgia Verrill, 1868

Anthogorgia Verrill, 1868: 413. Thomson & Simpson, 1909: 194. Nutting, 1910: 25.

Diagnosis

Calyces cylindrical. Sclerites of apex of calyx en chevron in eight points.

Sclerites of calyx body wall only slightly or indistinctly en chevron. Interior

coenenchyme in contact with the axis possesses small radiates.

Seven species of the Indo-Pacific.

Type species. Muricea divaricata Verrill, 1865; Indo-West Pacific.

Remarks

Thomson (1917: 22) recorded Muricella ramosa Thomson & Henderson,

1905 (known originally from the Persian Gulf to the Malay Archipelago at

40-113 m in depth), from Durnford Point, northern Natal, at 82 m in depth.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 209

Thomson’s specimen (SAM-—H975) and an additional specimen, also from Durn-

ford Point but from 165 m (SAM-H1003), are here considered to represent an

unidentified species of Anthogorgia. These specimens have cylindrical calyces

with sclerites from the calyx wall forming points that are only indistinctly en

chevron.

The present work represents a new record for the genus Anthogorgia in

southern Africa.

Family Plexauridae Gray, 1859

Axis composed of a chambered central core and a surrounding area of horn

that usually contains varying amounts of calcium carbonate not in the form of

sclerites. Polyps monomorphic, capable of retracting completely into coenen-

chyme or anthocodiae retracting into permanent calyces.

A diverse family of 30 genera; widespread in the Atlantic, Indian, and

Pacific Oceans. At least five genera in southern Africa.

Genus Echinomuricea Verrill, 1869a

Echinomuricea Verrill, 1869a: 285. Wright & Studer, 1889: 112. Thomson & Simpson,

1909: 198. Nutting, 1910: 56.

Diagnosis

Permanent calyces present, often conspicuous and more or less prickly.

Many sclerites of the calicular region are thorn stars with a single elongate taper-

ing spine projecting from a basal plate that is branched or lobed. Colony colour

usually brown or red.

Approximately 16 species of the Indo-West Pacific and Gulf of Mexico.

Type species. Nephthya coccinea Stimpson, 1855, Hong Kong.

Remarks

Two unidentified species have recently been collected from the Durban

region of Natal—a richly branched red form known from 10 m depth and a

sparsely branched light brownish-grey form from 53—88 m depth. Identification

to species has not been made at present.

The present work represents a new record for the genus Echinomuricea in

southern Africa.

Genus Euplexaura Verrill, 1869c

Plexaura Verrill, 1865: 186 (part.).

Euplexaura Verrill, 1869c: 75. Wright & Studer, 1889: 143. Kiikenthal, 1919: 215; 1924: 90.

Diagnosis

Calyces form low inconspicuous protuberances. Sclerites are spindles, many

approaching tuberculate ovals and spheroids.

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

A genus of perhaps 20 species of the Indian and Pacific oceans.

Type species. Euplexaura capensis Verrill, 1869c; South Africa.

Remarks

Two unidentified species have recently been collected from off Durban,

Natal, from 14 m and 68 m in depth. Both species are brownish-white in colour.

They have not been identified to species.

Verrill (1865) misidentified South African material as Plexauria friabilis

Lamouroux, 1816. Verrill (1869c) subsequently created the name Euplexaura ca-

pensis for the same taxon. Verrill’s material is not presently available for

comparison with the recently collected specimens.

Thomson (1911: 883, pl. 44 (figs 2a—c)) described Euplexaura media from

Cape Recife, south coast of South Africa, 31 m depth. Thomson (1917: 24)

identified a specimen from Stalwart Point, eastern Cape Province (97 m depth)

as Euplexaura parciclados Wright & Studer, 1889, originally described from

Japan. Examination of both of Thomson’s specimens has shown them to belong

to the genus Lophogorgia, owing to the predominance of symmetrical six- and

eight-radiates. Reliable identification to species is not possible since the material

is dry, fragmentary, and badly damaged.

Genus Menella Gray, 1870

Menella Gray, 1870: 407. Wright & Studer, 1889: 53. Nutting, 1910: 85.

Diagnosis

Sclerites are rooted leaves with broad rounded blades forming the distal

projection and a branched tuberculated handle forming the proximal portion.

Irregular spindles also present.

Three or four species of the Indo-Pacific.

Type species. Menella indica Gray, 1870; Japan.

Remarks

Two colonies representing this genus have recently been collected on the

sublittoral coral reefs at Sodwana Bay and Kosi Bay, northern Natal, at

18-24 m in depth (Williams 1989: 142; in press c). The material has not been

identified to species. The colonies are deep wine-red in colour.

The present work represents a new record for the genus Menella in south-

ern Africa.

Genus Astromuricea Germanos, 1896

Astromuricea Germanos, 1896: 175.

Diagnosis

Colonies planar, branching sparse to profuse. Axis thick, coenenchyme

thin. Anthocodiae with conspicuous crown and points. Permanent calyces with

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 211

thorn scales, some of which have an elongated basal root and a tuberculated or

serrated head that arises obliquely. Coenenchyme contains thorn scales or thorn

spindles.

A genus of at least two species from the Indo-West Pacifc.

Type species. Astromuricea polyklados Germanos, 1896; Ternate (Indonesia).

Astromuricea fusca (Thomson, 1911) comb. nov.

Figs 17-18, 19A—C

Muriceides fusca Thomson, 1911: 878, pl. 44 (figs 4a—c). Kikkenthal, 1924: 164. Stiasny, 1940:

20, pl. 2 (figs 11-12).

Villogorgia mauritiensis (non Ridley, 1882) Hickson, 1900: 87.

Material

SAM-—H3603, off Gonubie, eastern Cape Province (33°04,7’S 28°07,2’E),

90m, 17 July 1984, 4 dry colonies, dredge, G. C. Williams, R.V. Meiring

Naude. SAM-—H3821, off Whale Rock, Transkei (32°02,0’S 29°19,1’E),

200-210 m, 3 July 1985, 3 dry colonies, dredge, G. C. Williams, R.V. Meiring

Naude. SAM-—H3905, off Whale Rock, Transkei (32°02,0’S 29°19,1’E),

200-210 m, 3 July 1985, 4 colonies, dredge, G. C. Williams, R.V. Meiring

Naude. SAM-—H3606, off Park Rynie, Natal (30°20,05’S 30°51,4’E), 96 m,

10 July 1984, 5 colonies, dredge, G. C. Williams, R.V. Meiring Naude.

Description

Colonies examined range from 80 mm to 300 mm in height and 70 mm to

500 mm in breadth. Colonial growth form is planar, branching is bushy and

usually lateral; some anastomosis does occur between adjacent branches.

Colonies arise from a broad and bulky holdfast, often with a wood-like texture.

Polyps with permanent calyces, anthocodiae capable of total retraction into

calyces but are mainly preserved slightly exserted. Calyces 1-2 mm in length.

Polyps present on all sides of branches or restricted to opposite longitudinal

rows, parallel to plane of colony. Axis thick, coenenchyme thin and densely

spiculiferous. Anthocodia with crown and points composed of spindles arranged

en chevron. These spindles are usually somewhat bent, tuberculated, and

measure 0,12—0,35 mm in length.

Calyx sclerites are irregularly-shaped thorn scales, usually 0,15—0,20 mm in

length. The surfaces of these sclerites usually have several large tubercles.

Coenenchymal sclerites are 0,08—0,15 mm long. These are usually in the form of

butterflies or thorn scales with coarse tuberculation. Colony colour brown, or

grey in life and when preserved dry or in alcohol.

Distribution

Known only between East London, eastern Cape Province, and along the

Transkei coast to Durban, Natal; 90-360 m in depth (Williams in press a).

DD ANNALS OF THE SOUTH AFRICAN MUSEUM

Specific type locality not known. The present work represents a new record for

the genus in southern Africa.

Remarks

Hickson (1900: 87) identified a specimen from 155 m off East London as

Villogorgia mauritiensis Ridley, 1882. Examination of Hickson’s specimen shows

no significant differences with specimens of Astromuricea fusca from the same

locality. I therefore consider that Hickson’s material should be attributed to this

species. The South African material has characteristics conforming to the genus

Astromuricea and differing from those of Villogorgia or other plexaurid genera.

These are the possession of thorn scales that are mostly higher than wide and

consist of several diverging processes, not with distinct basal processes; four

radiates with central projections are not present in the coenenchyme; antho-

codiae have numerous relatively small sclerites forming a distinctive crown and

points.

Thomson (1911: 878) originally assigned this species to the genus Muri-

ceides Studer, 1887. However, the form of the sclerites is not typical of this

genus, but conforms rather to those of the genus Astromuricea Germanos, 1896

(Dr F. M. Bayer pers. comm.), and therefore Thomson’s generic placement

Fig. 17. Astromuricea fusca Thomson, 1911. An entire colony, 205 mm in height.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

—

Fig. 18. Astromuricea fusca (Thomson, 1911). A.A single polyp with partially retracted

anthocodia, showing placement of sclerites in anthocodia and calyx; total length of polyp

1,4 mm. B. Anthocodial sclerites. C. Calyx sclerites. D. Coenenchymal sclerites.

Scale bar = 0,15 mm.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

must be regarded as incorrect. The classification of paramuriceid genera within

the Plexauridae is presently unsettled. The placement of Thomson’s species in

the genus Astromuricea is here considered provisional, since it may possibly be a

synonym of Paramuricea Kolliker, 1865, or Placogorgia Studer, 1887. There are

also similarities in some respects to the genus Echinogorgia K6lliker, 1865

(Dr F. M. Bayer pers. comm.).

Genus Acanthomuricea Hentschel, 1903

Acanthomuricea Hentschel, 1903: 650.

Diagnosis

Colonies are very sparsely branched or unbranched. Polyps are mostly

arranged in two lateral rows. Calyces are prominent and conical in shape.

Sclerites of the calyces and coenenchyme are relatively large tuberculate

spindles. Those of the calyces are more or less longitudinally placed.

A genus of at least two species from the Indo-West Pacific.

Type species. Acanthomuricea biserialis Hentschel, 1903; Amboina

(Indonesia).

Acanthomuricea pulchra (Thomson, 1911) comb. nov.

Figs 19D-G, 20-21

Psammogorgia pulchra Thomson, 1911: 881, pl. 43 (fig. 5), pl. 45 (figs 3a—b). Ktikenthal, 1919:

240; 1924: 110. Stiasny, 1940: 18, text-fig. A.

Material

SAM-—H3610, off Gonubie, eastern Cape Province (33°04,7'S 28°07,2'E),

90 m, 17 July 1984, 4 colonies, dredge, G. C. Williams, R.V. Meiring Naude.

SAM-H3616, off East London, eastern Cape Province (33°15,0’S 27°58,0’E),

85 m, 17 July 1984, 2 colonies, dredge, G. C. Williams, R.V. Meiring Naude.

Description

Colonies examined measure 20—40 mm in height. Colonies are unbranched

or sparsely branched dichotomously. Anastomoses absent. Apices of branches

sometimes slightly clavate. Anthocodiae with conspicuous crown and points of

red, finely tuberculated spindles, some slightly bent, 0,20-—0,65 mm in length.

Surface of colony warty in appearance due to placement of numerous calyces

mostly on all sides of branches. Sclerites of calyces and coenenchyme are mainly

yellow, coarsely tuberculate clubs and spindles, 0,45—1,00 mm in length. Colony

colour yellow with red anthocodiae.

Distribution

Known only from the coasts of the eastern Cape Province and southern

Transkei, 85-100 m in depth (Williams in press a). Type locality is Cape

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA PD lis)

Fig. 19. Scanning electron micrographs. A-C. Astromuricea fusca (Thomson, 1911); calyx

sclerites. A. 0,225 mm. B.0,3 mm. C. Two sclerites, each 0,2 mm. D-—G. Acanthomuricea

pulchra (Thomson, 1911). D. Calyx sclerite, 0,6 mm. E. Anthocodial sclerite, 0,46 mm.

F. Anthocodial sclerite, 0,32 mm. G. Calyx sclerite, 0,78 mm.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

Morgan, border of Cape Province and Transkei (Thomson 1922: 881). The

present work represents the first record for the genus in southern Africa.

Remarks

This species is distinguished by its small size (<40 mm tall), sparse

dichotomous branching, yellow coloration with red anthocodiae, and coarsely

tuberculated coenenchymal sclerites that are robust spindles and clubbed

spindles.

Thomson (1911: 881) originally assigned this species to the genus

Psammogorgia Verrill, 1868. Psammogorgia is a Panamic genus, endemic to the

eastern Pacific, and shows no relation to the South African material (Dr F. M.

Fig. 20. Acanthomuricea pulchra (Thomson, 1911). A. An entire colony, 32 mm in length.

B. Colony, length 23 mm. C. Colony, length 18 mm. D. A single polyp showing placement of

sclerites in partially retracted anthocodia and calyx; total length of polyp 1,3 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 217

Bayer pers. comm.). Thomson’s generic placement must therefore be

considered incorrect. The present generic designation must be considered

provisional since several similar paramuriceid genera are presently considered

problematical, including Muricella of many authors but not Verrill, Astrogorgia

Fig. 21. Acanthomuricea pulchra (Thomson, 1911). A. Anthocodial sclerites. B. Sclerites

from the calyces and coenenchyme. Scale bar =0,3 mm.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Verrill, 1868, Anthomuricea Studer, 1887, and Acanthomuricea Hentschel, 1903

(Dr F. M. Bayer pers. comm.).

Other species of the family Plexauridae

Thomson (1917: 41, pl. 1 (figs 4, 7), pl. 4 (fig. 5)) described Stenogorgia

capensis from Algoa Bay. Thomson’s material is not presently available for

examination and no new material assignable to this species has been obtained.

Stenogorgia Verrill, 1883, is considered by Deichmann (1936), Grasshoff (1977)

and Bayer (1981: 945) as a junior synonym of Swiftia Duchassaing & Michelotti,

1864. Thomson’s description and figures of Stenogorgia capensis seem to fit the

genus Leptogorgia (family Gorgoniidae); it strongly resembles the three very

similar species (assignable to Leptogorgia by the presence of symmetrical tuber-

culation of the coenenchymal capstans) and described by Kikenthal (1919) as

Leptogorgia abietina, Leptogorgia tenuissima, and Leptogorgia pusilla, all from

the South African south coast. These three taxa mav in fact represent a single

species. However, the type material is not available for examination.

Family Gorgoniidae Lamouroux, 1812

Axis composed purely of horn with a narrow chambered central core, and

dense cortex. Polyps monomorphic, capable of total retraction into coenen-

chyme. Calyces present or absent. Anthocodia usually flattened rods, not

forming a conspicuous crown and points. Coenenchymal sclerites are spindles

with whorls of tuberculation, in the form of radiates, balloon clubs, scaphoids,

capstans, clubs, or double wheels.

Fourteen genera of the Atlantic, Mediterranean, Caribbean, and Indo-

Pacific. At least three genera in southern Africa.

Genus Eunicella Verrill, 18696

Eunicella Verrill, 1869b: 425. Kiikenthal, 1919: 255; 1924: 133.

Diagnosis

Surface layer of coenenchyme with numerous balloon clubs overlying an

interior layer of girdled spindles.

A genus of perhaps 12 species of the Atlantic, Mediterranean, southern

Africa, and Japan.

Type species. Gorgonia verrucosa Pallas, 1766; Spain.

Eunicella albicans (K6lliker, 1865)

Figs 1D, 22-24

Gorgonia palma var. alba (non Pallas, 1766) Esper, 1797: 153, pl. 40.

Gorgonia albicans Kolliker, 1865: 139. Hickson, 1904: 228. Thomson, 1917: 36.

Eunicella palma (non Pallas, 1766) Verrill, 1869b: 426.

Eunicella alba Tixier-Durivault, 1954: 624.

Eunicella albicans Kiikenthal, 1919: 269; 1924: 136. Velimirov, 1971: 267, figs la—b, 2.

219

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

2 = Ba ° SS

[oS TOI

oc BEE

SOO aaah

O50 (-) O90

Sa 0.00.09 30.00 6-00 8a

length.

, 300 mm

lony

ire CO

lla albicans (K6lliker, 1865). An ent

Fig. 22. Eunice

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM-H3257, Algoa Bay (33°59’S 25°40’E), 10 m, February 1984, 1 colony,

SCUBA, W.R. Liltved. SAM—H3262, Algoa Bay (33°59'S 25°40’E), 10 m,

February 1984, 1 colony, SCUBA, W. R. Liltved. SAM—H3917, Algoa Bay

(33°59'S 25°40'E), 10 m, 20 May 1984, 1 colony, SCUBA, G. C. Williams.

Description

Colonies examined range in length from 185 mm to 650 mm. Colonies are

upright, branching is dichotomous and planar. Branches arise from a single basal

main stem. Branches straight or slightly curved, not conspicuously sinuous.

Anastomosis apparently does not occur. Branches are conspicuously flattened in

the proximal region of the colony. These often have a series of longitudinal lines

evident on the surface. The plane of the flattened branches is parallel to the

plane of colony. The distal regions of the ultimate branches are cylindrical,

sometimes slightly flattened. Polyps present on all sides of branches, forming

hemispherical calyces or totally retracting into coenenchyme, forming slits

Fig. 23. Eunicella albicans (Kolliker, 1865). Coenenchymal sclerites. Scale bar = 0,15 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

Fig. 24. Scanning electron micrographs. Eunicella albicans (K6lliker, 1865); coenenchymal

sclerites. A.0,125 mm. B. Three sclerites 0,105—0,120 mm. C. 0,085 mm. D. 0,130 mm.

E. Four sclerites 0,975—0,195 mm. F. Six sclerites 0,105—0,190 mm. G. 0,085 mm.

2D ANNALS OF THE SOUTH AFRICAN MUSEUM

<0,5 mm long. Coenenchymal sclerites are medium-sized balloon clubs with two

crowns, 0,11—0,15 mm in length; and girdled spindles, 0,17—0,19 mm in length.

Colony colour in life is light orange, fading to white in alcohol.

Distribution

Presently known from only two localities along the south coast of South

Africa: False Bay and Algoa Bay; 10-14 m in depth (Williams in press a);

common at both these localities. The type locality is South Africa, exact locality

unknown.

Remarks

Eunicella albicans differs from other southern African species of Eunicella

by the possession of coenenchymal sclerites that are medium balloon clubs with

two crowns (0,11—0,15 mm in length).

Fig. 25. Eunicella papillosa (Esper, 1797). An entire colony, 180 mm in height.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 223

Eunicella papillosa (Esper, 1797)

Figs 1E, 25-28

Gorgonia papillosa Esper, 1797: 173, pl. 50. Kélliker, 1865 (part.): 139, pl. 18 (figs 25, 26,

27a).

Eunicella papillosa Verrill, 1869b: 425. Studer, 1879: 656. Wright & Studer, 1889: 145.

Hickson, 1900: 86, pl. 5 (fig. C). Thomson, 1911: 880, pl. 43 (figs 2-3); 1917: 24. Kiiken-

thal, 1919: 263, figs 126-128; 1924: 136. Molander, 1929: 11. Stiasny, 1938: 30, pl. 3

GigwI2) reply 7 (ig. 26); 1940: 20, pl 4 (fig. 20). Broch, 1939: 26, figs 22, 23. Tixier-

Durivault, 1954: 624. Day et al., 1970: 16. Velimirov, 1971: 271, figs 5-6. Day, 1974: 35,

fig. pg. 35. Branch & Branch, 1981, pl. 36.

Eunicella rigida Kikenthal, 1908: 503; 1919: 257, text-figs 118-119, pl. 30 (fig. 3); 1924: 134,

fig. 98

Material

SAM-—H3296, Hottentot’s Huisie, west coast Cape Peninsula (33°59’'S

18°21’E), 23 m, 15 June 1983, SCUBA, W. R. Liltved. SAM—H3303, off Llan-

dudno, west coast Cape Peninsula (34°00’S 18°20'E), 21 m, 24 January 1984,

1 colony, SCUBA, G. C. Williams. SAM—H9972, Seal Island, False Bay (34°08’S

18°35’E), 16-18 m, 30 October 1902, 1 colony, dredge, S.S. Pieter Faure survey,

PF 15801A. SAM—H3397, off Danger Point (34°40’S 19°20’E), 25 m, 28 April

1984, 2 colonies, SCUBA, W. R. Liltved, Sea Fisheries Research Institute, Line

Fish Survey. SAM—H3918, Algoa Bay (33°59’S 25°40'E), 12 m, 16 May 1984,

1 colony, SCUBA, G. C. Williams, R.V. Meiring Naude. SAM-—H3920, off

Nthlonyane River mouth, Transkei (32°16,7'S 29°06,0’E), 300 m, 5 July 1985,

several fragments, dredge, G. C. Williams, R.V. Meiring Naude. SAM—H3877,

off Port Edward (31°05,8’S 30°18,8’E), 140 m, 8 July 1985, 3 colonies, dredge,

G.C. Williams, R.V. Meiring Naude. SAM—H3918, off Umhlanga, Natal

(29°45’S 31°10’E), 68 m, 17 December 1984, 2 colonies, dredge, W. R. Liltved.

Description

Colonies examined measure 40—180 mm in length. Colonies grow upright or

in bushy masses, planar or multiplanar. Branching dichotomous to slightly

irregular, arising from a single basal main stem. Branches cylindrical, ultimate

branches sometimes slightly clavate. Branches usually straight or slightly curved,

not sinuous. Anastomosis apparently does not occur. Main stem cylindrical,

rarely somewhat flattened. Polyps numerous on all sides of branches, rarely

limited to two longitudinal tracts along opposite margins of branches. Polyps

usually form hemispherical to cylindrical calyces, 1-2 mm in length. Polyps also

capable of total retraction into coenenchyme, not forming calyces. Coenenchy-

mal sclerites are small balloon clubs with two crowns, 0,08—0,1 mm in length,

and girdled spindles, 0,10-—0,23 mm long. Colony colour pale yellow, pinkish, or

white in life, fading to white or greyish-white in alcohol.

Distribution

The most common and widespread member of the genus in southern

Africa; known from Cape Columbine on the Atlantic coast of the Cape Province

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. Eunicella papillosa (Esper, 1797). Coenenchymal sclerites. Scale bar = 0,1 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 225

to Durnford Point, northern Natal, intertidal to 360 m in depth (Williams in

press a). Type locality South Africa, exact locality not known.

Remarks

Material collected from deeper water (100 m to more than 300 m) from the

eastern Cape to southern Natal, differs slightly from typical shallow-water

material (0-100 m) in a number of respects. Kiikenthal (1908) described this

form as a separate species, E. rigida. Colonies from deeper water are more

Fig. 27. Scanning electron micrographs. Eunicella papillosa (Esper, 1797), coenenchymal scle-

rites. A.0,070mm. B.0,068mm. C. 0,065 mm. D. 0.075 mm. E. Two sclerites, each

0,060 mm. F. Three sclerites, 0,060—0,075 mm.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 28. Scanning electron micrographs. Eunicella papillosa (Esper, 1797), coenenchymal scle-

rites. A. 0,09 mm. B.0,08mm. C.0,10mm. D. 0,092 mm. E. Group of four sclerites

0,075-0,150 mm. F. 0,082 mm. G.Two sclerites 0,090-0,130 mm. H. Two sclerites,

0,0709-0,120 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA DD,

sparsely branched, often with a more lanky appearance and have fewer polyps,

these being spaced further apart, possess calyces that are very prominent

(1-2 mm long and 1-1,5 mm wide), and have a greater proportion of balloon

clubs with broad terminal crowns. Despite those differences, I consider the

various shallow- and deeper-water forms to be conspecific, since a gradual series

of morphological differences is evident and as clear-cut boundaries cannot be

established.

Eunicella papillosa differs from other southern African members of the

genus by the possession of coenenchymal sclerites that are mainly small balloon

clubs with two crowns (0,08—0,10 mm in length).

Fig. 29. Eunicella tricoronata Velimirov, 1971. An entire colony, 300 mm in length.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eunicella tricoronata Velimirov, 1971

Figs 1F, 29-32

Gorgonia papillosa (non Esper, 1797) Kolliker, 1865 (part.), pl. 18 (fig. 27b).

Eunicella albicans (non Kolliker, 1865) Molander, 1929: 10, figs 1-2. Broch, 1939: 27.

Eunicella alba (non Esper, 1796) Broch, 1939: 27.

Eunicella tricoronata Velimirov, 1971: 269, figs 3-4.

Material

SAM-—H3297, off Sunny Cove, False Bay (34°09’S 18°26’E), 10 m, 23 April

1983, 1 colony, SCUBA, G. C. Williams. SAM—H3346, Algoa Bay (33°50’'S

25°40’E), 11 m, 20 May 1984, 1 colony, SCUBA, G. C. Williams. SAM—H3333,

Algoa Bay (33°50’S 25°40’E), 11 m, May 1984, 1 dried colony, SCUBA, T. M.

Gosliner and G. C. Williams.

Description

Colonies examined range in length from 110 mm to 420 mm. Colonies grow

upright. Branching planar, dichotomous, arising from a single basal main stem.

Branches mostly flattened in plane of colony. Ultimate branches mostly cylindri-

Fig. 30. Eunicella tricoronata Velimirov, 1971. Coenenchymal sclerites. Scale bar = 0,15 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 229

cal to slightly flattened. Branches usually conspicuously sinuous, especially in

larger colonies (>25 mm in height). Anastomosis occurs only rarely. Polyps

numerous on all sides of branches, forming hemispherical calyces or low

rounded protuberances (<1,00 mm in length). Polyps also capable of total

retraction into coenenchyme. Coenenchymal sclerites are elongate balloon clubs

with three crowns, 0,10—0,15 mm long, and robust, often copiously tuberculated

girdled spindles, 0,12—0,20 mm long. Colony colour vivid orange in life, fading

to white or cream-white in alcohol.

Fig. 31. Scanning electron micrographs. Eunicella tricoronata Velimirov, 1971; coenenchymal

sclerites. A. 0,105 mm. B-C. 0,11 mm. D.0,10 mm. E. 0,115 mm. F. 0,11 mm.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

South coast of South Africa, from False Bay to East London, 9-36 m in

depth. Common in False Bay, off Cape Agulhas, and in Algoa Bay (Williams in

press a). The type locality is False Bay.

Remarks

Eunicella tricoronata differs from other southern African Eunicella species

by the possession of elongate balloon clubs (0,1—0,15 mm long) in the coenen-

chyme; these have three crowns.

Fig. 32. Scanning electron micrographs. Eunicella tricoronata Velinurov, 1971, coenenchymal

sclerites. A. 0,125 mm. B. 0,120 mm. C. 0,145 mm. D. 0,130 mm. E. 0,150 mm. F. 0,135 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 231

Genus Leptogorgia Milne Edwards & Haime, 1857

Leptogorgia Milne Edwards & Haime, 1857: 163. Kitkenthal, 1924: 323. Grasshoff, 1988: 97

(Lit!).

Lophogorgia Milne Edwards & Haime, 1857: 167. Kiikenthal, 1924: 322.

Filigorgia Stiasny, 1937: 307; 1939: 301. Bayer, 1956: F206.

Diagnosis

Sclerites of the coenenchyme are symmetrical radiates (capstans) or

spindles with whorls of tubercles that are symmetrically sculptured. Adjacent

tubercles do not fuse to form discs.

A genus of at least eight species of the Mediterranean, Atlantic, and

southern Africa and the subantarctic.

Type species. Gorgonia viminalis Pallas, 1766; Mediterranean Sea.

Leptogorgia barnardi Stiasny, 1940

Figs 33-36

Leptogorgia barnardi Stiasny, 1940: 26, text-fig. D, pl. 4 (figs 18-19).

Material

SAM-—H1285, St Francis Bay, southern Cape Province (34°05’S 25°05’E),

62 m, 11 May 1906, 1 colony, large trawl, S.S. Pieter Faure survey, PF 18834.

SAM-—H3266, Gansbaai, south-western Cape Province (34°25’S 19°15’E), 33 m,

10 October 1983, 1 colony, dredge, Sea Fisheries Research Institute. SAM-—

H3926, off Cape Agulhas (34°50’'S 20°10'E), 53 m, 24 February 1985, 1 colony,

SCUBA, W. R. Liltved, Sea Fisheries Research Institute, Line Fish Survey.

SAM-—H3927, off Cape Agulhas (34°50’S 20°10’E), 33 m, 24 February 1985,

2 colonies, SCUBA, W. R. Liltved, Sea Fisheries Research Institute, Line Fish

Survey. SAM—H3928, off Cape Agulhas (34°50’S 20°10’E), 30 m, 11 February

1985, 5 colonies, SCUBA, W. R. Liltved, Sea Fisheries Research Institute, Line

Fish Survey.

Description

The colonies examined measure 210-320 mm in length. Colonies are

upright, lank in appearance. Branching is dichotomous and planar. Anastomosis

not present. Most branching takes place in the proximal region of the colony

above the single basal stem. The terminal branches are long, thin, whip-like and

cylindrical (1,0—2,0 mm in diameter; up to 200 mm in length). Basal main stem

cylindrical, rarely slightly flattened. Polyps numerous, present on all sides of

branches, capable of total retraction into coenenchyme, rarely forming low

rounded calyces (<1,0 mm high). Coenenchymal sclerites are mainly elongate

capstans with whorls of tubercles well-separated (0,06—0,16 mm in length).

Colony colour permanent white.

DSP) ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Leptogorgia barnardi Stiasny, 1940. An entire colony, 240 mm in height.

Distribution

Presently known from four localities on the South African south coast—

Gansbaai, Cape Agulhas, St Francis Bay, and Cape Morgan. Stiasny (1940: 26)

recorded the holotype from Cape Morgan (border of Cape Province and Trans-

kei) from 93 m in depth.

Remarks

Leptogorgia barnardi can be distinguished from other southern African

Leptogorgia species by the possession of many coenenchymal sclerites that are

long thin capstans with well-separated whorls of tuberculation, white colony col-

oration, and dichotomous branching mostly in the proximal region of the colony

with long thin terminal branches.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

Fig. 34. Leptogorgia barnardi Stiasny, 1940. An entire colony, 350 mm in height.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Leptogorgia barnardi Stiasny, 1940. Coenenchymal sclerites. Scale bar = 0,15 mm.

Leptogorgia capensis (Hickson, 1900)

Figs 37-39

Gorgonia capensis Hickson, 1900: 83, pl. 5 (figs A, A’, A’’, A’’’). Thomson, 1911: 887.

Stiasny, 1940: 29, pl. 2 (figs 9-10).

Lophogorgia capensis Molander, 1929: 11, fig. 3. Tixier-Durivault, 1954: 626. Day et al.,

1S 70 Ge

Leptogorgia capensis Stiasny, 1940: 29, text-fig. F.

Material

SAM-H1315, lectotype (designated herein), off Cape St Blaize, south coast

of Cape Province (34°15’S 22°10'E), 73m; 8 June 1898, i icolomys larsexotten

trawl, S.S. Pieter Faure survey, PF 28. SAM-—H657, paralectotype (designated

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA IBS

Fig. 36. Scanning electron micrographs. Leptogorgia barnardi Stiasny, 1940, coenenchymal

Selenites 0,122 mm. B.05125mm. CC. 0,11 mm. D. 0,095 mm. E. Group of three scle-

rites 0,080-0,085 mm. F. Two sclerites 0,085—0,090 mm.

herein), same data as SAM-—H1315, 1 fragmented colony. SAM—H3226, False

Bay, south-western Cape Province (34°12,8'S 18°36,5'’E), 46 m, 10 September

1953, 1 fragmented colony, dredge, University of Cape Town Ecological Survey.

SAM-—H3921, Algoa Bay (33°50'S 25°50’'E), depth not recorded, October 1983,

1 dried colony, bottom trawl, Ocean Pearl.

Description

Colonies examined measure 130-330 mm in length. Colonies grow upright

but present a lank appearance due to the long and thin ultimate branches

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 37. Leptogorgia capensis (Hickson, 1900). An entire colony, 340 mm in height.

(1,5—2,5 mm in diameter and up to 170 mm long) that may be somewhat droop-

ing. Branching is planar and dichotomous to slightly lateral. Branches are

cylindrical and arise from a single basal stem. Anastomosis apparently does not

occur. Retracted polyps may form low rounded protuberances (<1,0 mm high)

or totally retract into coenenchyme leaving minute slits (<0,5 mm long) on the

surfaces of the branches. Polyps present on all sides of branches. Coenenchymal

sclerites are relatively large eight radiates (capstans) or girdled spindles,

0,07—0,17 mm in length. Colony colour permanent yellow due to sclerite color-

ation. Axis deep reddish-brown.

oS)

—

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

Fig. 38. Leptogorgia capensis (Hickson, 1900). Coenenchymal sclerites. Scale bar = 0,15 mm.

Distribution

Presently known from the south coast of South Africa from False Bay to

Algoa Bay, 46—73 m in depth. Type locality Cape St Blaize, Cape south coast.

Remarks

This species has sclerites with symmetrical whorls of tubercles and no scaph-

oids (C-shaped spindles with tubercles on the convex side modified or reduced)

are present. Thus it is assignable to the genus Leptogorgia (which may have

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Yj je“

Fig. 39. Scanning electron micrographs. Leptogorgia capensis (Hickson, 1900); coenenchymal

sclerites. A. 0,105 mm. B. 0,08 mm. C. 0,072 mm. D. 0,062 mm. E. 0,10 mm. F. 0,085 mm.

G. 0,070 mm. H. 0,085 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 239

some unilaterally spinose spindles), not Gorgonia (which has scaphoids as well

as capstans).

Leptogorgia capensis is distinguished from other southern African Lepto-

gorgia species by the possession of coenenchymal sclerites of relatively uniform

shape and size that are large robust capstans (mostly 0,10—0,17 mm in length),

lanky appearance with long thin ultimate branches (c. 2,0 mm in width and up

to 170 mm long), dichotomous to slightly lateral branching, and yellow colony

coloration.

Leptogorgia palma (Pallas, 1766)

Figs 1C, 40-45

Gorgonia palma Pallas, 1766: 189. Esper, 1791-99 (part.): 32, pl. 5. Kolliker, 1865: 139.

Gorgonia flammea Ellis & Solander, 1786: 80, pl. 11. Hickson, 1900: 81, pl. 5 (fig. B).

Thomson, 1911: 888; 1917: 36, pl. 1 (figs 3, 7), pl. 4 (fig. 6).

Lophogorgia palma Milne Edwards & Haime, 1857: 167.

Lophogorgia crista Mobius, 1861: 7, pl. 2 (figs 1-7). Kikenthal, 1919: 638, text-fig. 288, pl. 32

(fig. 18), pl. 33 (fig. 19); 1924: 322. Stiasny, 1940: 22, text-fig. B, pl. 3 (figs 13-15). Tixier-

Durivault, 1954: 625.

Leptogorgia crista Verrill, 1869b: 421.

Leptogorgia flammea Verrill, 1869b: 421.

Lophogorgia flammea Wright & Studer, 1889: 150, 151. Bielschowsky, 1918: 18; 1929: 77.

Molander, 1929: 11. Stiasny, 1940: 24, text-fig. C, pl. 3 (figs 16-17). Tixier-Durivault,

iat -o2>- Way et al., 1970: 16. Day, 1974: 35, fig. p. 35. Branch & Branch, 1981: 151,

jOlle 335

Leptogorgia palma Grasshoff, 1988: 115, pl. 7 (fig. 4), pl. 10 (fig. 6), pl. 13 (fig. 3).

Material

SAM-—H3670, off East London (33°02’S 27°56'E), 30-35 m, 16 July 1984,

1 colony, dredge, G. C. Williams, R.V. Meiring Naude. SAM-—H1326, Algoa

Bay (33°53’S 25°51'E), 47 m, 6 December 1898, 1 dried colony, large trawl, S.S.

Pieter Faure survey, PF 706B. SAM-—H3343, Algoa Bay (33°50'S 25°40’B),

11 m, 20 May 1984, 5 colonies, SCUBA, G. C. Williams. SAM—H3344, Algoa

Bay (33°50'S 25°40'E), 11 m, 20 May 1984, 4 colonies, SCUBA, G. C. Wil-

hams. SAM—H3342, off Danger Point, south-western Cape Province (34°35'S

19°20’E), 25 m, 24 April 1984, 4 colonies, SCUBA, W. R. Liltved, Sea Fisher-

ies Research Institute, Line Fish Survey. SAM—H3600, Sunny Cove, False Bay

(34°09’S 18°27’E), 10 m, 23 April 1983, 1 colony, SCUBA, G. C. Williams.

Description

Colonies examined range in length from 120 mm to | 100 mm. Colonies are

upright. Branching is sparse to copious, planar and pinnate, dichotomous, or

lateral. Branches arise from a single basal stem that is usually conspicuously flat-

tened parallel to the plane of the colony and up to 30 mm wide. Primary

branches also usually flattened. Ultimate branches are mostly cylindrical or

slightly flattened, often very flexible (2-4 mm in width, 25-200 mm in length).

Anastomosis does not occur. Retracted polyps form very low rounded protuber-

ances or retract completely into coenenchyme, forming minute slits on the

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

surfaces of the branches (0,5 mm long). Polyps are present on all sides of ulti-

mate branches, but are usually present only on opposite edges of the flattened

lower branches and basal stem, and not present or only scattered sparingly on

the flattened faces of these branches. Coenenchymal sclerites are mainly com-

pact capstans (0,04—0,10 mm long). A few capstans may be less compact with

whorls of tubercles well spaced. Colony colour is vivid orange, red-orange,

or deep brick-red; permanent due to sclerite coloration. Axis is dark brown to

black.

Distribution

Commonly encountered along the coast of South Africa from the west coast

of the Cape Peninsula to Durban, Natal; intertidal to 100 m in depth (Williams

in press a). Type locality Cape of Good Hope, but exact locality not known.

Fig. 40. Leptogorgia palma (Pallas, 1766). An entire colony, 250 mm in height, showing

pinnate branching.

241

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

PoDGOOS'

ay a LY

. Y

0 ¥ f- 0 4°

showing

;

350 mm in height

An entire colony,

, 1766).

Leptogorgia palma (Pallas

Fig. 41.

pinnate and lateral branching.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Leptogorgia palma is a highly variable species. Lophogorgia crista was pre-

viously differentiated on the basis of dichotomous branching and dark-red

coloration, whereas L. palma was defined by pinnate branching and red-orange

coloration. After having examined a large suite of specimens from throughout

the ranges of both forms (both older material as well as a great many recently

collected colonies), I conclude that the two forms are conspecific as no clear-cut

morphological boundaries can be substantiated. Some colonies have dichoto-

mous branching and are orange in colour, whereas others are dark-red with

pinnate branching. Also, some colonies do not exhibit distinct dichotomous or

pinnate branching but are intermediate, with irregular or lateral branching. The

size and shape of the coenenchymal capstans of the two forms are virtually

indistinguishable.

Thomson & Henderson (1906: 432, pl. 32 (figs 5—7)) recorded Lophogorgia

crista from the Cape Verde Islands, and noted several differences between their

material and South African specimens. I consider this a dubious record.

Ca”

Fig. 42. Leptogorgia palma (Pallas, 1766). An entire colony, 210 mm in height, showing

dichotomous branching.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 243

Leptogorgia palma is distinguished from other southern African members of

the genus by the possession of red-orange to wine-red coloration, coenenchymal

sclerites that are mainly compact capstans (0,04-0,10 mm long), ultimate

branches that are 25-200 mm long, and anastomosis 1s not present. Older col-

onies may exceed 2 m in height, this being the largest of all southern African

gorgonians.

The orange pinnately branched form is a common and conspicuous constitu-

ent of horizontal reefs and the sides of boulders in False Bay. Both the orange

Fig. 43. Leptogorgia palma (Pallas, 1766); coenenchymal sclerites. Scale bar = 0,1 mm.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 44. Scanning electron micrographs. Leptogorgia palma (Pallas, 1766); coenenchymal scle-

rites. A. 0,048 mm. B. 0,037 mm. C. 0,04 mm. D. 0,042 mm. E. 0,039 mm. F. 0,065 mm.

pinnate form and the wine-red, dichotomously branched form are common on

shallow reefs in Algoa Bay.

Leptogorgia gilchristi (Hickson, 1904) comb. nov.

Figs 46—49

Eugorgia Gilchristi Hickson, 1904: 230, pl. 9 (figs 15, 19). Thomson, 1917: 38.

Leptogorgia alba var. capensis Thomson, 1917: 29.

Leptogorgia aurata Thomson, 1917: 32, pl. 1. (fig. 5), pl. 4 (fig. 2).

Eugorgia lineata Thomson, 1917: 39, pl. 2 (fig. 3), pl. 5 (fig. 2).

Eugorgia gilchristi: Kikenthal, 1924: 347. Stiasny, 1940: 27, text-fig. E.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 245

Fig. 45. Scanning electron micrographs. Leptogorgia palma (Pallas, 1766); coenenchymal scle-

rites. A.0,04mm. B. 0,045 mm. C.0,i2mm. D.0,08mm. E. Group of three sclerites,

0,035—0,060 mm. F. Group of three sclerites, 0,06—0,08 mm.

Material

SAM-—H3382, off Danger Point, south-western Cape Province (34°41’S

19°20’E), 44 m, 16 April 1984, 1 colony, SCUBA, W. R. Liltved. SAM—H3925,

Oi Cape Aculhas (34°50’S 20°10'E), 45m, 27 February 1985, 1 colony,

SCUBA, W.R. Liltved, Sea Fisheries Research Institute, Line Fish Survey.

SAM-—H969 (holotype of Eugorgia lineata), off Cape St Blaize (34°12’S

PP Nie) 27-33 m, ls July 1898) 5 colonies, dredge, SS. Rieter Faure survey,

PF 210. SAM-H3340, Algoa Bay (33°50’'S 25°40'E), 11m, 20 May 1984,

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

1 colony, SCUBA, G. C. Williams. SAM—H978 (holotype), St Francis Bay

(34°5'20"S_ 25°43'E), 95 m, 1 November 1898, 1 partial colony, dredge, S.S.

Pieter Faure survey, PF 622. SAM—H982 (holotype of Leptogorgia aurata) and

SAM-—H983 (type specimen of Leptogorgia alba var. capensis), Durnford Point,

Natal-(28°25'S 32°30 E), 82 mi) 28) February 19015 1 colony cach dredecussns:

Pieter Faure survey, PF 12164 and PF 12165.

Description

Colonies examined range in length from 110 mm to 360 mm. Colonies grow

upright. Branching is planar and pinnate, bushy or arising from a single basal

stem. Branching is very copious, anastomoses are frequent. Proximal branches

and main stem often flattened. Ultimate branches mostly cylindrical, 5-25 mm

in length, often curving and ascending. Polyps present on all sides of ultimate

branches, usually forming hemispherical protuberances during retraction. Flat-

tened faces of proximal branches mostly free of polyps. Coenenchymal sclerites

are capstans exhibiting a wide range of sizes, 0,03—0,13 mm in length. Some of

the larger ones may be slightly bent. Colony colour highly variable: bright

WSK

ad)

Yi,

SILK

SRS

iL [A

Oy,

LQy,

Fig. 46. Leptogorgia gilchristi (Hickson, 1904). An entire colony, 300 mm in height.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 247

Fig. 47. Leptogorgia gilchristi (Hickson, 1904). An entire colony, 165 mm in height.

yellow, white, pink, mauve, rust-orange, brick-red. Some colonies are bicol-

oured white and pink, yellow and orange, on white and red. Colour is conserved

in alcohol.

Distribution

Recorded from the south and east coasts of South Africa from Danger

Point to northern Natal, 10-95 m in depth. Type locality is St Francis Bay, Cape

south coast.

Remarks

Hickson (1904: 230) described Eugorgia gilchristi from St Francis Bay

(southern Cape Province) and Thomson (1917: 39) described Eugorgia lineata

from Cape St Blaize (also southern Cape Province). Examination of both Hick-

son’s and Thomson’s type specimens show that neither belong to the genus

Eugorgia Verrill, 1868, as double wheel sclerites (modified capstans with tuber-

cles of two whorls fused into discs or wheels) are absent. The capstans are

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 48. Leptogorgia gilchristi (Hickson, 1904). Coenenchymal sclerites. Scale bar = 0,1 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 249

virtually all symmetrically sculptured (even though some may be slightly bent

along the longitudinal axis). Both specimens must therefore be allocated to the

genus Leptogorgia. Comparison of the two specimens shows no appreciable dif-

ferences, except for the size of the calyces, which vary depending on the state of

retraction. I therefore consider Eugorgia lineata to be a junior synonym of

Leptogorgia gilchristi. Comparison of the types of Leptogorgia alba var. capensis

Thomson, 1917 (p. 29), and Leptogorgia aurata Thomson, 1917 (p. 32) with

Hickson’s type shows very minor differences in calyx and sclerite size, differ-

ences that can be accounted for by degree of retraction and intraspecific

Fig. 49. Scanning electron micrographs. Leptogorgia gilchristi (Hickson, 1904); coenenchymal

sclerites. A. 0,11 mm. B. 0,05 mm. C. 0,035 mm. D. 0,08 mm. E. Group of four sclerites,

0,065—0,100 mm. F. 0,11 mm.

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

variation. I therefore consider these two species to be conspecific with Lepto-

gorgia gilchristi.

Leptogorgia gilchristi is distinguished from other southern African species

of the genus by the possession of copious pinnate branching with ascending and

curved ultimate branches (<25 mm long) and anastomosis common, a broad

range of coenenchymal sclerite size (0,03—0,13 mm in length), and extremely

variable colony coloration.

Genus Rumphella Bayer, 1955

Rumphella Bayer, 1955: 212.

Diagnosis

Sclerites of the coenenchyme are mainly conspicuous wart clubs, some

radiates also present.

An Indo-Pacific genus of three or four species.

Type species. Plexaura aggregata Nutting, 1910; Malay Archipelago.

Remarks

An unidentified species has recently been collected at Sodwana Bay, north-

ern Natal. It is present in sandy gullies or depressions in coral reefs at 15-18 m

in depth (Williams 1989: 152; in press c).

This represents a new record for the genus Rumphella in southern Africa.

Other species of the family Gorgoniidae

At present, only three gorgoniid genera are known from southern Africa:

Eunicella, Leptogorgia, and Rumphella. Several other species have been

described or reported from the region. These include Lophogorgia luetkeni

Wright & Studer, 1889 (in Thomson, 1917), Leptogorgia africana Thomson,

1917, Leptogorgia rigida Verrill, 1868-1870 (in Thomson, 1917), Leptogorgia

tenuissima Kikenthal, 1919, Leptogorgia pusilla Kikenthal, 1919, and Lepto-

gorgia abietina Kikenthal, 1919. Of these, the only material currently available

for examination is Thomson’s specimen identified as Lophogorgia luetkeni (from

False Bay), and his types of Leptogorgia africana (East London) and Lepto-

gorgia rigida (border of eastern Cape and Transkei). The three specimens are

dried and badly fragmented; thus positive identification of the material is very

difficult. However, an examination of branching pattern and sclerites shows

Leptogorgia africana to be similar to Leptogorgia gilchristi, whereas Leptogorgia

rigida and Lophogorgia luetkeni resemble Leptogorgia palma.

Several specimens present in the South African Museum collection (both

older ones as well as recently collected material) cannot with certainty be allo-

cated to a particular species. The full extent of morphological variability present

within the four presently recognized southern African species of Leptogorgia has

not been fully assessed but a considerable amount of intraspecific variation is

apparent, particularly in Leptogorgia flammea and L. gilchristi.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 251

Family Ellisellidae Gray, 1859

Axis composed of calcified concentric layers; sclerites and horn not present;

central core not chambered. Colonies unbranched and whip-like, or branching

dichotomous, pinnate or lyrate. Polyps monomorphic. Sclerites are primarily

double heads, double cones, double stars and clubs.

About seven genera of the Atlantic and Indo-Pacific with at least two

genera in southern Africa. The family is in need of revision.

Genus Junceella Valenciennes, 1855

Junceella Valenciennes, 1855: 14. Kiikenthal, 1919: 857; 1924: 361 (Lit!).

Diagnosis

Colonies are flagelliform or sparsely dichotomously branched. Sclerites of

the outer coenenchyme are distinct clubs, somewhat flattened, with rounded

tubercles covering the head, and a whorl of tubercles surrounding the handle.

At least seven species of the Indo-Pacific.

Type species. Gorgonia juncea Pallas, 1766; Philippines.

Remarks

A single colony assignable to this genus has recently been collected from

Sodwana Bay in northern Natal, at 37 m depth (Williams 1989: 142, in press c).

The specimen has not been identified to species. It is orange-red in colour.

This represents a new record for the genus Junceella in southern Africa.

Other species of the family Ellisellidae

Thomson (1917: 43, pl. 5 (fig. 6)) identified material from Natal (113-

146 m depth) as Verrucella bicolor Nutting, 1908, originally described from

Hawaii. Two specimens in the South African Museum collection are identified

as Scirpearia furcata and S. flagellum (Scirpearia is a synonym of Ellisella

(Bayer, 1981: 945). At least three other ellisellid species (excluding Junceella

sp.) have recently been collected from southern African waters (including

material conforming to Thomson’s description as well as the Scirpearia

material). Because of the confused state of the taxonomy of the family Elliselli-

dae, none of this material can at present be reliably assigned to genus or species.

Proper identification of southern African ellisellids requires extensive revisions

of most of the recognized genera involving a large collection of specimens (Dr

F. M. Bayer pers. comm.). The family Ellisellidae is presently known in south-

ern Africa from East London, eastern Cape Province, to Sodwana Bay,

northern Natal (37-567 m in depth).

Family Chrysogorgiidae Verrill, 1883

Axis composed of entirely calcified material in concentric lamellae. Surface

smooth, glossy with a metaliic or iridescent lustre. Polyps monomorphic. Scle-

Dy) ANNALS OF THE SOUTH AFRICAN MUSEUM

rites are rods, scales, or plates, sometimes needle-like; others flat and broad.

Holdfast is root-like (forms living in soft substrata) or disc-like (forms attached

to hard objects).

Twelve genera of cosmopolitan distribution (many from great depth). At

least five genera in southern Africa.

Genus Chrysogorgia Duchassaing & Michelotti, 1864

Chrysogorgia Duchassaing & Michelotti, 1864: 13, 21. Wright & Studer, 1889: 23. Versluys,

1902: 17. Kiikenthal, 1919: 505; 1924: 388 (Lit!). Bayer, 1956: F216. Bayer & Stefani,

OSS 259)

Diagnosis

Colonies are erect and often bottlebrush-like in appearance. Branching is

sympodial. The main stem has a zigzag appearance with the lateral branches

arranged in a spiral fashion. Lateral branches relatively short, dichotomously

branched. Polyps sparsely arranged on the lateral branches.

Sclerites are scales and prickly spindles. Main stem often bronze or copper-

coloured with a lustrous iridescence, lateral branches and polyps whitish or

yellowish.

A large genus of probably more than 50 species, mostly from deep water of

the Atlantic and Indo-Pacific oceans.

Type species. Chrysogorgia desbonni Duchassaing & Michelotti, 1864;

Lesser Antilles.

Remarks

An unidentified species of Chrysogorgia occurs from off Durban to north-

ern Natal (722-1 200 m in depth).

The present work represents a new record for the genus in southern Africa.

Genus Radicipes Stearns, 1883

Radicipes Stearns, 1883: 97. Kiikenthal, 1919: 540; 1924: 410 (Lit!). Bayer, 1956: F216.

Lepidogorgia Verrill, 1884: 220. Versluys, 1902: 5.

Strophogorgia Wright & Studer, 1889: 2.

Diagnosis

Colonies are very thin and whip-like, unbranched, and arise from a root-

like holdfast. The roots ramify dichotomously. Polyps are arranged singly,

sparsely and uniserially along the axis. The coenenchyme is often very thin.

Sclerites are irregularly-shaped scales or spindles with smooth or slightly rough-

ened surfaces. Colony colour usually whitish.

At least five species of the Atlantic and Indo-Pacific, usually from deep

water.

Type species. Radicipes pleurocristatus Stearns, 1883; western Pacific.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 253

Remarks

An unidentified species of Radicipes has recently been collected near Cape

Vidal, northern Natal, at 600—810 m in depth. This represents a new record for

the genus in southern Africa.

Genus Simpsonella Stiasny, 1940

Hicksonella Simpson, 1910: 682; preoccupied by Hicksonella Nutting, 1910.

Simpsonella Stiasny, 1940: 31.

Helicogorgia Bayer, 1981: 902, 938.

Diagnosis

Colonies usually unbranched and whip-like, or rarely with sparse lateral

branching. Polyps uniserial to muitiserial, longitudinally placed on one side of

stem, leaving a naked track on the opposite side. Polyps clavate, non-retractile.

Fig. 50. Simpsonella capensis (Simpson, 1910). A. An entire colony; scale bar= 10 mm.

B. Surface detail from middle of colony showing the side containing the polyps; length of figure

9,0 mm. C. Surface detail from middle of colony showing the side containing the bare tract;

length of figure 9,2 mm.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anthocodiae retractile into body of polyps. Sclerites are mainly tuberculated |

spindles and plates, often of irregular shape.

A genus of at least four species, eastern African coast from Zanzibar to

East London. There is some doubt as to the familial status of this genus

(Dr F. M. Bayer pers. comm.); the sclerites are unlike those of other chryso-

gorgiids.

Fig. 51. Simpsonella capensis (Simpson, 1910). A. Sclerites from the calyx wall. B. Coenen-

chymal sclerites. Scale bar = 0,2 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA EDS

Type species. Juncella spiralis Hickson, 1904; South Africa.

Simpsonella capensis (Simpson, 1910)

Figs 50-52

Hicksonella capensis Simpson, 1910: 685, 688, pl. 13 (figs 6-7).

Simpsonella capensis Williams, in press b.

Material

SAM-—H3932, off Gonubie, eastern Cape Province (33°04,7’S 28°07,2’E),

90 m, 17 July 1984, 1 colony, coll. G. C. Williams, R.V. Meiring Naude.

Description

The colony examined is 140 mm in length. The colony is flagelliform,

unbranched, and stands upright but curved with a spiral twist toward the distal

end. The axis is c. 1,5 mm in width. The maximum diameter of the colony,

including the polyps, is 2,5 mm. The polyps are arranged multiserially along a

longitudinal tract that covers approximately three-quarters of the circumference

of the axis. The polyps are elongate and narrow and thus appear less crowded

relative to other species. The polyps are slightly incurved and somewhat clavate

1,5 mm long, 0,32 mm wide proximally, and 0,46 mm wide distally. Sclerites of

the polyps (0,09—0,25 mm long) are irregular plates with scalloped margins and

rounded tubercles that are covered by fine punctations. Coenenchymal sclerites

are elongate and mostly tuberculated spindles 0,06-0,20 mm in length. Colony

colour cream-white.

Distribution

East London region to southern Natal (67-90 m in depth). Type locality

southern Natal (red cliff near Morewood Cave).

Remarks

This species is distinguished by the possession of elongate and narrow

polyps that are relatively openly spaced on the longitudinal polypiferous tract,

and irregularly shaped, coarsely tuberculated spindles of the coenenchyme

(0,06—0,20 mm iong). (See remarks under Simpsonella spiralis (type species)

regarding the generic designation adopted for this species.)

Simpsonella flagellata (Simpson, 1910)

Figs 53-55

Hicksonella flagellata Simpson, 1910: 684, 688, pl. 13 (figs 44-5).

Simpsonella flagellata Williams, in press b.

Material

SAM-—H3929, off Port Edward, northern Transkei (31°06’S, 30°18’E),

125 m, 8 July 1985, 20 colonies, dredge, G. C. Williams, R.V. Meiring Naude.

ANNALS OF THE SOUTH AFRICAN MUSEUM

, 1910); sclerites.

H. 0,075 mm.

19 mm. D. 0,125 mm.

impson

11 mm

Cc:

impsonella capensis

B. 0,11 mm.

F. 0,15 mm. G

hs. §

. 0,15 mm.

175 mm.

icrograp

lectron m

tes from the polyp walls

tes

ing e

Scann

Coenenchymal scler

Fig. 52

A-D. Scler

E-H.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 257

—<—— a

Fig. 53. Simpsonella flagellata (Simpson, 1910). A. An entire colony, 300 mm in length.

B. Surface detail from middle of colony; total length of figure 16 mm. C. Surface detail from

middle of colony; total length of figure 6 mm.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 54. Simpsonella flagellata (Simpson, 1910). A. Sclerites from the calyx wall. B. Coenen-

chymal sclerites. Scale bar = 0,1 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 259

Fig. 55. Scanning electron micrographs. Simpsonella flagellata (Simpson, 1910); sclerites.

A-D. Sclerites from the polyp walls. A. 0,08 mm. B.0,11 mm. C. 0,08 mm. D. 0,21 mm.

E-H. Coenenchymal sclerites. E. 0,062 mm. F. 0,055 mm. G. 0,075 mm. H. 0,15 mm.

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-—H3930, off Nthlonyane River mouth, Transkei (32°17’S 29°06’E), 300 m,

5 July 1985, 7 colonies, dredge, G. C. Williams, R.V. Meiring Naude.

Description

Colonies examined range in length from 60 mm to 500 mm. Colonies are

flagelliform, unbranched, curved or ascending in a loose open spiral. Axis

0,5-—2,0 mm wide. Coenenchyme very thin. Polyps crowded and multiserial, in a

longitudinal band covering about one half of the circumference of the axis. A

naked tract of coenenchyme is present on the opposite side. Polyps face upward

and curve strongly inward, strongly clavate, mostly 2,0-2,5 mm long and

0,7-1,5 mm wide. Polypiferous tract present almost throughout entire length of

colony, only the proximal-most 10-30 mm is polyp free. Holdfast discoid to

deltoid, 5-10 mm in diameter. Width of polypiferous portion of colony usually

3-4 mm. Polyp sclerites (0,04-0,2 mm long) are irregularly-shaped, tuber-

culated plates with mostly scalloped margins; also present are some double discs

or unilaterally foliate forms. Coenenchymal sclerites (0,1—-0,25 mm long) are

robust irregular spindles with large pustule-like tubercles; also present are many

double discs and unilaterally foliate forms. Colony colour white to cream-white

with bare tract greyish-white. Axis is graphite coloured with a lustrous sheen.

Distribution

Transkei coast, 125-300 m in depth. Type locality Cape Morgan (border of

Cape Province and Transkei).

Remarks

This species is distinguished from other members of the genus by the pos-

session of double discs and other unilaterally foliate sclerites (0,04—0,06 mm in

length). (See remarks under Simpsonella spiralis (type species) regarding the

generic designation adopted for this species.) :

Simpsonella spiralis (Hickson, 1904)

Figs 56-58

Juncella spiralis Hickson, 1904: 231, pl. 8 (figs 6-9).

Junceella spiralis: Stiasny, 1940: 31.

Hicksonella spiralis Simpson, 1910: 682, pl. 13 (figs 1, 3). Thomson, 1911: 889. Toeplitz, 1929:

241, 273. Hickson, 1938: 607.

Simpsonella spiralis Stiasny, 1940: 31. Williams, in press b.

Helicogorgia spiralis Bayer, 1981: 938.

Material

SAM-H1245 (syntype) of Cape Morgan, southern Transkei (32°45’45"S

28°26'15"E), 66 m, 12 January 1899, 4 partial colonies, dredge S.S. Pieter Faure

survey, PF 858. SAM—H3931, off Mbashe River mouth, Transkei (32°21'S

29°00'E), 100 m, 5 July 1985, 10 colonies both partial and whole, dredge, G. C.

Williams, R.V. Meiring Naude.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 261

Description

Colonies examined range in length from 260 mm to 370 mm. The colonies

are flagelliform and form relatively tight spirals, resembling curved corkscrews.

Axis is 1,0—1,5 mm wide. Coenenchyme thin. Polyps are crowded and arranged

multiserially in a longitudinal band covering approximately three-quarters of the

circumference of the axis. A narrow naked tract of coenenchyme is present

opposite the polyp band. Polyps 1,0—1,5 mm long and 0,3—0,5 mm wide, clavate,

facing upward and slightly incurved. Polyp sclerites (0,10—0,17 mm long) are

mostly broad and irregularly-shaped plates, with large rounded tubercles or fine

punctation only. Coenenchymal sclerites are robust ovoid or elongate spindles

with large pustule-like tubercles (0,07—0,28 mm long). Colour of polyps mostly

cream-white; bare tract is orange-red; axis graphite coloured and glossy.

Fig. 56. Simpsonella spiralis (Hickson, 1904). A. An entire colony, 150 mm in length.

B. Surface detail from middle of colony; total length of figure 11 mm.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 57. Simpsonella spiralis (Hickson, 1904). A. Sclerites from calyx wall. B. Coenenchymal

sclerites. Scale bar =0,15 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 263

Fig. 58. Scanning electron micrographs. Simpsonella spiralis (Hickson, 1904); sclerites.

A-D. Sclerites from the polyp wall. A.0,13 mm. B.0,09 mm. C. 0,085 mm. D. 0,14 mm.

E-H. Coenenchymal sclerites. E. 0,09 mm. F. 0,075 mm. G. 0,07 mm. H. 0,125 mm.

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Southern Transkei coast, 66-100 m in depth. Type locality Cape Morgan

(border of Cape Province and Transkei).

Remarks

Simpsonella spiralis is differentiated from other members of the genus by

the tightly spiralled, corkscrew-like growth form and possession of broad ovoid

to elongate spindles (0,07—0,28 mm long) in the coenenchyme.

Simpson (Dec. 1910: 682) considered Hickson’s allocation of this species to

Junceella (Juncella of Hickson is a misspelling) to be incorrect and proposed the

generic name Hicksonella. Stiasny (1940: 31) found this name preoccupied by

Hicksonella Nutting (May 1910), a genus in the family Gorgoniidae, and pro-

posed the name Simpsonella instead. Bayer (1981: 938), having not seen

Simpson’s proposal, assigned the generic name Helicogorgia for the same. Simp-

sonella is therefore the valid and accepted generic designation.

Simpsonella squamifera (Kikenthal, 1919)

Figs 59-61

Radicipes squamiferus Kikenthal, 1919: 545, figs 240-243; 1924: 412.

Simpsonella squamifera Williams, in press b.

Material

SAM-—H3886, off Port Edward, northern Transkei (31°05,8’S 30°18,8’E),

140 m, 8 July 1985, 2 whole and 3 partial colonies, dredge, coll. G. C. Williams,

R.V. Meiring Naude. SAM—H3888, off Qolora River mouth, Transkei

(32°45,8'S 28°36,4’E), 240-150 m, 14 July 1984, 2 whole colonies, dredge, coll.

G. C. Williams, R.V. Meiring Naude. SAM—H3933, off Gonubie, eastern Cape

Province (32°55,0'S 28°31,0’E), 630 m, 25 May 1978, 6 colonies, heavy dredge,

South African Museum, R.V. Meiring Naude.

Description

Colonies examined range between 65 mm and 330 mm in length. Colonies

are flagelliform, very thin, almost hair-like; upright, and are curved or form a

loose open spiral. Axis 0,3-1,5 mm in diameter, arising from a disc-like

holdfast, 4-5 mm in diameter. Polyps arranged uniserially along a sinuous

longitudinal tract along the axis. Some colonies have polyps more crowded,

appearing biserial in places. Polyps usually present in distal half of colony only,

0,6-0,8 mm long by 0,16-0,30 mm wide; short, robust, somewhat clavate,

facing upward and incurved. Polyp sclerites are irregular plates with scalloped

margins and tuberculated surfaces (0,10-—0,22 mm long). Coenenchymal sclerites

are elongated coarsely tuberculated spindles that may be bent or curved

(0,12—0,33 mm long). Colony colour cream-white or reddish-orange.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 265

Distribution

East London region to Durban, Natal, in depths 90-775 m (Williams in

press a), and also Tanzania (type locality).

Remarks

This species was originally described by Kikenthal (1919: 545) from the

region of Dar es Salaam and Zanzibar (Tanzania), 463 m in depth.

Fig. 59. Simpsonella squamifera (Kiikenthal, 1919). A. An entire colony, 240 mm in length.

B. Surface detail from the middle of one colony; total length of figure 3 mm. C. Surface detail

from middle of another colony; total length of figure 4 mm.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 60. Simpsonella squamifera (Kikenthal, 1919). A. Sclerites from calyx wall. B. Coenen-

chymal sclerites. Scale bar = 0,15 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 267

Fig. 61. Scanning electron micrographs. Simpsonella squamifera (Kikenthal, 1919); sclerites.

A-C. Sclerites from the polyp wall. A. 0,22 mm. B. 0,12 mm. C. 0,175 mm. D-F. Coenen-

chymal sclerites. D.0,10 mm. E. 0,35 mm. F. 0,20 mm. G. Detail of a polyp wall sclerite;

length of portion shown 0,06 mm. H. Detail of a coenenchyme sclerite; length of portion

shown 0,04 mm.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is distinguished from other Simpsonella species by the uniserial or biserial

arrangement of the polyps and the possession of large elongate coarsely

tuberculate spindles in the coenenchyme (0,12—0,33 mm in length).

Kikenthal (1919: 545) allocated this species to the genus Radicipes Stearns,

1883. However, species of Radicipes are distinguished by polyp sclerites that are

minutely sculptured rods or scales with smooth margins and without large

tubercles, and root-like projections of the holdfast that are dichotomously

branched. Simpsonella squamifera, on the other hand, possesses scalloped plates

and coarsely tuberculated spindles, and small disc-like holdfasts; both these

features are characteristic of the genus Simpsonella.

This species shows some variation regarding both colour and arrangement

of polyps. Colony colour is usually cream-white but some may be at least

partially rust-orange to reddish-brown. The longitudinal tract that contains the

polyps forms a sinuous pattern along one side of the axis. The polyps may be

sparsely arranged in a uniserial fashion along this tract or they may be more

crowded with adjacent polyps in close proximity, appearing biserial in some

places.

Genus Trichogorgia Hickson, 1904

Trichogorgia Hickson, 1904: 222.

Malacogorgia Hickson, 1904: 226.

Diagnosis

Colonies lyrate, flabellate, and planar, with secondary branching dicho-

tomous. Length of terminal branches often one third to one half of the total

colony length. Sclerites when present are ovoid scales.

A genus of four species from the western Atlantic, Caribbean Sea, and

South Africa.

Type species. Trichogorgia flexilis Hickson, 1904; South Africa.

Trichogorgia flexilis Hickson, 1904

Figs 62, 71A—B

Trichogorgia Flexilis Hickson, 1904: 222, pl. 8 (fig 13), pl. 9 (figs 16-17).

Trichogorgia flexilis: Stiasny, 1940: 36, text-fig. I.

Material

SAM-H581 (syntype), off Cape Recife (34°07'S 25°43’E), 102 m, 1 dried

colony. SAM—H1047, off Cape Vidal (28°10’S 32°45’E), 146-183 m, 27 Feb-

ruary 1901, dredge, 2 colonies, S.S. Pieter Faure survey, PF 11967. SAM—

H3704, off East London (33°10,3’S 20°06,2’E), 100 m, 17 July 1984, 2 colonies,

dredge, G. C. Williams, R.V. Meiring Naude. SAM—H3935, Mbashe River

mouth, Transkei (32°20,0’S 29°00,9’E), 110 m, 5 July 1985, 1 colony, dredge,

G. C. Williams, R.V. Meiring Naude.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 269

Fig. 62. Trichogorgia flexilis Hickson, 1904. A. An entire colony, 90 mm in height.

B. A single polyp 0,7 mm in length. C. Detail of a terminal branch 3,5 mm long.

D. Polyp sclerites. Scale bar =0,1 mm.

Description

The colonies examined range in length from 70 mm to 90 mm. Colonies are

planar, branching is dichotomous, relatively sparse from a single basal stem.

Polyps are densely arranged in two rows along each branch and are placed

opposite or alternate to one another. The contracted polyps are relatively robust

and somewhat clavate, and rarely exceed 1 mm in length. Sclerites of the

coenenchyme and surface of the polyps are scales that vary in length from

0,04mm to 0,17 mm. Many are somewhat bone-shaped with very fine

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

denticulations along the margin. Colony colour greyish-white, or light yellowish-

brown.

Distribution

Cape Recife near Port Elizabeth to Cape Vidal, northern Natal, 24-183 m

in depth (Williams in press a). Type locality is Cape Recife (south coast of South

Africa).

Remarks

Trichogorgia capensis (Hickson, 1904: 226) (= Malacogorgia capensis), was

described from near the type locality of T. flexilis at Algoa Bay (36 m depth). Its

distribution extends from False Bay to Algoa Bay (Utinomi 1964: 11). I have

examined Hickson’s holotype (SAM-—H651), which is dried and damaged. It

appears identical to T. flexilis, except that it completely lacks sclerites. Thus,

two species of Trichogorgia are known to occur in southern Africa, T. flexilis

with sclerites, and JT. capensis without sclerites. Utinomi (1964: 9) recorded

T. capensis as Malacogorgia capensis from Antarctica (Gunnerus Bank and

Cape Cook at 500-590 m in depth).

Genus Xenogorgia Bayer & Muzik, 1976

Xenogorgia Bayer & Muzik, 1976: 85.

Diagnosis

Colonies bottlebrush-like in appearance with lateral branches arising

irregularly from the main axis. The polyps are cylindrical and are situated on all

sides of the branches. Sclerites are thin, mostly oval scales.

One species of the genus is presently known (from the west coast of South

Africa in deep water).

Type species. Xenogorgia sciurus Bayer & Muzik, 1976; west coast of South

Africa.

Remarks

Xenogorgia sciurus was described from off Lambert’s Bay on the west coast

of South Africa (680-800 m in depth). The species has apparently not been

encountered since the type material was originally collected. See Bayer & Muzik

(1976: 85-89, figs 8-10) for a detailed description of the type material.

Family Primnoidae Gray, 1857

Axis entirely calcified, often longitudinally grooved on surface. Polyps

monomorphic, armed with large plates or scales. Retracted anthocodiae

protected by an operculum composed of eight deltoid scales. Branches covered

by a layer of scales or plates.

Twenty-four genera of cosmopolitan distribution.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 271

Genus Callogorgia Gray, 1858

Callogorgia Gray, 1858: 286. Bayer, 1982: 119 (Lit!).

Xiphocella Gray, 1870: 56.

Caligorgia: Wright & Studer, 1889: 75. Kiikenthal, 1919: 362; 1924: 267.

Diagnosis

Colonies with planar and pinnate branching, branchlets mostly alternate,

only rarely opposite. Polyps face upward, in whorls of 3—6. Each polyp is curved

inward toward the axis. Polyp sclerites are curved scales. Coenenchymal

sclerites are ovate, polygonal or elongate plates.

Approximately 20 species of the Atlantic, Mediterranean, and Indo-Pacific.

Type species. Gorgonia verticillata Pallas, 1766; eastern Atlantic and

Mediterranean Sea.

Remarks

Several colonies of a presently undetermined species of Callogorgia have

recently been collected from the southern Transkei (Indian Ocean coast of

southern Africa) between 200 m and 250 m in depth.

The present work represents a new record of the genus Callogorgia to

southern Africa.

Bayer (1982: 116) distinguished two other genera that are similar to

Callogorgia. These are Fanellia Gray, 1870 (eight species from the Pacific

Ocean; see Bayer & Stefani 1988: 471 for a key to the species of Fanellia), and

Ascolepis Thomson & Rennet, 1931, with perhaps four species in the Southern

oceans.

Genus Calyptrophora Gray, 1866

Calyptrophora Gray, 1866: 25. Wright & Studer, 1889: 50. Kiikenthal, 1919: 468; 1924: 317

(Lit!). Bayer, 1956: F221.

Diagnosis

Colonies with branching that is mostly planar and loosely dichotomous,

although sometimes unilaterally pinnate and fan-shaped. Polyps are in whorls

and face upward. Each polyp is surrounded by a ring composed of the fusion of

two pairs of body scales.

A genus of approximately six species of the Indo-Pacific and Atlantic

oceans, in deep water.

Type species. Calyptrophora japonica Gray, 1866, Japan.

Remarks

Material representing a unidentified species Calyptrophora has recently

been collected from northern Natal (Indian Ocean coast of southern Africa)

between 722 m and 1 200 m in depth.

The present work represents a new record for the genus Calytrophora to

southern Africa.

DED, ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Narella Gray, 1870

Narella Gray, 1870: 49.

Calypterinus Wright & Studer, 1889: 53.

Stachyodes Wright & Studer, 1889: 54.

Diagnosis

Colonies planar, branching often lyrate, pinnate, or dichotomous. Polyps

face downwards and are arranged in whorls. Each polyp is conspicuously armed

with 3—4 pairs of large abaxial body scales. Perhaps 20 species of the Atlantic,

Indo-Pacific, and Antarctic.

Type species. Primnoa regularis Duchassaing & Michelotti, 1860; West

Indies.

Narella gilchristi (Thomson, 1911) comb. nov.

Figs 1G, 63-65

Stachyodes gilchristi Thomson, 1911: 885, pl. 44 (fig. 1), pl. 45 (figs 2a, 2b). Stiasny, 1940: 34,

text-fig. H, pl. 1 (fig. 8).

Stachyodes gilberti (?misspelling of S. gilchristi) Kikenthal, 1919: 468; 1924: 316.

Stachyodes capensis Thomson, 1917: 25, text-fig. 5, pl. 3.

Material

SAM-H985 (holotype), off Cape Vidal, northern Natal (28°10’S 32°40’E),

146-183 m, 27 February 1901, 1 fragmented colony, dredge, S.S. Pieter Faure

survey, PF 11966. SAM-—H3657, Park Rynie, Natal (30°20,7’S 20°51,6’E),

105 m, 1° July 1984, 3 whole colonies, dredge, G. C. Williams, R.V. Meiring

Naude. SAM-H1108 (holotype of Stachyodes capensis), off Cape Morgan

(border Cape Province and Transkei), 141m, 26 July 1901, 1 dried and

fragmented colony, dredge, S.S. Pieter Faure survey, PF 13159. SAM—H3656,

off Ngabara Point, Transkei (32°25,0’S 28°58,3’E), 330-340 m, 13 July 1984,

2 partial colonies, dredge, G. C. Williams, R.V. Meiring Naude. SAM-—H3660,

off East London (33°10,3’S 28°06,2'E), 100 m, 17 July 1984, 1 whole colony,

dredge, G. C. Williams, R.V. Meiring Naude.

Description

The colonies examined are 90-140 mm in height and 70-150 mm in width.

Branching is lyrate (branching in which the secondary branches are straight,

upright, and parallel) and planar. Colonies may have as many as 20 upright and

parallel secondary branches. Some of these may be dichotomously branched.

Polyps appear in whorls of 4—8 polyps per whorl. Whorls are congested along all

branches, and vary from 4 mm to 9 mm in diameter. Polyps 2-3 mm in height

and 1,0-—1,5 mm in width, composed of 3 (rarely 4) pairs of abaxial scales. Basal

and medial scales rounded, pointed, or somewhat serrated terminally. Buccal

scales usually rounded, more-or-less smooth terminally. Coenenchyme of

branches covered with a layer of scales, exhibiting a cobblestone-like appear-

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA DAS

ASS &

ERY

ie ey

<i i ae

ay SO Sih oY

SIS LD AY (X

RG fx KY vA

INES BD SI a oS

oS GAR BS £

FINS SY BY $

ny OE :

my & mS aii

Wy NY At a

Ms ie aS 3 i ‘)

Ne Eh i Ry Is

kay 2N BM Say Fey

aN a3 ae A Ley a i ,

AS 9 GR ae MES

MS OR "a se

owt Bote

Oe sp

Fig. 63. Narella gilchristi (Thomson, 1911). A. An entire colony, 150 mm in height. B. Detail

of a group of three polyps; scale bar=1,0 mm. Abbreviations: ba—basal scale, bu—buccal

scale, m—medial scale, o—opercular scale.

ance. Colony scales 0,65—2,0 mm in length. Colony colour pink in life, fading to

white in alcohol.

Distribution

East London to Cape Vidal, northern Natal, 90-340 m in depth (Williams

in press a). Type locality is Cape Vidal.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 64. Scanning electron micrographs. Narella gilchristi (Thomson, 1911); polyps. A. Group

of three polyps showing basal, buccal, medial, and opercular scales; total length of micrograph

4,0 mm. B. Two polyps showing basal, medial, and buccal scales; length of polyps 1,9 mm.

C. A single polyp enlarged from B, 1,9 mm in length. D. Detail of a polyp showing buccal and

opercular scales; total length of micrograph 1,2 mm. E. A whorl of polyps; polyp on left has

two pairs of medial scales; total length of micrograph 3,6 mm. F. A single polyp, 2,4 mm long.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 2715

Fig. 65. Scanning electron micrographs. Narella gilchristi (Thomson, 1911). A. Group of

three polyps, each 1,2 mm in length. B. Detail of polyp showing terminal margins of medial

scales. C-H. Scales from polyps and surface of colony. C.1,25mm. D.0,8 mm.

EE 0.95mme Ee mm, (Ge? mm. Be 12 mm:

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Bayer (1956: 222) considered Stachyodes Studer, 1887, a synonym of

Narella Gray, 1870. I have compared the holotypes of Narella gilchristi and

Stachyodes capensis Thomson, 1917 and can find no substantial differences to

justify maintaining them as two species. Variability is evident in the orna-

mentation of the terminal margin of the polyp body scales. Some polyps have

rounded scales with margins lacking spines; others have prominent deltoid

serrations or spines. The more rounded terminal margins of the polyp scales in

S. capensis can certainly be explained by intraspecific variation. I therefore

consider S. capensis a junior synonym of N. gilchristi.

Genus Primnoeides Wright & Studer, 1889

Primnoeides Wright & Studer, 1889: 90.

Diagnosis

Colonies are feather-like with planar and pinnate branching. The lateral

branches are arranged in pairs, originating opposite one another. The polyps

face upwards and are arranged in pairs along the lateral branches. Sclerites of

the polyps and coenenchyme are thin and flat scales.

Perhaps two species known from the south-western Indian Ocean and

adjacent Subantarctic.

Type species. Primnoeides_ sertularoides Wright & Studer, 1889;

Subantarctic.

Remarks

Several colonies representing a currently unidentified species of the genus

Primnoeides have recently been collected from the east coast of southern Africa

(southern Transkei to northern Natal), 400—450 m in depth.

The present work represents a new record for the genus Primnoeides to

southern Africa.

Genus Primnoella Gray, 1858

Primnoella Gray, 1858: 286. Kiikenthal, 1919: 384; 1924: 279 (Lit!). Bayer, 1956: F220.

Diagnosis

Colonies mostly unbranched and whip-like or rarely sparsely dichotomously

branched. Polyps are in whorls and are closely appressed to the axis. The polyps

face upwards. The opercular scales are inconspicuous and the more conspicuous

marginal scales often fold over them.

Approximately fifteen species of the Southern Hemisphere: Southern

Ocean, Atlantic, and Indo-Pacific. )

Type species. Primnoa australasiae Gray, 1849, Australia, Tasmania, and

New Zealand.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA DAG.

Remarks

Material representing a presently unidentified species of the genus Prim-

noella has recently been collected from the east coast of southern Africa off the

southern Transkei, 710-775 m in depth.

The present work represents a new record of the genus Primnoella for

southern Africa.

Genus Thouarella Gray, 1870

Thouarella Gray, 1870: 45. Kikenthal, 1919: 405; 1924: 287 (Lit!). Bayer, 1956: F220.

Amphilaphis Wright & Studer, 1889: 70.

Euthouarella Kikenthal, 1915: 149.

Diagnosis

Colonies often resemble bottlebrushes. Branching pinnate and multiplanar,

mostly profuse. Branches short, and arise from a single main stem. Large scales

present on all sides of a polyp. Eight marginal scales in two rings of four that

alternate below the opercular scales. Marginal scales fold inward over the

smaller opercular scales. A genus of approximately thirty species inhabiting the

Subantarctic, southern Atlantic and southern Indo-Pacific.

Type species. Primnoa antarctica Valenciennes, 1846; Southern oceans.

Thouarella hicksoni Thomson, 1911

Figs 1H, 66-68

Thouarella hicksoni Thomson, 1911: 886, pl. 44 (figs 3a, 3b), pl. 45 (fig. 1). Kikenthal,

1919: 439; 1924: 301. Stiasny, 1940: 32, text-fig. G, pl. 4 (fig. 21).

Thouarella Hicksoni: Tixier-Durivault, 1954: 625.

Material

SAM-—H1276, off Lion’s Head, Cape of Good Hope Peninsula (33°57’'S

18°10’E), 110 m, 22 March 1900, shrimp trawl, 3 colonies, S.S. Pieter Faure

survey, PF 2256. SAM-—H3636, off Stony Point, Transkei (32°30,9’S 28°45,0’E),

360 m, 12 July 1984, many colonies, dredge, G. C. Williams, R.V. Meiring

Naude.

Description

Colonies examined range in length from 70 mm to 230 mm. Branching is in

a bottlebrush pattern, pinnate, multiplanar. Branches arise approximately per-

pendicular to main stem. Branching profuse along a single main stem. Some

secondary branches may also be branched. Polyps are crowded on branches,

arranged on all sides of a branch but not in whorls. Polyps are 1,0—1,5 mm in

length. The eight marginal scales that subtend the opercular scales are broad

and somewhat circular with a deltoid and blunt terminal spine, which may be

somewhat serrated laterally. Other polyp scales are deltoid tc somewhat circular

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 66. Thouarella hicksoni Thomson, 1911. A. An entire colony, 160 mm in height.

B. Detail of a terminal branch showing ten polyps; total length of figure 6 mm.

in shape, and with coarse tuberculation proximally. Coenenchymal scales are

thin and irregularly ovoid. Colony colour pale yellowish to cream-white.

Distribution

Namaqualand coast (South African west coast) to the coast of the Transkei,

100-860 m in depth (Williams in press a). Type locality is Cape St Francis (Cape

south coast).

Family Isididae Lamouroux, 1812

Axis composed of nodes of horn and internodes of non-spicular calcium

carbonate. Holdfasts are root-like for soft substrata or disc-like for hard

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 279

Fig. 67. Scanning electron micrographs. Thouarella hicksoni Thomson, 1911; polyp body

scales. A. 0,25 mm. B. 0,40 mm. C. 9,43 mm. D. 0,24 mm. E. 0,45 mm. F. 0,20 mm.

substrata. Branching arises from nodes or internodes. Polyps monomorphic.

Sclerites of spindles, needles, rods, plates or scales.

Twenty genera in six subfamilies, cosmopolitan distribution.

Subfamily Keratoisidinae Gray, 1870

Polyps non-retractile. Sclerites of the polyps include needles, rods or

spindles.

At least four genera.

: Se

CG Aw

“

ANNALS OF THE SOUTH AFRICAN MUSEUM

280

< Se

WES

ion of micrograph 0,9 mm.

length. F. A single polyp, 1,2 mm long.

imens

A, total length of micrograph 0,56 mm.

linear d

imum

Thouarella hicksoni Thomson, 1911. A. A single

Detail of polyp

le polyp, max

five polyps, 1,2 mm

ing

ing electron micrographs.

length.

fans

| branch show

ion oO

Scann

1,3 mm

ddle port

. Termina

C. Mi

Fig. 68.

polyp,

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 281

Genus Keratoisis Wright, 1869

Keratoisis Wright, 1869: 23. Bayer, 1956: F222.

Ceratoisis: Verrill, 1883: 20 (unjustified emendation). Wright & Studer, 1889: 26. Kiikenthal,

1919: 585; 1924: 423 (Lit!).

Diagnosis

Colonies with branches arising from the internodes. Colonies large and

robust. Polyps with longitudinally placed sclerites not arranged en chevron.

Terminal ends of polyps with spine-like projections of spindles or needles

between the tentacular bases.

A genus of perhaps twenty or more species of circumtropical distribution in

deep water.

Type species. Keratoisis grayi Wright, 1869; north-eastern Atlantic.

Remarks

An undetermined species of the genus Keratoisis occurs off the Natal coast,

600-810 m in depth. This represents a new record for the genus Keratoisis in

southern Africa.

Genus Acanella Gray, 1870

Acanella Gray, 1870: 16. Nutting, 1910: 14. Kukenthal, 1919: 573; 1924: 418 (Lit!). Bayer,

9562 F222.

Diagnosis

Colonies with branches arising from the nodes. Colonies are bushy, with the

branches often arising in whorls. Spicules of the polyps are mainly spindles and

some flattened rods, with surfaces often finely denticulated.

A genus of ten or more species from the Atlantic and Indo-Pacific oceans.

Type species. Mopsea arbusculum Johnson, 1862; North Atlantic.

Remarks

An undetermined species of the genus Acanella is known from the Natal

coast, 549-1 200 m in depth. Thomson (1917: 19) identified this Acanella species

as A. eburnea (Pourtales, 1860), originally described from the northern Atlantic.

I consider this a dubious identification.

The present work represents a new record for the genus Acanella to

southern Africa.

Subfamily Mopseinae Gray, 1870

Polyps non-retractile, with an operculum. Sclerites of the polyps are

exclusively scales.

At least three genera.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Chathamisis Grant, 1976

Chathamisis Grant, 1976: 9, 10, 43. Bayer, 1981: 941.

Diagnosis

Colonies with branches arising from the internodes. Branching is bushy or

bottlebrush-like, multiplanar. Sclerites are scales with scalloped or serrated

margins and varying amounts of tuberculation.

A genus of at least nine species from the Southern oceans.

Type species. Chathamisis bayeri Grant, 1976; New Zealand.

Chathamisis ramosa (Hickson, 1904)

Figs 69-70, 71C—F

Ceratoisis ramosa Hickson, 1904: 224, pl. 7 (figs 3-4), pl. 8 (fig. 12). Thomson, 1911: 877,

pl. 43 (fig. 1). Stiasny, 1940: 35.

Primoisis ramosa Kikenthal, 1919: 616; 1924: 436.

Chathamisis ramosa Bayer & Stefani, 1987: 966.

Material

SAM-—H1054, off East London (33°10'S 27°50'E), 357 m, 23 April 1901,

1 colony, shrimp trawl, S.S. Pieter Faure survey, PF 12826. SAM-—H3665, off

Stony Point, Transkei (32°39’S 20°45’E), 360 m, 12 July 1984, 2 colonies, G. C.

Williams, R.V. Meiring Naude.

Description

Colonies range in length from 60 mm to 95 mm. Colonies are attached to

hard substrata with disc-like holdfasts. Colonies somewhat flabellate. Main stem

short (<10 mm long). Branching is bushy and multiplanar, but the colonies are

somewhat laterally compressed in one plane. Polyps scattered and present on all

sides of branches, usually arising perpendicular to axes of branches. Polyps are

1,0-1,2 mm long and 0,5 mm wide, clavate. Anthocodiae retractile into body of

polyp. Nodes are metallic gold or bronze in colour (0,3—1,5 mm in length).

Internodes white (3-7 mm in length). Polyp sclerites are irregular plates with

serrated or scalloped margins and some surface tuberculation (0,17—0,30 mm

long). Coenenchymal sclerites are similar but mostly narrow (0,08—0,30 mm in

length), some smaller ones being crescent shaped. Colony colour cream-white,

pale-yellowish, or pale-pinkish.

Distribution

Off the Cape of Good Hope Peninsula to northern Natal, 146-900 m in

depth (Williams in press a). Type locality is off Cape Point (south-western Cape

Province).

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 283

“e)

S o/Q

i ‘b

~) “A ,

Fig. 69. Chathamisis ramosa (Hickson, 1904). A. An entire colony, 70 mm in length.

B. Detail of the distal end of a branch showing six polyps and opercular sclerites; length of

figure 5,1mm. C.A single polyp showing placement of sclerites; total height of polyp

0,6 mm. D. A single opercular sclerite 0,2 mm in length.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 70. Chathamisis ramosa (Hickson, 1904). A. Sclerites from the calyx wall. B. Coenen-

chymal sclerites. Scale bar = 0,1 mm.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 285

Fig. 71. Scanning electron micrographs. A-B. Trichogorgia flexilis Hickson, 1904, sclerites

from the polyp wall. A.0,1 mm. B. 0,114 mm. C-F. Chathamisis ramosa (Hickson, 1904);

sclerites from the polyp wall and surface coenenchyme. C.0,12mm. D. 0,145 mm.

E.02i-mms £0519 mm:

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This species is differentiated from other southern African isidids by the

possession of disc-like holdfasts for attachment to hard substrata, profuse

branching with branches arising from internodes, and transversely arranged

scale-like or plate-like sclerites surrounding the polyps.

Morphological variability is evident in this species. Some colonies have a

bushy growth form and are multiplanar, others appear planar with dichotomous

branching but none-the-less possess short branches perpendicular to the plane of

the colony. In many of these near-planar colonies, the polyps are mainly

restricted to one face of the colonial plane. Sclerite ornamentation also varies.

Some colonies have scales that are relatively smooth with only a few scattered

tubercles. Others possess strongly tuberculated scales.

KEY TO THE GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

1A. Axis absent, colony composed of one or two outer layers (cortex) and

an inner layer (medulla). Cortex and medulla separated by a ring of

longitudinal canals. Medulla composed entirely of free but densely-set

SCIEHILES aoc ee Wes Hebe: olen ewe es Wace dao ee D,

1B. Axis present, composed of fused sclerites united by horn-like material

and/or calcium carbonate, or entirely proteinous and permeated by

varying amounts of non-spicular calcium carbonate, or totally calcified

without sclerites or-horm.....o......000.605.405. 0. ee +

2A. Colonies unbranched; almost digitiform or clavate to capitate (swollen or

widening distally), <50 mm in length (Fig. 7) ..... Diodogorgia capensis

2B. Colonies elongate and tapering distally; usually branched; mature colonies

>50 mm in length (Figs 1,5) ..6.. 40s .sc see hoe ee 3

3A. Colonies uniform brownish-tan or whitish. Calyces permanent and con-

spicuous, with eight external longitudinal grooves. Anthocodiae com-

MnOmlky joeeseinvec! @xseriec! TRY. Z)) oaccoosccsosee: Anthothela parviflora

3B. Colonies wine-red, orange with yellow calyces, or uniform pinkish-white

or yellow. Calyces present or absent, without eight external longitudinal

grooves. Anthocodiae commonly preserved totally retracted into calyces

On colomy (Pies 5). cag ey nee, SeM ccm Homophyton verrucosum

4A. Axis composed of sclerites that are united together by horn-like material

or non-spicular calcium carbonate or both (Figs 10-11)..................

Pre ere et rer ed. | Athen family Melithaeidae (incl. Acabaria rubra)

4B. Axis composed of proteinous material, which to varying degrees is per-

meated by non-spicular calcium carbonate, or axis composed purely of

non-spicular calcium carbonate without any proteinous material........ 5

SA. Axis continuous and uniform; with a central core that is hollow and cross-

chambered. Central core may be very narrow and inconspicuous in some

FOTIA: 205s) Pre oko eg thn a lee eee Ge bldg once ou ee als, aa 6

SB:

6A.

6B.

TA.

7B.

8A.

SB.

9A.

OB:

10A.

10B.

11A.

11B.

12A.

12B.

13A.

138:

14A.

14B.

15A.

155:

16A.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 287

Axis solid throughout (in forms with a continuous and uniform axis). If

axis jointed, then internodes with hollow central core................ 3)

Chambered core of axis surrounded by smooth sclerites bound together by

sheaths of horn-like material. Sclerites of coenenchyme primarily compact

tiea@naren (RCS A= VO) eid io vs eA es mae oak dea Ideogorgia capensis

Chambered core of axis surrounded by horn-like layers that may contain

non-spicular calcium carbonate. Sclerites not primarily triradiates ...... i

Calyces permanent; with spindles, needles or crown spines arranged into

CICMECINCHEV ROM POMMS 2 Je v.38 i440 6s. be eG ace pte eae yaa 8

Calyces without eight en chevron points; or entire polyp retractile into

SMoMeMme MMe OleCOLOMY:..2/4 irae bias sa Seen hed sk eves conde? Se ue eee ety oes 9

Calyces tall and cylindrical (3-4 mm long, 1,5 mm wide), with distal

thorny crown composed of projecting crown spines . Acanthogorgia spp.

Calyces short and conical (approximately 1 mm long), without distal

HMMOniy chown! Of projecting crown spines. ......:...... Anthogorgia sp.

Coenenchyme contains many sclerites that are balloon clubs (Eunicella

SME oc a an ee cee se iat eet oe tae ere ae 10

CSocnencnyme without balloomelubs 27... 5..4.05..-6- 4254.5. 505- 12

PANOoncubsicomposed Ol TWO CLOWNS . 2... 22.8. .2.22 6 see ae te ee 11

Balloon clubs <0,1 mm in length. Surface of colony papillose, due to the

numerous rounded calyces. Branches cylindrical, without longitudinal

lines on surface. Branching multiplanar (Figs 25-26). . Eunicella papillosa

Balloon clubs 0,1—0,15 mm long. Surface of colony not conspicuously pap-

illose. Branches often somewhat flattened. Surface of branches often with

distinct longitudinal lines or grooves. Branching planar (Figs 22-23)

Nc kis La) a hes he hua Beane Eunicella albicans

Sclerites of coenenchyme are primarily capstans (Leptogorgia spp.) ... 13

Goenenchymal'sclerites are nOt Capstans ...«.....06..05.5225.2.-20 Ny

rane nmecichotomousOnlateral ..2.4.2. 222. 4easus nse eee ee yee oe 14

Elec PAICHIMIMCRTDITUIN ALE ery cesta ce -gel 7 aside ropa ee cue yeah ent area Lea ee oe tate Cat 16

Colony colour red-orange to wine red. Capstans 0,04—0,10 mm long

MISUSE ee ua Haneda peice hea ae ieee mes Leptogorgia palma

Colony colour white or yellow. Capstans 0,06—-0,17 mmlong.......... 15

Colour white. Capstans mainly elongate (Figs 33-35).................-

1 6 8 0 0.0°0. 0-67. G ERT RE ERROR RENO ie Sear ea eee ERIS Leptogorgia barnardi

Colour yellow. Capstans mainly robust (Figs 37-38) .Leptogorgia capensis

Colony colour orange, red-orange, or wine-red. Capstans 0,04—0,10 mm

Ronee tO SAO E43) hs oats au gs ci gue s Sieve tam ou nieya ees Leptogorgia palma

288

16B.

JEN,

17B.

18A.

18B.

19A.

19B.

20A.

20B.

21A.

21B.

22A.

22B.

23A.

23B.

24A.

24B.

25A.

25B.

260A.

26B.

27A.

27B.

28A.

28B.

DON:

ANNALS OF THE SOUTH AFRICAN MUSEUM

Colony colour highly variable, usually yellow, white, pink, or mauve,

rarely red. Many capstans 0,03 mm long, in addition to some larger ones

up to 0,13 mm long (Figs 46-48) ................ Leptogorgia gilchristi

Many coenenchymal sclerites are wart clubs, 0,08—0,12 mmlong .........

ee eS los COM ROR Re oe ee Inte REIN EA ACN Sd Boal No oF otc Rumphella sp.

Sclenites are not want-clubsee.. 2.50 say eee eee ane 18

Calyces low, indistinct or not present. Sclerites are robust tuberculated.

spindles, many ovoid or spheroid in shape ............ Euplexaura spp.

Calyces low or conspicuous. Sclerites predominantly thorn stars, thorn

scales, rooted leaves, or coarsely tuberculate clubs andspindles ....... i)

Calyx sclenitestaremooredileavesm an ar. oe a eee -. Menella sp.

Calyx sclerites are thorn scales, thorn stars or coarsely tuberculate clubs

aNG'SpINCES: i. oa g oe adss Weekes Wed Wise Soa 6 lee ep eee 20

Calyx sclerites are coarsely tuberculate clubs and spindles (Figs 20-21)

sdaeinr oui etl Lait a peepee anh AS Ban Sieh cam aad, Ree a a Acanthomuricea pulchra

Calyx sclerites are mainly thorn scales or thornstars ................. 21

Many calicular sclerites are thorn scales (0,2—0,35 mm in length). Calyces

not armed with thorn stars (Figs 17-18) ............ Astromuricea fusca

Calyces armed with distinct projecting thorn stars (approximately 0,4 mm

GMS MSI) eset tlhe ea a eee eet re Cae wea Echinomuricea spp.

Axis jointed, composed of proteinous nodes and longer internodes of non-

spicular calcium Carbomate . 2.2.6 6056524 2.0<00 502950 eee ree ZS

Axis continuous, not differentiated into nodes and internodes......... US

Branches anise tronmhory, nodes. ..50) 4: 464 eee Acanella sp.

Branches arise from calcareous internodes = 7552-4... ea eee 24

Branching profuse. All polyp sclerites are scales arranged transversely

CEG S6O 97) a2 hee one ek malate, eevee icine ses ern Chathamisis ramosa

Branching sparse. Largest polyp sclerites are needles arranged longitudi-

BaD Ny es seciiarce ere tare neers, Sam eet aie ellen cam fe eae Keratoisis sp.

Sclerites are primarily tuberculated double heads or flattened clubs .... 26

Sclentes are not doublesheads onflattened’clubs2 =... 445 Joes aes Zi

Many sclerites are flattened clubs.......... eee eee Junceella sp.

Many sclentes are double ieads nao ea ee ae family Ellisellidae

Colonies whip-like, unbranched .... 2. ..2.... 0:22.52 24 Sees 28

Colonies sparsely or proiusely branched) 344.540: aan eee 33

Polyps clustered into interrupted whorls. Polyps present on all sides of the

STM ONS SGC BING 22 ie. Ch Dae Ses een os Plame ema Pees fa Primnoella sp.

Polyps arranged longitudinally along one side of colony only. A bare tract

free of polyps runs longitudinally opposite the polyp-bearing side ...... 29

Single polyps longitudinally spaced with intervals of naked stem between

SACHAPOMY MeN oa. ose Holy seer ees Fee Cae oe ee Radicipes sp.

29B.

30A.

30B.

31A.

31B.

32A.

32B.

33A.

33B.

34A.

34B.

35A.

35B.

36A.

36B.

37A.

37B.

38A.

38B.

39A.

39B.

40A.

40B.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 289

Polyps arranged continuously in a longitudinal tract. Adjacent polyps

eloscLOpetherunimternupltedcy, S405. tees co dl ebs a cures mm ane 30

Polyps arranged uniserially along polyp-bearing longitudinal tract

(FS SPO) cumaen 6 Sewers Aiea ae Pee Pare pee Simpsonella squamifera

Polyps arranged multiserially along polyp-bearing longitudinal tract ... 31

Colony forming a tight spiral, corkscrew-shaped. Colony bicoloured;

polyps cream-white, naked tract yellow, orange, or red (Figs 56-57) ....

ey Ss kk ae 2 site ODE edad dees ee Simpsonella spiralis

Colony whip-like or forming a loose open spiral. Colonies cream-white or

Die ON OUC CMe No evi cin lard ae Gas ee ee ee ene 52

Many sclerites are double discs or unilaterally foliate forms

(0,04—0,07 mm long) (Figs 53-54) ............... Simpsonella flagellata

Double discs or unilaterally foliate forms absent (Figs 50-51) ............

PE es Gtk. 8 Bice 2 Sica axmnandars snacb ai Simpsonella capensis

Sclerites are relatively smooth ovoid scales, often slightly constricted

medially with both ends rounded, or sclerites absent .../.............- 34

Sclerites are broad scales or plates sometimes with one end pointed, with

coarse or granular surface tuberculation covering at least one portion of

ECAC CLC INI ME er epee erent Cbd Lol P Gay aban te, « cae saa tetera geno ENG eens 31)

Branching planar, more or less lyrate (with secondary branches upright

and parallel) or dichotomous. Polyps are arranged biserially along

[SVEISLES oo 0 Se geene ee en ate ie nee aE iPr rela NUL Cesare coer ten, Sane 35)

Branching pinnate (bottlebrush-like) or sympodial. Polyps arise on all

SPAS RO IMONAM CIC Sa yee snot) oye satin t bey ke Huei Roe agent a cee ee cue ee een 36

Sclerites present, sparse to numerous (Fig. 62) ..... Trichogorgia flexilis

DelemiesaDSENt .. 6... eee ee ee ee eS Whore: aN Trichogorgia capensis

Branching pinnate (bottlebrush-like). Axis not zigzag-shaped ..........

yn eee Aiea. aire Bec a dees PRD Se Xenogorgia sciurus

Branching sympodial. Axis zigzag-shaped with bronze or gold metallic iri-

SCC ICC MIMN fi ce Ais eee ase amie Meee Gane Los & Caren Chrysogorgia sp.

PS ACANAC MMOS MINT AUS ee ae eet eee el a lee ee ca ts eee oN cannes 38

RAN eMMeichOtOMOUS OF lyrate .... 2s i oe sea ss see ee 39

Branching bottlebrush-like, multiplanar. Polyps clavate, placed individu-

ally along lateral branches (Fig. 66).............. .. Thouarella hicksoni

Branching planar. Polyps arranged in pairs, cylindrical and incurved ...... |

RU tei ue eclede clot a 0) ost a a hs 4. act Primnoeides sp.

Branching lyrate. Polyps downward-facing (Fig. 63).... Narella gilchristi

Branching dichotomous. Polyps upward-facing ..................... 40

Branching sparse. Polyps in whorls arranged on all sides of stem ........

PP inchs Se ain St Nae a. dake a ak) ee arrays Calyptrophora sp.

Branching profuse. Polyps in pairs, each placed on opposite sides of stem

I eee ia et a aP an sate od Sie lull A a amp eon as apats Callogorgia sp.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

LIST OF GORGONIAN OCTOCORALS RECORDED FROM SOUTHERN AFRICA

Order ALCYONACEA Lamouroux, 1816; amended by Bayer, 1981

Family Anthothelidae

Subfamily Anthothelinae

Subfamily Spongiodermatinae

O°

Homophyton verrucosum (Mobius, 1861)

Diodogorgia capensis (Thomson, 1911)

Family Melithaeidae

Acabaria capensis (= Isidella capensis Studer, 1879) [incertae sedis]

Acabaria rosea Tixier-Durivault, 1954

* Acabaria valdiviae Kikenthal, 1908

** Melitodes faurii Thomson, 1917

** Melitodes grandis Thomson, 1917

Mopsella singularis Thomson, 1917

Wrightella coccinea Gray, 1870

** Wrightella fragilis Thomson, 1917

** Wrightella furcata Thomson, 1917

** Wrightella sp. Thomson, 1917

* Wrightella trilineata Thomson, 1917

Family Keroeididae

Ideogorgia capensis (Simpson, 1910)

Family Acanthogorgiidae

** Acanthogorgia spp.

** Anthogorgia sp.

Family Plexauridae

** Echinomuricea spp.

** FEuplexaura spp.

** Menella sp.

° Astromuricea fusca (Thomson, 1911)

Acanthomuricea pulchra (Thomson, 1911)

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA

Family Gorgoniidae

Eunicella albicans (KOlliker, 1865)

Eunicella papillosa (Esper, 1797)

Eunicella tricoronata Velimirov, 1971

Leptogorgia abietina Kukenthal, 1919

Leptogorgia africana Thomson, 1917

Leptogorgia barnardi Stiasny, 1940

Leptogorgia capensis (Hickson, 1900)

Leptogorgia gilchristi (Hickson, 1904)

Leptogorgia luetkeni Wright & Studer, 1889

Leptogorgia palma (Pallas, 1766)

Leptogorgia pusilla Kiikenthal, 1919

Leptogorgia rigida Verrill, 1868-1870

Leptogorgia tenuissima Kikenthal, 1919

Rumphella sp.

Family Ellisellidae

Ellisellidae spp.

Junceella sp.

Family Chrysogorgiidae

[o)

Chrysogorgia sp.

Radicipes sp.

Simpsonella capensis (Simpson, 1910)

Simpsonella flagellata (Simpson, 1910)

Simpsonella spiralis (Hickson, 1904)

Simpsonella squamifera (Kikenthal, 1919)

Trichogorgia capensis Hickson, 1904

Trichogorgia flexilis Hickson, 1904

Xenogorgia sciurus Bayer & Muzik, 1976

Family Primnoidae

O =p +

Callogorgia sp.

Calyptrophora sp.

Narella gilchristi (Thomson, 1911)

Primnoella sp.

Primnoeides sp.

Thouarella hicksoni Thomson, 1911

29]

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Isididae

Subfamily Keratoisidinae

+ Acanella sp.

+ Keratoisis sp.

Subfamily Mopseinae

° Chathamisis ramosa (Hickson, 1904)

°—described and illustrated in the present account

*—material not available for examination.

**—material available but species not presently considered identifiable or valid.

#—material available but damaged or fragmentary and not adequate for valid determination or

detailed description.

+—deep water forms (> 200 m) beyond the scope of the present study and not presently iden-

tified to species.

SUMMARY AND CONCLUSION

Of the thirty genera here considered valid and known to occur in southern

Africa, 17 are new records. These are Diodogorgia, Anthogorgia, Echinomuri-

cea, Menella, Astromuricea, Acanthomuricea, Rumphella, Junceella, Chryso-

gorgia, Radicipes, Callogorgia, Calyptrophora, Primnoella, Primnoeides, Chat-

hamisis, Acanella, and Keratoisis. Six new combinations are presented here:

Diodogorgia capensis, Acabaria rubra, Astromuricea fusca, Acanthomuricea

pulchra, Leptogorgia gilchristi, and Narella gilchristi.

ACKNOWLEDGEMENTS

I am grateful to the following individuals for their helpful comments and

suggestions. Phil Alderslade (Northern Territory Museum, Darwin), Frederick

M. Bayer (Smithsonian Institution, Washington, D.C.), Manfred Grasshoff

(Forschungs-Institut Senckenberg, Frankfurt), George M. Branch and Jenny

Day (University of Cape Town).

I extend my appreciation to Dane Gerneke and Klaus Schultes of the

Electron Microscope Unit (University of Cape Town) for their assistance in the

preparation of scanning electron micrographs.

I thank Liz Hoenson, Sheryl Ozinsky, and Michelle van der Merwe (S.A.

Museum) for curatorial assistance and the preparation of black-and-white prints,

and Marcelle Scheiner and Sandra Saven for typing the manuscript.

The following figures are by Virgilio Branco (South African Museum): 5B,

5C, 7, 10, 11A, 14, 17, 22, 25, 29, 33, 34, 37, 40, 41, 42, 46, 47, 62A, 63A, 66A,

69A. All other figures, photographs, and scanning electron micrographs are by

the author.

_ ————

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 293

REFERENCES

Bayer, F. M. 1955. Contributions to the nomenclature, systematics, and morphology of the

Octocorallia. Proceedings of the United States National Museum 105 (3357): 207-220.

Bayer, F. M. 1956. Octocorallia. Jn: Moore, R. C. ed. Treatise on invertebrate paleontology.

Part F. Coelenterata: 166-231. New York: Geological Society of America; Lawrence:

University of Kansas Press.

Bayer, F. M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata,

Anthozoa), with diagnoses of new taxa. Proceedings of the Biological Society of

Washington 94 (3): 902-947.

Bayer, F. M. 1982. Some new and old species of the primnoid genus Callogorgia Gray, with a

revalidation of the related genus Fanellia Gray (Coelenterata: Anthozoa). Proceedings of

the Biological Society of Washington 95 (1): 116-160.

Bayer, F. M., GrassHorr, M. & VERSEVELDT, J. 1983. Illustrated trilingual glossary of

morphological and anatomical terms applied to Octocorallia. Leiden: E. J. Brill/Dr

W. Backhuys.

Bayer, F. M. & Muzix, K. M. 1976. New genera and species of the holaxonian family

Chrysogorgiidae (Octocorallia: Gorgonacea). Zodlogische mededeelingen 50 (5): 65-90.

Bayer, F. M. & STEFANI, J. 1987. New and previously known taxa of isidid octocorals

(Coelenterata: Gorgonacea), partly from Antarctic waters. Proceedings of the Biological

Society of Washington 100 (4): 937-991.

Bayer, F. M. & STEFANI, J. 1988. Primnoidae (Gorgonacea) de Nouvelle-Calédonie. Bulletin

du Muséum national ad histoire naturelle (4) 10 (3): 449-518.

BretscHowsky, E. 1918. Eine Revision der Familie Gorgoniidae. Inaugural-Dissertation zur

Erlangung der Doktorwiirde der Hohen Philosphischen Fakultat der Schlesischen

Friedrich-Wilhelms Universitat zu Breslau. Breslau: Buchdruckerei H. Fleischmann.

Brancu, G. M. & Branco, M. 1981. Living shores of southern Africa. Cape Town: Struik.

BrocH, H. 1939. Some South African shallow water octactinians. Kungliga Fysiografiska

sdllskapets i Lund forhandlingar 9 (6): 48-79.

Day, J. H. 1974. A guide to marine life on South African shores. 2nd ed. Cape Town: A. A.

Balkema.

Day, J. H., Fietp, J. G. & PENRITH, M. J. 1970. The benthic fauna and fishes of False Bay,

South Africa. Transactions of the Royal Society of South Africa 39 (1): 1-108.

DEICHMANN, E. 1936. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of

the Museum of Comparative Zoology at Harvard College 53: 1-317.

DucHASSAING, P. 1870. Revue des zoophytes et des spongiaires des Antilles. Paris: Victor

Masson et Fils.

DUCHASSAING, P. & MICHELIN, H. 1846. Note sur deux polypiers de la famille des couraux

appartenant aux genres Solanderia et Pterogorgia. Revue Zoologique Société Cuvierienne

(1) 9: 218-220.

DUCHASSAING, P. & MICHELOTTI, J. 1864. Supplément au mémoire sur les coralliaires des

Antilles. Memorie della Accademia delle Scienze di Torino (2) 23: 97-206.

E.uis, J. & SOLANDER, D. 1786. The natural history of many curious and uncommon

zoophytes, collected from various parts of the globe by the late John Ellis, systematically

arranged and described by the late Daniel Solander. London: Benjamin White and Son.

Esper, E. J. C. 1794-1806. Fortsetzungen der Pflanzenthiere (Theil 1). Nurnberg: in der

Raspischen Buchhandlung.

GERMANOS, N. K. 1896. Gorgonaceen von Ternate. Jn: KUKENTHAL, W. ed. Ergebnisse einer

zoologischen Forschungreise in den Molukken und Borneo, im Auftrage der Sencken-

bergischen -naturforschenden Gesellschaft ausgeftiihrt von Dr. Willy Kikenthal, Teil 2,

Band 1. Abhandlungen hrsg. von der Senckenbergischen Naturforschenden Gesellschaft 23

(1): 145-187.

GrassHorF, M. 1977. Die Gorgonarien des 6éstlichen Nordatlantik und des Mittelmeeres. III.

Die Familie Paramuriceidae (Cnidaria, Anthozoa). Meteor Forschungsergebnisse (D) 27:

5-76.

GrassHorF, M. 1982a. Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der

Biskaya (Cnidaria, Anthozoa). Ergebnisse der franzdsischen Expeditionen Biogas,

Polygas, Geomanche, Incal, Noratlante und Fahrten der ‘Thalassa’. I. Ailgemeiner Teil.

Bulletin du Muséum national d’histoire naturelle (4) 3 (A, 3): 731-766.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

GrassHorF, M. 1982b. Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der

Biskaya (Cnidaria, Anthozoa). Ergebnisse der franzdsischen Expeditionen Biogas,

Polygas, Geomanche, Incal, Noratlante und Fahrten der ‘Thalassa’. II. Taxonomischer

Teil. Bulletin du Muséum national @histoire naturelle (4) 3 (A, 4): 941-978.

GrRassHoFrF, M. 1988. The genus Leptogorgia (Octocorallia: Gorgoniidae) in west Africa.

Atlantide Report 14: 91-147.

Gray, J. E. 1858. Synopsis of the families and genera of axiferous zoophytes or barked

corals. Proceedings of the Zoological Society of London 25: 278-294.

Gray, J. E. 1859. Description of some new genera of lithophytes or stony zoophytes.

Proceedings of the Zoological Society of London 27: 479-486.

Gray, J. E. 1866. Description of two new forms of gorgonioid corals. Proceedings of the

Zoological Society of London 1866: 24-27.

Gray, J. E. 1868. Descriptions of some new genera and species of alcyonoid corals in the

British Museum. Annals and Magazine of Natural History (4) 2: 441-445.

Gray, J. E. 1870. Catalogue of the lithophytes or stony corals in the collection of the British

Museum. London: British Museum.

HENTSCHEL, E. 1903. Gorgonacea von Amboina und Thursday Island. Zoologische For-

schungsreisen in Australien und dem Malayischen Archipel. Ausgeftihrt in den Jahren

1891-1893 von Richard Semon 5 (6). Denkschriften der Medizinisch-Naturwissenschaft-

lichen Gesellschaft zu Jena 8: 643-652.

Hickson, S. J. 1900. The Alcyonaria and Hydrocorallinae of the Cape of Good Hope.

Marine Investigations in South Africa 1: 67-96.

Hickson, S. J. 1904. The Alcyonaria of the Cape of Good Hope. Part II. Marine

Investigations in South Africa 3: 211-239.

KOLLIKER, R. A. 1865. Icones histiologicae oder Atlas der vergleichenden Gewebelehre. Zweite

Abtheilung. Der feinere Bau der hdheren Thiere. Erstes Heft. Die Bindesubstanz der

Coelenteraten. Leipzig: Verlag von Wilhelm Engelmann.

KOLLIKER, R. A. 1871. Beitrage zur Kenntniss der Polypen. Verhandlungen der Physikalisch-

medizinischen Gesellschaft zu Wurzburg 2: 11-29.

KUKENTHAL, W. 1908. Die Gorgoniden Familie der Melitodidae Verr. (5. Mitteilung). Zoo-

logischer Anzeiger 33 (7-8): 189-201.

KUKENTHAL, W. 1915. System und Stammesgeschichte der Primnoidae. Zoologischer Anzeiger

46 (5): 142-158.

KUKENTHAL, W. 1919. Gorgonaria. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-

Expedition auf dem Dampfer ‘Valdivia’, 1898-1899 13 (2): 1-946.

KUKENTHAL, W. 1924. Gorgonaria. Tierreich 47. Berlin & Leipzig: Walter de Gruyter

& Co.

Mirne Epwarps, H. & Hare, J. 1857. Histoire naturelle des coralliaires ou polypes

proprement dits 1. Paris: a la Libraire Encyclopédique de Roret.

Mosius, K. 1861. Neue Gorgoniden des naturhistorischen Museums zu Hamburg. Nova Acta

Acad. Caesareae Leopoldino-Carolinae Germanicae Nat. Curiosorum 29: 1-12.

Mo.anper, A. R. 1929. South and West African Octactiniae in the Gothenburg Natural

History Museum. Goteborgs Kungliga vetenskaps- och vitterhetssamhdalles handlingar (B) 1

(7): 1-16.

Nuttinc, C. C. 1910. The Gorgonacea of the Siboga Expedition. IV. The Plexauridae.

Siboga-Expeditie (Livr. 48) Monograph 13"!: 1-20.

Patias, P. S. 1766. Elenchus zoophytorum sistens generum adumbrationes generaliores et

specierum cognitarum succinctas descriptiones cum selectis auctorum synonymis. WHagae

Comitum.

RIpLey, S. O. 1882. Contributions to the knowledge of the Alcyonaria with descriptions of

new species from the Indian Ocean and Bay of Bengal. Annals and Magazine of Natural

History (5) 9: 184-193.

Rip.ey, S. O. 1884. Alcyonaria. In: Report on the zoological collections made in the Indo-

Pacific Ocean during the voyage of H.M.S. ‘Alert’ 1881-2. London: British Museum.

Sars, M. 1856. Nouveau polypes. Jn: Sars, M., Koren, J. & DANIELSSEN, D. C. eds. Fauna

littoralis norvegiae 2: 63-79.

Simpson, J. J. 1910a. On a new pseudaxonid genus— Dendrogorgia. Proceedings of the Royal

Physical Society of Edinburgh 18: 62-67.

GORGONIAN OCTOCORALS OF SOUTHERN AFRICA 295

Simpson, J. J. 19106. Hicksonella, a new gorgonellid genus. Journal of the Royal Micro-

scopical Society 1910 (6): 681-692.

STEARNS, R. E. C. 1883. Descriptions of a new genus and species of alcyonoid polyp, from

Japanese waters, with remarks on the structure and habits of related forms, etc.

Proceedings of the United States National Museum 6: 96-101.

ST1IASNY, G. 1937. Gorgonaria von Cap Blanco, gesammelt durch Dr. Theodore Monod.

Temminckia 11: 297-315.

STIASNY, G. 1939. Gorgonaria von Kap Blanco, Senegal und Rio d’Ouro (Aus dem

Zoologischen Museum, Amsterdam). Revue de Zoologie et de Botanique Africaines 32

(3-4): 285-322.

STiasny, G. 1940. Alcyonaria und Gorgonaria von Siid Afrika aus der Sammlung des South

African Museum, Cape Town. Verhandelingen der K. nederlandsche akademie van

wetenschappen (Afdeeling Natuurkunde, Tweede Sectie) 39 (3): 1-37.

ST1IASNY, G. 1943. Die Gorgonarien-Familie Acanthogorgiidae Kuikenth. & Gorz. mit

besonderer Beriicksichtigung des Materials der Siboga-Expedition. Vorlaufige Mit-

teilung. Zoologische Anzeiger 141 (5-6): 127-133.

STiasny, G. 1947. De Gorgonarien-Familie Acanthogorgiidae Kikenth. & Gorz. (Met

bijzondere Inachtneming van het Materiaal der Siboga-Expeditie.) Verhandelingen der K.

nederlandsche akademie van wetenschappen (Afdeeling Natuurkunde, Tweede Sectie) 43

(2): 1-93.

STuDER, T. 1879. Ubersicht der Anthozoa Alcyonaria, welche wahrend der Reise S.M.S.

Gazelle um die Erde gesammelt wurden. Monatsberichte der Koniglichen Preussischen

Akademie der Wissenschaften zu Berlin 1878: 632-688.

STUDER, T. 1887. Versuch eines Systemes der Alcyonarien. Archiv ftir Naturgeschichte 53 (1):

1-74.

Tuomson, J. A. 1927. Alcyonaires provenant des campagnes scientifiques du Prince Albert I*

de Monaco. Résultats des campagnes scientifiques accomplies par le Prince Albert I 73:

1-77.

THomson, J. A. & HENDERSON, W. D. 1906. The marine fauna of Zanzibar British East

Africa, from collections made by Cyril Crossland, M.A., B.Sc., F.Z.S., in the years 1901

and 1902. Alcyonaria. Proceedings of the Zoological Society of London 1906 (1): 393-443.

THomson, J. A. & RENNET, N. I. 1931. Alcyonaria, Madreporaria, and Antipatharia.

Scientific Reports of the Australasian Antarctic Expedition, 1911-14 (C) 9 (3): 1-46.

Tuomson, J. A. & Simpson, J. J. 1909. An account of the alcyonarians collected by the Royal

Indian Marine Survey Ship Investigator in the Indian Ocean. II. The alcyonarians of the

littoral area. Calcutta: Trustees of the Indian Museum.

TuHomson, J. S. 1911. The Alcyonaria of the Cape of Good Hope and Natal—Gorgonacea.

Proceedings of the Zoological Society of London 1911 (2): 870-893.

THomson, J. S. 1917. South African Gorgonacea. Memoirs and Proceedings of the

Manchester Literary and Philosophical Society 61 (1): 1-56.

Tuomson, J. S. 1923. Charts and comparisons of the distribution of South African Alcyonaria.

With a statement of some of the problems of their dispersal Transactions of the Royal

Society of South Africa 11 (1): 45-84.

TrxteR-DurRIVAULT, T. A. 1954. Les octocoralliaires d’Afrique du Sud. Bulletin du Muséum

national d/histoire naturelle (2) 26 (5): 526-533, 624-631.

Torpiitz, C. M. 1929. Die Gorgonarien Westindiens. Kap. 7, die Famile Gorgonellidae,

Zugleich eine Revision. Zoologische Jahrbticher (Suppl.) 16 (2): 235-376.

Utinom!, H. 1964. Some octocorals from the Antarctic waters off Prince Harald coast. Japa-

nese Antarctic Research Expedition 1956-1962, Scientific Reports (series E) 23: 1-14.

VALENCIENNES, A. 1855. Extrait d’une monogrphie de la famille des Gorgonidées de la classe

des polypes. Compte rendu hebdomadaire des séances de l' Académie des Sciences 41:

7-15.

VAN OFWEGEN, L. P. 1987. Melithaeidae (Coelenterata: Anthozoa) from the Indian Ocean

and the Malay Archipelago. Zoologische verhandelingen 239: 1-57.

VELIMIROV, B. 1971. Beitrag zur Systematik der siidafrikanischen Gorgonien. Zoologischer

Anzeiger 187 (314): 266-273.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

VERRILL, A. E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expe-

dition, under Commodore C. Ringgold and Captain John Rodgers, U.S.N., from 1853

to 1856, collected by Dr. Wm. Stimpson, naturalist to the Expedition. With descriptions of

some additional species from the west coast of North America. Proceedings of the Essex

Institute, Salem, Mass. 4: 181-196.

VERRILL, A. E. 1868. Critical remarks on the halcyonoid polyps in the Museum of Yale

College with descriptions of new genera. American Journal of Science and Arts (2) 45:

411-415.

VERRILL, A. E. 1868-1870. Notes on Radiata in the Museum of Yale College. 6. Review of the

polyps and corals of the west coast of America. Transactions of the Connecticut Academy

of Arts and Science 1: 377—422 (1868); 423-502 (1869); 503-558 (1870).

VERRILL, A. E. 1869a. Critical remarks on halcyonoid polyps. No. 3 American Journal of

Science and Arts (2) 47: 282-285.

VERRILL, A. E. 1869b. Critical remarks on the halcyonoid polyps with descriptions of new

species in the Museum of Yale College, no. 4. American Journal of Science and Arts (2)

48: 419-429.

VERRILL, A. E. 1869c. Synopsis of the polyps and corals of the North Pacific Exploring Expe-

dition, under Commodore C. Ringgold and Captain John Rodgers, U.S.N., from 1853 to

1856. Collected by Dr. Wm. Stimpson, naturalist to the expedition. Additions and cor-

rections. Proceedings of the Essex Institute, Salem, Mass. 6: 75-104.

VERRILL, A. E. 1878. Notice of recent additions to the marine fauna of the eastern coast of

North America, No. 2. American Journal of Science and Arts (3) 16: 371-378.

VERRILL, A. E. 1879. Notice of recent additions to the marine Invertebrata, of the north-

eastern coast of America, with descriptions of new genera and species, and critical remarks

on others. Part I— Annelida, Gephyraea, Nemertina, Nematoda, Polyzoa, Tunicata, Mol-

lusca, Anthozoa, Echinodermata, Porifera. Proceedings of the United States National

Museum 2: 165-226.

VERRILL, A. E. 1883. Report on the Anthozoa, and on some additional species dredged by the

‘Blake’ in 1877-1879, and by the U.S. Fish Commission Steamer ‘Fish Hawk’ in 1880-82.

Bulletin of the Museum of Comparative Zoology at Harvard College 11 (1): 1-72.

VERRILL, A. E. 1884. Notice of the remarkable marine fauna occupying the outer banks off the

southern coast of New England. No. 9. Brief contributions to zoology from the Museum of

Yale College. No. 55. American Journal of Science and Arts (3) 28: 213-220.

VeERRILL, A. E. 1922. Alcyonaria and Actiniaria. Report. Canadian Arctic Expedition

1913-18 8: 1-164.

VersLuys, J. 1902. Die Gorgoniden der Siboga-Expedition. I. Die Chrysogorgiiden. Siboga-

Expeditie Monograph (Livr. 6) Monograph 13: 1-120.

WILuiAMs, G. C. 1989. A provisional annotated list of octocorallian coelenterates occurring on

the sublittoral coral reefs at Sodwana Bay and Kosi Bay, northern Natal, with a key to the

genera. South African Journal of Science 85 (3): 141-144.

WILLIAMS, G. C. 1990. The Pennatulacea of southern Africa (Coelenterata, Octocorallia).

Annals of the South African Museum 99 (4): 31-119.

WILLIAMS, G. C. 1992. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft

corals (Coelenterata, Anthozoa). Annals of the South African Museum 100 (3): 249-358.

WiLuaMs, G. C. (in press a). Biogeography of the octocorallian coelenterate fauna of south-

ern Africa. Biological Journal of the Linnean Society.

WILLIAMS, G. C. (in press b). Revision of the gorgonian genus Simpsonella (Octocorallia:

Chrysogorgiidae) from the western margin of the Indo-Pacific, with the description of a

new species from southeastern Africa. Zoological Journal of the Linnean Society.

Wiiams, G. C. (in press c). Coral reef octocorals: an illustrated guide to the genera of soft

corals, sea fans, and sea pens inhabiting the coral reefs of northern Natal. Durban: Durban

Natural Science Museum.

Wricut, E. P. 1869. Ona new genus of Gorgonidae from Portugal. Annals and Magazine of

Natural History (4) 3: 23-26.

WriGcut, E. P. & StuDER, T. 1889. Report on the Alcyonaria collected by H.M.S. Challenger

during the years 1873-1876. Report of the Scientific Results of the Voyage of H.M.S. Chal-

lenger 1873-1876 31 (1): i-Ixxu, 1-314.

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid- ae region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

i SEECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

ee) - 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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

GARY C. WILLIAMS

THE ALCYONACEA OF SOUTHERN AFRICA.

GORGONIAN OCTOCORALS

(COELENTERATA, ANTHOZOA)

VOLUME 101 PART 9 SEPTEMBER 1992 ISSN 0303-2515

_ ANNALS

‘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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

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

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

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

bered consecutively.

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

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

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

abbreviations.

Footnotes should be avoided unless they are short and essential.

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

should be left to editor and publisher.

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

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

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

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

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

into consideration.

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

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

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

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

ing, if possible.

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

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

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes...’

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

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

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

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

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

each name, with suffixes a, b, etc., to the year for more than one paper by the same author in

that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 19692).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

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

last pages of article).

Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 101 # Band

September 1992 September

Part 9 Deel

7 try

INES

S.-VVVWVV

aug ngy 68

NOVI

REVISION OF THE FRESHWATER

DIAPTOMID GENUS LOVENULA

(CRUSTACEA, COPEPODA) IN AFRICA

NANCY A. RAYNER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

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

OUT OF PRINT/UIT DRUK

iL, Wales, SO), LD, 2-5, Sits, SE 8, S, TD)

Gy toe, MED), &, UY, 7), UGS), WG-2, 5, 71.9.1.

1413), 15425), 242, 5), 27) 313), 326), 33.

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 132 6

Printed in South Africa by In Suid-Afrika gedruk deur

he Rustica weressaetye etd. Die Rustica-pers, Edms., Bpk.,

Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap

D1511

REVISION OF THE FRESHWATER DIAPTOMID GENUS LOVENULA

(CRUSTACEA, COPEPODA) IN AFRICA

Nancy A. RAYNER

Department of Zoology and Entomology, University of Natal, Pietermaritzburg,

South Africa

(With 9 figures and 1 table)

[MS accepted 20 December 1990}

ABSTRACT

In this revision of the African freshwater diaptomid genus Lovenula, keys to the freshwater

calanoid families, the African diaptomid genera and Lovenula species are presented, as well as

diagnoses of the family Diaptomidae and the subfamilies Paradiaptominae and Diaptominae.

Full synonymies, diagnoses, a record of all material examined and a map of distribution, have

been included for each of the four species, L. falcifera, L. africana, L. excellens and

L. simplex. Lovenula (Neolovenula) alluaudi and Lovenula natalensis have been transferred to

the genus Paradiaptomus and the systematic status of the two related genera, Lovenula and

Paradiaptomus, is discussed.

CONTENTS

PAGE

SMOG MCW ON sso odoaaddeon sag dead ae Mage eO COR ECE Se OP rica te a MEER Tasha Chern arenes 297)

SAMMI HCAS CUSSION pee neat einace ee eaatin ckartceet aajec aaa rncmmeme geaeyaee aa osiseieluee 300

henmmologyand abbreviations .-2...- 212... say. e2e sso abso soe en ote seeone 300

Keyatonthe families of freshwater Calanoida..2...............-.+...--.---- 303

ecmtlya Dap tom ae ers aerate eee once eee eon oasis reawio ws as see aeee 304

Subfanuly,Paradiaptominlac.. wy. 2see-ceeceae acess ce scene seen ene een 304

SubfannilyaDiap lOminaea- caja cas Gosccee so eces aessnss lucie os see ars os 304

Keys to the African genera of the family Diaptomidae .................. 305

Genuspeoverntla Schimmel S98y ce nneeccaces ceceneaseasccaceroeiisencns: 306

Kevatoneispecies of the cenus Loventla sea. ...0 eens. sons tn es eee 323

Speciesremoved iromthe eenus Wovenvla .-.......5-.2-s es eee eo 324

Systematic status of Lovenula and Paradiaptomus................. eae 326

PROC 1 COS Ree eee He Gere se A Sania Re Shs we eo aaterac tn cieccive ac tvmcelediva raat 327

RGA C ee ers Ses, Be co ah GRO secrete nO ee Atlan a Sec w SNE Ne 331

INTRODUCTION

Four Lovenula species are endemic to South, East and North Africa:

Lovenula falcifera (Lovén), L. africana (Daday), L. excellens Kiefer and

L. simplex Kiefer. All are comparatively large predators and the possession of

well-developed raptorial maxillipeds is not only a striking morphological feature

but also a highly-weighted taxonomic character. The type species, Broteas

falcifer, was collected by J. Wahlberg from a saline pan near the junction of the

297,

Ann. S. Afr. Mus. 101 (9), 1992: 297-332, 9 figs, 1 table.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crocodile and Apies rivers in the Transvaal (Fig. 1) and deposited in the

Swedish State Museum, where it was described by Loven.

The taxonomy of the genus has been confused for nearly a century, mainly

because research has been undertaken by scientists from European institutions,

most of whom never visited South Africa. They relied on small samples of

animals, or sediment from temporary waters, sent to them by colleagues. An

outstanding contribution was made to the knowledge of South African fresh-

water micro-crustaceans by G. O. Sars, who described new species of Clado-

cera, Ostracoda and Copepoda from southern Africa. Although he did not desig-

nate types, he returned the material to the South African Museum in Cape Town,

where it is still available for study. Other material collected in the early part of

this century was housed in European museums and, in many cases, it has been lost

ANGOLA

_ ZAMBIA

‘ - Cahora Bassa _ L.Chilwa ==i15

Dam

Cunene Owambo L.KariBa “Oy

Etosha Pan Caprivi fits

: ZIMBABWE |

Okavango Delta

Omatako

Makgadikgadi

oWindhoek BOTSWANA

; s

Kalahari owe™

AMIENS . TRANSVAAL

Vaal

Dam Pongola

WALVIS BAY

LUDERITZ

Bloemhof

» Dam

2 Y

om -OES.

P.K.leRoux

Dam_4.F.Verwoerd

Dam

Fig. 1. Map of southern Africa showing countries, cities, major river systems, and some of the

freshwater locations mentioned in the text (see Gazetteer, p. 331).

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 299

or records are not traceable. Tribute must be paid also to Frederich Kiefer, one of

the greatest of all freshwater copepod taxonomists, who, together with Sars, laid

the foundation of the taxonomy of freshwater micro-crustaceans in southern

Africa. Kiefer (1934) reviewed the South African freshwater Copepoda and so

provided a firm basis for current taxonomic research on copepods in southern

Africa. In addition, the world list of calanoid copepods compiled by Dussart &

Defaye (1983) is an important reference work.

The four closely related Lovenula species are among the largest of the

freshwater diaptomids and it is surprising that there has been so much confusion

about the identity of these distinctive predatory copepods. However, the con-

stant changes to the generic name, as well as the large number of synonyms,

testify to the lack of any in-depth study, and Kiefer (1934) was largely dis-

Fig. 2. Map showing river systems and distribution of Lovenula falcifera (@), L. africana (A),

L. excellens (WM) and L. simplex (x).

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

regarded in his attempt to create stability. Sars (1899) redescribed ‘L. falcifera’

from material that had been sent to him from the Cape Flats by W. F. Purcell,

but the specimens used by Sars were L. simplex (yet to be recognized as a new

species by Kiefer in 1929). Rithe (1914) also did not recognize L. simplex as a

new species when he identified it as Paradiaptomus falcifer. Although these two

species are closely related taxonomically, the geographical separation of L. falci-

fera and L. simplex, which is so obvious today (Fig. 2), would surely have been

questioned by early taxonomists if more material had been available for study.

Communication was also a problem as shown by Lovén (1845) giving the type

locality of L. falcifera as the Magaliesberg (Transvaal) on the road to Port

Natal. Van Douwe (1912a) suggested that Sars’ (1899) redescription was not

that of L. falcifera but related to a new species. However, despite Van Douwe’s

misgivings, Sars (1927) again recorded L. simplex as L. falcifera. In his collect-

ing expeditions in Ovamboland in the 1920s, K. H. Barnard had collected a

species of Lovenula from temporary pools in this arid area of Namibia. Sars

(1927), realizing that this was a different species from his L. falcifera (i.e.

L. simplex), named the new species L. barnardi. From this time onwards,

L. simplex was known as L. falcifera and L. falcifera as L. barnardi, an error

perpetuated by Harding & Smith (1967). In addition to the confusion over

L. falcifera and L. simplex, there was a lack of authentic information on

L. excellens, a Lake Chrissie species described by Kiefer (1929). Methuen

(1910) failed to recognize it as a new species, identifying it as ‘a local variety’ of

Broteas falcifer. It is hoped that this revision will create stability in the genus

Lovenula and lead to a reassessment of the Paradiaptominae of Africa. Dussart

(1980) stressed the importance of revising, genus by genus, the African copepod

fauna.

SYSTEMATIC DISCUSSION

TERMINOLOGY AND ABBREVIATIONS

Names of collectors

KHB—K. H. Barnard (S. A. Museum, 1911-1956); FMC—F. M. Chutter;

JAD—J. A. Day; AJCG—A. J. C. Gardiner; RCH—R. C. Hart; JMK—

J. M. King; WFP—W. F. Purcell (S. A. Museum, 1896-1905); MTS—M. T.

Seaman.

Repositories

AM—Albany Museum, Grahamstown; NM—Natal Museum, Pieter-

maritzburg; SAM—South African Museum, Cape Town; TM—Transvaal

Museum, Pretoria; SMN—State Museum, Windhoek, Namibia; BMNH—

British Museum (Natural History); ZMUH— Zoological Museum, University of

Hamburg; HMNH— Hungarian Natural History Museum.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 301

Terminology (see Fig. 3)

Some of the terms are well known to copepodologists but have been

included for non-specialists who may be unfamiliar with copepod terminology.

All measurements are in millimetres. Measurements and coloration are affected

by age and period of time of preservation of the specimen. Curvature of female

antennae affects measurements relative to body length. Thoracic ‘wings’ and

shape of the tip of last segment of male right Al may be affected by orientation

on the microscope slide. Antennule segment numbers refer to existing segments

and do not necessarily relate to homologous ancestral numbers. Material

examined consisted of a number of adult males and females, unless otherwise

stated. Details are presented in the following order: museum catalogue number

(if relevant), locality, collector’s name, date.

per

cephalon

PROSOME

metasome

UROSOME

spiniform

process

Fig. 3. Terminology and abbreviations (see p. 302 for glossary). A. Somites and

body divisions. B. Position of appendages. C. P2. D. Male P5. E. Female P5.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

C. V.—coefficient of variation (%); X = mean.

aesthetask—small, rod-shaped sense organ on Al

anal somite (AS)—somite proximal to furca (Ur2 or 3 in female; Ur5 in male)

antenna 1 (Al)—antennule, the first prosomal limb

antenna 2 (A2)—antenna, the second prosomal limb

basis (B2)—basal stem segment of endopodite and exopodite

body length (TL)—total length excluding furcal setae

cephalon (cephalosome)—head, including maxilliped somite

coxa (B1)—the middle stem segment of a biramous limb

egg sac, brood sac—receptacle for eggs, forming part of genital complex

endopodite (Enp)—inner ramus of biramous appendage beginning with

segment Enp1

exopodite (Exp)— outer ramus of biramous appendage beginning with segment

Exp1

furca (FR)—caudal rami

furcal setae (FS)—five spiny terminal setae and one slender ‘dorsal’ seta arising

from each furcal ramus

genital somite (GS)—first urosomal double somite in female

length: breadth (L:B)—ratio maximum length to breadth of a structure

mandible (Md)—the third prosomal limb

maxilla 1 (Mx1)—maxillule, the fourth prosomal limb

maxilla 2 (Mx2)—maxilla, the fifth prosomal limb

maxilliped (Mxp)—the sixth prosomal and first thoracic limb

metasome—pedigerous somites between cephalon and major articulation of the

body

natatory legs (P1—-P4)—swimming legs, relating to pedigerous somites (Pdg1—4)

non-natatory legs (P5)—modified legs, used in male for spermatophore

transference, in female for egg protection

pedigerous somites (Pdg1—5)—leg-bearing somites

precoxa (PCX)—the proximal stem segment of a biramous limb

prosome (Pr)—complete body section anteriad to the major articulation

(Cephalon + Pdg1—5)

rostrum (R)—a beak-like projection on anterior of the prosome

segment (Seg)—fundamental division of a limb

seta—relatively long, flexible process, tapering to a point, generally with a

double row of fine hairs (setules), giving the appearance of a feather

somite—fundamental body division

spine—short, strong, more or less inflexible, usually armed on each side with a

row of small denticles giving it a saw-like edge

spiniform process—an extension of a segment having the appearance of a spine

spinule—very small spine

thoracic somites (Pdg1—5)—pedigerous thoracic somites

thoracic ‘wings’ (Pdg5 wings)—expansion of last prosomal somite in female to

form wing-like processes

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 303

thorn—a hollow, usually pointed, extension of a limb segment

urosome (Ur)—complete body section posteriad to the major articulation, the

first somite being the non-pedigerous genital double somite

Limnological terms of local origin (Fig. 1)

dam—a man-made impoundment. This term has become common usage in

South Africa, although in the Northern Hemisphere it relates to a wall that

impounds a river.

pan—as defined by King (1951: 81) ‘Pans, in the South African usage of the

term, are shallow depressions varying from a few square feet up to several

Square miles in area which occur typically along a belt stretching north—

north-east from Calvinia, through the northern Cape and the western Orange

Free State to the Transvaal. This belt is about 600 miles long and 100 miles

wide, with a branch into South-West Africa and an outlying patch about Lake

Chrissie in the Eastern Transvaal. After rain they may contain water; but

often they are dry, broad floors of muddy and salty accumulation’.

vlei—as defined by King (1951: 92) ‘The beds of intermittent rivers have in

many cases been so choked by wind-drifted sand or silt that the original

gradients have been lost, and a series of shallow lakes or ‘“‘vleis’”” now

appears after rains instead of a flowing river’.

KEY TO THE FAMILIES OF FRESHWATER CALANOIDA

Adapted from Gurney (1931), Dussart (1967), and Kiefer (19782).

1. Pl endopodite with 3 segments; P2—P4 endopodite with three segments. . 2

— P1 endopodite with fewer than three segments; P2—P4 endopodite with

=) SSL UCIT seas AAs Meme cerca alae dae eee ani en an ae RRr eee: oma 2 3

2. P5in both sexes identical to P1—P4, with three-segmented endopodite ......

NM ieee sk Se Peek is ON ws Ua woe os ee Family Centropagidae

— P5 modified in both sexes, female without endopodite*, male with one-

Sepmmlemled CNGOPOGHe” 2.6. ice et ee Family Pseudodiaptomidae

3. Pl endopodite two-segmented; P2—P4 endopodite with 2-3 segments 4

— Plendopodite one-segmented; P2—P4 endopodite one- or two-segmented ..

Ps Okc hat dah co UME wea aks gE Family Temoridae

4. P2-—P4 endopodite three-segmented; PS with endopodite in both sexes......

MI I OO re esas hs SMI SAAC ealed Family Diaptomidae

— P2-—P4 endopodite two-segmented; P5 without endopodite in both sexes ....

PO ce vt an ita tec Hed acest) iekacta cre Seca eek Family Acartiidae

* Some exceptions (Dr T. C. Walter, Smithsonian Institute, Washington, D.C., U. S. A., pers.

comm.).

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Diaptomidae G. O. Sars, 1903

Centropagidae (part.): Giesbrecht, 1892: 58. Schmeil, 1896: 5-6. Giesbrecht & Schmeil, 1898:

52. Van Douwe, 1909: 4. Thiebaud, 1915: 8.

Diaptomidae Sars, 1903: 83. Pesta, 1928: 27-28. Rylov, 1930: 78; 1935: 169. Gurney, 1931: 108.

Kiefer, 1932a: 460; 1960: 23-24; 1978a: 70. Damian-Georgescu, 1966: 40. Dussart, 1967: 88.

General description of the Diaptomidae

The cephalon is distinct from Pdgl. Pdg4 and Pdg5 are fused and often

expanded in female to form ‘wings’. Female urosome with two or three somites;

urosome of male with five somites and may be asymmetrical. Genital somite of

female is expanded laterally, sometimes with prominent asymmetrical lobes.

Furcal rami are usually longer than broad, typically symmetrical in female,

sometimes asymmetrical in male, with lateral setae on right furcal ramus some-

times modified in male. Female Al and male left Al are 25-segmented (for details

of individual segments with setae, aesthetasks and sensory spines see Kiefer

1978a). Male right Al has 21 or 22 segments, enlarged and modified between

segments 13 and 18, with geniculation between segments 18 and 19. The

21-segmented Al has three terminal, post-geniculate segments; the 22-

segmented Al has four. A2 has a seven-segmented exopodite that is longer than

endopodite. P1 has a two-segmented endopodite and three-segmented exopodite,

Exp2 lacks spine, Exp3 with one or two spines; P2—P4, endopodite and exopodite

with three segments, Exp3 with one spine. P5 is non-natatory and biramous in

both sexes. Female P5 has Exp2 produced mediad into a large, inner, spiniform

process; Exp3 is reduced or absent or represented by one or two movable spines.

Male P5, as well as basal segments of limb, highly asymmetrical, right leg much

longer than left. Right leg has a lateral spine on Exp2; Exp3 is represented by a

long, hinged, terminal blade-like claw. Male left P5 with Exp2 and Exp3 fused

and variously modified, with sensory pads, spinelike outgrowths and setae. A

single egg-sac in female, rod-shaped spermatophore in male.

Kiefer (1932a, 1932b) divided the Diaptomidae into two subfamilies,

Paradiaptominae and Diaptominae.

Subfamily Paradiaptominae Kiefer, 1932a

P1, Exp3 with two outer spines; male right Al with three post-geniculate

segments, terminal segment may have beak-like extension; male left P5, Exp2

with strong, well-developed lateral or medial spiniform process. There are three

genera in the subfamily Paradiaptominae: Lovenula Schmeil, Paradiaptomus

Sars, and Metadiaptomus Methuen. They include some large predatory species

that occur in temporary waters in drier regions and also in permanent waters.

The majority of species are endemic to Africa.

Subfamily Diaptominae Kiefer, 1932a

P1, Exp3 with one outer spine; male right Al with four post-geniculate

segments, terminal segment without extension; male left P5 often with circular,

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 305

serrated Exp2 and spine, if present, short and broad-based. The Diaptominae

are cosmopolitan and include all the rest of the diaptomids, some 45 genera

(Dussart & Defaye 1983).

KEYS TO THE AFRICAN GENERA OF THE FAMILY DIAPTOMIDAE

KEY TO FEMALES

WW RGSOMERWALIEWONSOMITES” Ye uid keh cena sa ee ok oben aes wee Sele y,

Inesome with three SOMILES «02.263 6. c 458 ee pas ta woes Metadiaptomus

Body length usually >2,0 mm, robust build. Maxillipeds raptorial, with

ADR MPIESe Se OMENS Vor taetnc Soe ce fa cs a yet etm os aeihe 3 = oo ensue 3

Body length seldom > 2,0 mm, small build. Maxillipeds not modified, with

taORMlLIBTIVC SECIMENES sce Sete s/s ets wets ne Mee Maw prod oe eos eee 4

Body length 3,0—4,0 mm. Maxillipeds scythe-shaped (Fig. 51). P5, Exp3

represented by two closely-applied spines, outer longer than inner, which

project across large spiniform process of Exp2. Furcal rami twice as long as

Ipn@admsctac without bulbous bases..:.4.............+-...--0- Lovenula

Body length 2,0-3,5 mm. Maxillipeds not as above. P5, Exp3 represented

by two small spines, outer shorter than inner, spines not projected across

Exp2 spiniform process. Furcal rami sometimes lamelliform, expanded dis-

fallyeysetae may have bulbous bases’........:5.........- Paradiaptomus

Eaicndopodite with two'terminal setae..............-.-- Tropodiaptomus

Esrendopodite without terminal setae................. Thermodiaptomus

KEY TO MALES

Right Al, 21-segmented, geniculate between segments 18 and 19, no articu-

lation between segments 20 and 21, diagnostic spiniform processes on

segments 8, 10, 11, 13. Left P5, Exp2 with spiniform process well devel-

OMe CmlatenaOmimedial <5. 6 ae esd d wea onked ocyag eeleneauG see ama Soe 2

Right Al, 22-segmented, geniculate between segments 18 and 19, articu-

lation between segments 20 and 21, diagnostic spiniform processes on

segments 10, 11, 13, 15. P5, Exp2 without spiniform process............ 3

Maxillipeds not raptorial, with endopodite of five segments. Right P5, B2

expanded on inner margin and fringed with small knobs, spines or setae

PIPE ents en ace ina oie lately arate es aus Metadiaptomus

Maxillipeds raptorial with endopodite of <5 segments. Right P5, B2 not

expandedand without knobs, spines\or setae .....-...-.22..-52.22.+-2-5: 4

P5, Exp2 parallel to long axis of body with lateral spine adjacent to Exp3

Dba Ge CRC lA Wines hee oe ee ais Wee tae aia viola alana eas Tropodiaptomus

P5, Exp2 at right angles to long axis of body, with lateral spine separated

inom xis: Diade-like Claw 2... 0g.4..0..cesefee ce ee Thermodiaptomus

* The North African species Paradiaptomus alluaudi has three somites.

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

4. Body length 2,5-4,0 mm. Maxillipeds scythe-shaped (Fig. 51). Left P5,

Exp2 with strong, terminal spiniform process. Setae of right furcal ramus

modified with three lateral setae separated from two medial setae.........

15 Rs RRS EE Bac PRE es i Geeta Siar MR els Mi ORCA Lovenula

— Body length 2,0-3,5 mm. Maxillipeds not as above. Left P5, Exp2 with

short, lateral spiniform process. No separation of setae on right furcal

ramus but outermost seta may be enlarged .............. Paradiaptomus

Genus Lovenula Schmeil, in Giesbrecht & Schmeil, 1898

Broteas Lovén, 1845: 436 (non C. L. Koch, 1839). Sars, 1899: 6-24.

Lovenula Schmeil, 1898 (in Giesbrecht & Schmeil, 1898: 105). Grochmalicki, 1913: 524. Kiefer,

1932a: 461; 1934: 108-110; 1978a: 74. Damian-Georgescu, 1966: 43. Dussart, 1967: 88.

Dussart & Defaye, 1983: 54.

Paradiaptomus Sars, 1907: 3. Stebbing, 1910: 531. Van Douwe, 19126: 25; 1914: 96. Brady,

1913: 467. Gurney, 1929: 581; 1931: 109. Lowndes, 1930: 164. Barnard, 1935: 490.

Paradiaptomus (Lovenula) Dussart, 1989: 32.

Type species. Lovenula falcifera (Lovén, 1845).

Remarks

The generic name of this taxon was originally Broteas, assigned by Lovén

(1845) when he described Broteas falcifer. Schmeil (in Giesbrecht & Schmeil,

1898) changed the name to Lovenula because Broteas was preoccupied by an

arachnid genus. Sars (1895) described Paradiaptomus lamellatus, the type species

of the genus Paradiaptomus. Sars (1907) gave the generic name Paradiaptomus

to both B. falcifer and P. lamellatus as he considered them to be congeneric and

also the name Paradiaptomus was older than Schmeil’s Lovenula. Sars (1927)

conceded that Lovenula and Paradiaptomus were two distinct genera but it was

still a few years before the name Lovenula Schmeil gained full acceptance.

Characteristics of the genus (Figs 4—6)

Length 2,5-4,0 mm, females more robust than males, prosome approxi-

mately two-thirds total length in both sexes (Table 1).

Prosome, urosome and furca. Female with two urosomal somites; last

thoracic double somite (Pdg4—5) of female expanded as ‘wings’ (Fig. 4A).

Female genital somite expanded laterally, usually more pronounced on left

(Fig. 4C). Male urosome of five somites, slender and cylindrical, often curving

to the right or directed ventrally (Fig. 41). Furcal rami in both sexes asymmetri-

cal, right slightly longer than left, each furcal ramus with five strong apical setae

and a slender ‘dorsal’ seta; a very small dorsolateral seta on each furcal ramus

(Fig. 4D); three outer setae on male right furcal ramus are spine-like, lack

setules on outer margin and separate from two inner setae (Fig. 4J).

Antennae. Female Al (Fig. 5A) and left Al of male, 25-segmented with

notably two very short spines on segments 8 and 12. Male right Al (Fig. 5B-D)

21-segmented, geniculate between segments 18 and 19, with terminal segment

307

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA

vl =? 1861 er TOTASTOYITS

£ ) 1861 — IO[ASTOYIIS

02 & 1861 = Te[AQpuoy JOOIH

v 6 1861 sae To]Aopuoy JOOIH

xXajduns “7

0) ee 2 0961 eunt | C6LITTIVA SISSY 2xe']

9 6 096] ount | C6LITTIVA oISSHY oye]

SUI]JaIXA “TT

v 2 9¢C6I «= SI-9° ET V CLOT HNN Ojo e1OH

v } 976I =SI-9' EC CLOT HNNG Too P10H

(sopumoyT DIDWID.A1G

snwojdoipoivd “uKS)

= 6 LS61 APW tb — ued Ipesyxipesyeyy

(jeLayeur Aepeq)

ll 8061 67°C OT 8S61l HNN PMANY OCT

(jeLojeur Kepeq)

[be 8061 6¢°C OI 8861 HNN PMANY OCT

DUDILLD “TJ

LG €76I “JPN ELSTIV-NVS purjoquivaQ

8S 6 €76I “IPN ELSTIV-NVS purjoquivaO

Ol 6 S86I “JEW EI = wed Uls}UOP{IO}S

9 2 L86I Id2S 6 = weq doyusords

GS 8 L86I “dag 6 = weg doyusords

Dlafizof °7

“AD

(TL) Wisu9] [RIO] u—s- xa ajeq Joquinu ongoyeyea Ayipeso']

“‘DINUIAOT JO Satoads Ino} JO (WU UT) sJUSOINSeOyY

1 ATaVL

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

with a beak-like extension; longest diagnostic spiniform process on segment 11,

with spiniform process on segment 10 shorter and curved in towards it; a strong,

broad-based spiniform process on segment 13; the smallest process on segment

8. A2 biramous (Fig. 5E), with two-segmented endopodite and seven-segmented

exopodite; basis with 1-2 setae; Enp1, two setae, lateral border denticulate dis-

tally; Enp2 with two distal lobes with approximately 16 setae (not all shown in

figure); Exp2 and Exp7 long, Exp3—6 compacted; a single, strong seta arises

from Exp3, Exp5, Exp6, and proximal end of Exp7; reduced Exp4 lacks a seta

and Exp7 has three long, terminal setae.

Mouthparts. Mandible (Fig. 5F) consists of a large gnathobasal coxa with a

single tooth and 5-6 smaller teeth separated by a V-shaped cutting edge; man-

dibular palp with four inner setae on basis, two-segmented endopodite with four

setae on Enp1, six setae on Enp2; five-segmented exopodite with one inner seta

on each of Exp1—4, two setae on ExpS. Maxilla 1 (Fig. 6) with precoxa with

13 spine-like setae with bristle-like setules; coxa with two lobes on inner border,

one lobe with three strong setae, the other with two long and two short setae;

basis with one proximal seta and four distal setae of different lengths; one-

segmented endopodite with two lobes, with three short, one long, and four very

long setae, one-segmented exopodite with seven long setae; nine very long setae

on outer border of coxa. Maxilla 2 (Fig. 5G) with precoxa with two lobes with

5 + 3 setae, coxa with 3 + 3 setae, basis with three long and one very small

setae, Enp 1-3 each with spine-like seta; one seta on precoxa, two on coxa and

one on basis are claw-like with fine bristles (Fig. 5H). Maxilliped (Fig. 51) con-

sists of precoxa, coxa, basis and four-segmented endopodite; groups of 1, 2, 3

and 3 setae on coxa and a spiny process at articulation with basis; basis with

fringe of small denticles on inner border with two and one strong setae; Enp1

with two long and two shorter setae; three terminal segments (Enp2-—4) pro-

duced distally to form spiniform processes, smooth on outer margin, denticulate

on inner.

Natatory legs. First natatory leg (P1) (Fig. 4E) with two-segmented endo-

podite and three-segmented exopodite (spine formula: 1, 0, 2); second, third and

fourth natatory legs (P2—P4) (Fig. 4F) with endopodite and exopodite three-

segmented, spine formula of exopodite: 1, 1, 2; endopodites of P2—P4 same

length; exopodites of P3 and P4 longer and sturdier than P2, length being

achieved by extension of Exp1.

Non-natatory P5. Female P5 (Fig. 4G, H), endopodite one-segmented with

two terminal setae, 5—6 sub-apical spinules at base of outer seta; Exp2 and Exp3

fused, Exp2 with an inner spiniform process and an outer broad-based spine;

Exp3 represented by two closely applied spines, outer spine longer than inner,

both projecting across Exp2 spiniform process. Male PS (Fig. 4K—M), right leg,

Exp1 with an inner rounded process, Exp2 with an outer distal spine and Exp3

represented by a terminal, medially denticulate, blade-like claw; endopodite

two-segmented, the last segment ball-shaped with a semi-circular fringe of 6-8

spinules; left leg with one-segmented endopodite, Exp2 produced laterally into a

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 309

strong, medially denticulate spiniform process; on medial margin of Exp2, a

triangular pad fringed with knobs or setae, and between outer spine and inner

pad, a slender apical spine extending diagonally between the two structures.

Kiefer (1932a) divided the genus Lovenula into two subgenera, Lovenula

s.s. and Neolovenula for Diaptomus alluaudi Guerne & Richard, 1890 (see

p. 324).

Lovenula falcifera (Lovén, 1845)

Figs 4-6

Broteas falcifer Lovén, 1845: 436, pl. 6 (figs 1-16). Guerne & Richard, 1889: 118-121,

figs 41-43.

Lovenula falcifera Giesbrecht & Schmeil, 1898: 105, fig. 25. Daday, 19106: 118. Kiefer, 1932a:

481, 484 (figs 6-8), 486; 1932b: 214-215, fig. 1; 1934: 110-115, figs 1-8; 1939: 324,

figs 1-3. Hutchinson et al., 1932: 17-150. Brehm, 1958: 34. Loffler, 1961: 356; 1964: 187;

1968. Kok, 1974: 153-183. Dussart & Defaye, 1983: 54.

non Broteas falcifer Sars, 1899: 6-24, pl. 1 (figs 1-15).

Lovenula mea Gurney, 1904: 300-301, pl. 18 (figs 7-13) (part.).

Diaptomus bouvieri Daday, 1910a: 187, 188, 195, pl. 5 (figs 1-11), text-fig. la—b.

Paradiaptomus falcifer Stebbing, 1910: 532. Tollinger, 1911: 189. Van Douwe, 1912a: 2, pl. 1

(figs 1-5); 1912b: 29, 31; 1914: 95-96. Brady, 1913: 468, pl. 35 (figs 7-10). Gurney, 1929:

582.

Paradiaptomus meus Stebbing, 1910: 533 (part.).

Paradiaptomus gurneyi Tollinger, 1911: 188 (part.). [syn. nov. ]

Lovenula barnardi Sars, 1927: 92, pl. 6 (figs 6-9). Kiefer, 1928: 8. Harding & Smith, 1967: 518.

Paradiaptomus barnardi Lowndes, 1931: 1291; 1933: 308. Barnard, 1935: 490.

Lovenula furcata Brehm, 1958: 34, fig. 29. [syn. nov. ]

Lovenula excellens Brehm, 1958: 35-37 (figs 34-37). Hart, 1984: 1602-1607; 1985a: 151-178;

19855: 17-26; 1986: 351-371; 1987: 287-318 (misidentification).

Paradiaptomus (Lovenula) falcifera Defaye, 1988: 115, table 1, figs 13-16. Dussart, 1989:

33-34, 126, figs 20, 71.

Paradiaptomus (Lovenula) furcata Dussart, 1989: 39-40, 130, fig. 24C.

Material examined (Figs 1-2, Gazetteer)

BMNH 1904.9.21.23-24, BMNH 1951.8.10.68—69, wet specimens labelled

Lovenula mea Gurney, 1904. Identity of male: Lovenula falcifera, female:

Paradiaptomus schultzei van Douwe, 1912b; L. mea placed into synonymy with

L. falcifera (see above). SAM-1501, Port Elizabeth, no details. NM, pond,

Harrismith, E. Warren, February 1908. SAM-—A3787, Kimberley, no details.

HMNH-K8896, Farm Frauenstein, near Neudamm, 50 km east-north-east

Windhoek, Namibia, W. Michaelsen, 13 May 1911. HMNH-K8895, Farm

Otjituezu, near Neudamm, 66km_ north-east of Windhoek, Namibia,

W. Michaelsen, 13 May 1911. SAM-—A4791, Grootfontein, S. H. Haughton,

August 1917. SAM-A11573, Ukualonkathi, Ovamboland, KHB, March 1923.

SAM-A11574, Ongka, Ondangua, Ovamboland, KHB, February 1923.

SAM-A11575, Onambeke, Ovamboland, KHB, April 1923. SAM—A11576,

Ukualuthi, Ovamboland, KHB, April 1923. SAM—A5915, Junction Marico and

Crocodile rivers, R. W. Tucker, 1918. SAM-—A5943, Beaufort West, S. H.

Haughton, August 1917. SAM-—A11581, Aliwai North, S. H. Haughton, 24 July

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

1921. SAM-A11582, Cornets Kop, Molteno, Beaufort West, S. H. Haughton,

June 1921. SAM-—A11584, west of Gouritzrivier bridge, G. E. Hutchinson,

1927. SAM-A11590, Lobatsi, Bophuthatswana, J. H. Power, 1927. SAM-—

A11537, SAM-A11538, SAM-A11539, SAM-A12446, SAM-—A12448, SAM-—

A12449, SAM-—A12450, SAM-—A12451—all these vials contain material from

Ovamboland, identified in G. O. Sars’ handwriting as ‘L. barnard? and relate

to SAM-A11573-6. Barnard sent this material for identification to Sars in

1925 (correspondence files, S. A. Museum). TM-—1225, 1 mile north-east of

Tsotsoroga Pan, 23 June 1930, Vernay-Lang Kalahari Expedition. TM-—1260,

1 mile north-east of Tsotsoroga Pan, 3 July 1930, VLKE. TM—1508, N’Kate Pan,

7 August 1930, VLKE. BMNH 1959.3.2.3-4, Dodoma, Tanganyika, H. J. de

S. Disney, undated. AM—VAL 907A, Vaal River catchment, Stn 3, stones-in-

current, FMC, 11 November 1959. AM-VAL 912A, Vaal River catchment, Stn

VD 17, stones-in-current, FMC, 10 November 1959. AM-GEN 591D, pan

between Middleburg and Belfast, J. Agnew, 23 June 1960. AM-—GEN 595B,

Dewetsdorp, J. Agnew, 29 March 1961. AM—VAL 1379F, Vaal River catch-

ment, Stn SV 9, stones-in-current, FMC, 14 April 1964. AM—VAL 1390B, Vaal

River catchment, Stn SV 9, marginal vegetation, FMC, 14 April 1964. Witbank

Dam, MTS, 2 March 1978. Welbedacht Dam, MTS, July 1980. Mazelspoort

Dam, MTS, August 1981. P. K. le Roux Dam, MTS, 4 July 1981. P. K. le Roux

Dam, RCH, 12 January 1982. SMN-50727, waterhole, eastern Caprivi,

Namibia, S. Bethune, 8 December 1982. SMN-50772, 65 km east of Windhoek,

Namibia, C. Meyer, 23 January 1984. Greyvenstein Dam, MTS, 11 May 1984

and 15 June 1984. Omatako Dam, Namibia, W. A. Smit, 20 September 1984.

AM-GFR, Witmos, J. O’Keefe and F. de Moor, 19 February 1985. Sterk-

fontein Dam, W. Dorgeloh, 13 March 1985. White Hill Pan, 42 km from

Hwange, Zimbabwe, A. J. C. Gardiner, July 1985. Elandsdrift, Great Fish

River, R. W. Palmer, 20 January 1986. Spioenkop Dam, RCH, 9 September

1987. SMN-—51328, Gautscha Pan, Boesmanland, Namibia, B. A. Curtis,

17 March 1988. SMN-51330, Tjokwe Pool, Boesmanland, Namibia, B. A. Curtis,

17 March 1988. Quaggaplaat, Sutherland, temporary pool under snow, JMK,

31 August 1988. D510/02, Brak River airfield, JMK, 31 August 1988. Nuweleeu-

rivierdam, RCH, 18 January 1989. D560/02, Quaggaplaat, JMK, 2 August 1989.

Description

Measurements are given in Table 1 and abbreviations on p. 302.

Female (Figs 4-6). Large, robust build (Fig. 4A); Al extends to posterior

border of GS; antennae and mouthparts (Figs 5A, E-I, 6); thoracic ‘wings’

markedly expanded and backwardly directed, left longer than right, two points

on left ‘wing’, one on right with a small additional spine on each ‘wing’; GS

expanded more on left than right, slightly shorter than AS (Fig. 4C); furca

(Fig. 4D); P1 (Fig. 4E); P2 (Fig. 4F); P5 strongly developed, spiniform process

of Exp2 often carried at right angles to Exp1; endopodite same length as Exp1,

with two strong terminal setae, more than half length of endopodite; small

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA S11

5 RE

CULL

K =

POR, Kl

SS =

: SK KK

Fig. 4. Lovenula falcifera. A-H. Adult female. A. Dorsal view. B. With egg sac and

spermatophore. C. Thoracic ‘wings’ and GS, dorsal view. D. Furca, dorsal view.

E. Left P1, posterior view. F. Left P2, posterior view. G. P5, posterior view. H. PS,

Exp2 and Exp3, posterior view. I-M. Adult male. I. Dorsal view. J. Furca, dorsal

view. K. P5, posterior view. L. P5, left leg, Exp2, anterior view. M. P5, right leg,

EnupZ- ventral view. Bar scales n mm. A, B, I=1. D, F, K, L=0;5: C, E, G, J

= 0,25. H, M=0,05.

S12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Lovenula falcifera. Antennae and mouthparts. A. Female left Al. B. Male right Al.

C. Male Al, segments 8-13. D. Male A1, end of terminal segment. E. Antenna 2. F. Man-

dible. G. Maxilla 2. H. Maxilla 2, claw-like setae of coxa and basis, ventral view.

I. Maxilliped. Bar scales in mm. A, B, I=0,5. C, E, F, G=0,25. D, H=0,05.

313

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA

PCX

Fig. 6. Lovenula falcifera. Maxilla 1.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

process on outer margin of Exp2 lateral spine (Fig. 4G—H). Egg sac (Fig. 4B),

details from three localities:

1. One female, c. 258 eggs, bilobed sac, L. 0,8 x B. 1,2 mm (SAM-A4791,

temporary pool, Grootfontein, 1917).

2. Range 9-43 eggs, k = 21,6, P. K. le Roux Dam (Hart 1987).

3. One female, c. 116 eggs, sac L. 0,85 x B. 1,0 mm (D560/02, Quaggaplaat,

1989).

Male (Figs 4-5). Slender build (Fig. 41); right Al strongly developed,

segments 13-18 expanded and flattened (Fig. 5B—D); furca (Fig. 4J); PS, right

leg, Enp2 spinules short and rounded; left leg endopodite well developed, with

2-3 apical spinules; Exp2 with strong, semi-circular spiniform process

(Fig. 4k—M). Spermatophore length, range 0,75—0,95 mm.

Coloration

In freshly collected specimens, there may be indigo pigment deposited

internally in appendages and furca. J. M. King, University of Cape Town (pers.

comm.), noted that L. falcifera collected from a snow-bound pond at

Quaggaplaat, near Sutherland, was blue-green. Barnard (1935) recorded

L. falcifera from near Tsotsoroga Pan, Kalahari, as having a sky-blue body and

anterior and posterior appendages bright red. Specimens collected by Warren in

1908 from a pond near Harrismith, were described as follows: ‘. . . the body

milky white, the antennae purple, the furca bright red round the base and

purple terminally, the setae purple’ (Brady 1913).

Historical

Lovenula falcifera was collected by J. Wahlberg, from a salt pan in the

Magaliesberg, between the Crocodile and Apies rivers, Transvaal. Lovén gave

the specific name ‘falcifer’—bearing a scythe or sickle—referring to the sickle-

shaped, large raptorial maxillipeds. Swedish museum authorities declined to

supply any information regarding types.

Distribution (Fig. 2)

In South Africa, Lovenula falcifera occurs in temporary, often saline,

waters on the highveld, major impoundments on the Orange and Vaal rivers;

Ovamboland and other areas of Namibia; and Kalahari (Barnard 1935).

Hutchinson et al. (1932) collected the species in 1928 from Transvaal pans:

Bothasrust, Brakpan 1 and 2, Eliazar, Florida Grass, Leeuwkraal, Rietfontein 1

and 2, Weltevreden East and West; from O. F. S.: Morgenson farm dam; from

Eastern Cape: Stormberg Dam. Also recorded from Kenya (Van Douwe

1912a), Uganda (Lowndes 1931), East African high mountain lakes (Loffler

1964, 1968), and Ethiopia (Defaye 1988).

Lovenula falcifera is widely distributed, especially in drier areas in

temporary pools but it also has the ability to colonize major man-made water

bodies (Hart 1984, 1985a, 1985b, 1986, 1987; Defaye 1988). Despite its wide

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA S15

distribution, it has not been recorded from low-lying areas, vleis, coastal lakes

or coastal plains (Fig. 2). It appears to be confined to the high plateau of Africa

above 1 000 m, much of which lies in the pan belt (see p. 303), as well as high

mountain lakes in East Africa. It is obviously well adapted to the extreme

temperature ranges of these regions.

Lovenula falcifera often co-occurs with a species of Metadiaptomus as prey

species. It has been recorded as co-existing with M. meridianus (Witmos,

Sterkfontein Dam, P. K. le Roux Dam, Greyvenstein Dam, Mazelspoort Dam,

Quaggaplaat Pan, Nuweleeurivier Dam), M. colonialis (Spioenkop Dam, White

Hill Pan in Zimbabwe, Omatako Dam and Neudamm in Namibia) and

M. transvaalensis (a pan between Middleburg and Belfast, Tvl.). In Witbank

Dam in the eastern Transvaal, L. falcifera co-exists with Thermodiaptomus

syngenes.

Remarks

In southern Africa, the confusion of L. falcifera with L. simplex and

L. excellens has detracted from correct interpretation of the biology of this

important diaptomid.

Lovenula africana (Daday, 1908)

Biss 7

Diaptomus africanus Daday, 1908: 45, 46 (figs 25a—e); 1910a: 111, pl. 5 (figs 1-13). Tollinger,

1911: 55. Cunnington, 1920: 559. Gurney, 1929: 579-580.

Paradiaptomus biramata Lowndes, 1930: 163, pl. 1 (figs 1-7), pl. 3 (figs 1-8); 1933: 308-309.

Lovenula africana Daday, 1910b: 118. Kiefer, 1932a: 461, 486, 489 (figs 15-17); 1934: 119-122,

figs 19-23; 1939: 324. Brehm, 1958: 37. Loffler, 1961: 356 (table 2), 363. Dussart &

Defaye, 1983: 55. Green, 1986: 496-498.

Paradiaptomus africanus Lowndes, 1936: 6, figs 1A—H, 2A-B. LaBarbera & Kilham, 1974:

461-464.

Paradiaptomus (Lovenula) africanus Harding, 1942: 180.

Paradiaptomus (Lovenula) africana Defaye, 1988: 112, table 1, figs 8-12. Dussart, 1989: 37,

129, figs 23A, 70.

Material examined (Figs 1-2, Gazetteer)

HMNH-1908 (Daday’s material), Lake Rukwa. BMNH 1932.4.23.6-15,

syntypes of Paradiaptomus biramata Lowndes, Hora Keloli, Ethiopia, J. Omer

Cooper, 1926. BMNH 1941.5.16.50-60, Lake Rukwa, East Africa, C. K.

Ricardo and R. J. Owen, undated. BMNH 1958.10.2.29, Makgadikgadi main

pan, Botswana, D. H. Eccles, 4 May 1957. Sowa Pool, north-eastern tip of

Sowa Pan, Makgadikgadi, Botswana, R. C. Hart, 10 July 1990; males and

females with egg sacs and spermatophores.

Description

Measurements are given in Table 1 and abbreviations on p. 302.

Female (Fig. 7A—E). Moderate size (Fig. 7A); Al extends to posterior

border of GS; thoracic ‘wings’ asymmetrical with three dissimilar processes on

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

KEE ESCO CREEK EEC KEE

Fig. 7. Lovenula africana. A-E. Adult female. A. Dorsal view. B. Thoracic ‘wings’

and GS, dorsal view. C. Furca, dorsal view. D. P5, posterior view. E. P5, Exp2 and

Exp3, posterior view. F-M. Adult male. F. Dorsal view. G. Furca, dorsal view.

H. Right Al. I. Al, segments 8-13. J. Al, end of terminal segment. K. PS, posterior

view. L. PS, left leg, Exp2, posterior view. M. P5, right leg, Enp2, posterior view. Bar

scalesinmm. A, F=1. B, H=0,5. C, D, G, I, K=0,25. L=0,1. E, J, M=0,05.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA Sil

left side, outer process pointed, medial finger-like, inner rounded; GS slightly

expanded, same length as AS (Fig. 7B); furca (Fig. 7C); P5, endopodite weakly

developed, distinctly shorter than Expl, setae small, one-quarter length of

endopodite; spiniform process of Exp2 directed backwards; Exp2 lateral spine

shorter than in other species and with barely discernible process (Fig. 7D-E);

egg sac, 13-14 eggs (Daday 1908; paralectotype).

Male (Fig. 7F—M). Moderate size (Fig. 7F); Al moderately developed

(Fig. 7H—J); furca (Fig. 7G); P5, right leg, Enp2 spinules slender and pointed;

left leg endopodite well developed, with 2—3 apical spinules; Exp2 produced to

form a straight, backwardly-directed spiniform process (Fig. 7K—M).

Type material

Daday (1908) did not designate type material (Dr Laszlo Forr6, HMNH,

pers. comm.). Type locality, Lake Rukwa, Tanzania. Lectotype and paralecto-

types designated from Daday’s original material, deposited by author in 1989,

with the Hungarian Natural History Museum, Budapest. Catalogue numbers:

lectotype III/P—382; paralectotypes, male III/P—383, female III/P—384. Examin-

ation of syntypes of Paradiaptomus biramata Lowndes (BMNH 1932.4.23.6-15)

confirmed its synonymy with L. africana (Kiefer 1934).

Distribution (Fig. 2)

Only one southern African record, Makgadikgadi Pan, Botswana; other

African records: in Ethiopia: Hora Keloli, Hora Horeso (Lowndes 1930);

in Kenya: Lake Elementeita, Rift Valley (Lowndes 1933); crater lake, 2 miles

west of Lake Naivasha (Lowndes 1936); Lake Rukwa (Harding 1942); in

Kenya, Uganda, Tanzania, Rwanda: Lakes Big Momela, El Kekhotoito,

Embagai, Eyasi, Kusare, Magad, Manyara, Mikuyu, Nakuru, Reshitani, and

Tulusia (LaBarbera & Kilham 1974); in Ethiopia: Lakes Paulo, Bishoftu,

Arenguade, Kilotes, Abijata, Langano, and Arenguade (Green 1986; Defaye

1988).

Lovenula africana occurs in waters with high conductivity from Ethiopia

through East Africa, with the southernmost record being Makgadikgadi Pan,

Botswana. The conductivity range of 11 East African lakes was 3 350-15 000

umhos cm“! (x = 11 439 + 5 528), the highest conductivity range recorded for

any of 11 common species of copepods (LaBarbera & Kilham 1974). This

suggests that L. africana has an affinity for saline waters, confirmed by its

occurrence in Makgadikgadi salt pan. The salt concentration in this pan may

reach proportions such that flamingos’ legs are encrusted with salt deposits

(R. Kennard, pers. comm.). Metadiaptomus transvaalensis has been recorded as

the prey species of Lovenula falcifera in Makgadikgadi Pan. In Lake Bunyonyi,

Metadiaptomus aethiopicus co-occurs with L. africana (Lowndes 1936). Cun-

nington’s (1920) record of L. africana from Lake Malawi is doubtful, because

of the low conductivity of the lake (210 wmhos cm~').

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lovenula excellens Kiefer, 1929

Fig. 8

Lovenula excellens Kiefer, 1929: 309-310, figs 1-3.

Broteas falcifer Methuen, 1910: 159, pl. 16 (fig. 45a—b). [syn. nov.]

Lovenula excellens Kiefer, 1932a: 461, 487-488, figs 12-14; 1934: 102, 106-107, 116-119,

figs 15-18. Hutchinson et al., 1932: 1-150, pl. 8 (fig. 2). Loffler, 1961: 356 (table 2), 363.

Dussart & Defaye, 1983: 56.

Paradiaptomus excellens Gurney, 1929: 582.

Paradiaptomus (Lovenula) excellens Dussart, 1989: 38-39, 130, figs 24A, 70.

Material examined (Figs 1-2, Gazetteer)

AM-VAL 676B, Lake Chrissie, bottom sediment, FMC, 10 December

1958. AM-VAL 720D, Vaal River catchment, Stn VD 2, Eckman grab, FMC,

10 December 1958. AM-VAL 1179D, Lake Chrissie, bottom sediment, FMC,

1 June 1960.

Description

Measurements are given in Table 1 and abbreviations on p. 302.

Female (Fig. 8A—-E). Moderate size, robust build (Fig. 8A); Al extends to

posterior border of AS; thoracic ‘wings’ not markedly expanded, left slightly

longer than right; GS with cone-shaped expansion on left side; AS slightly

longer than GS (Fig. 8B); bases of FS widely separated, setae fine, long and

tapering (Fig. 8C); P5, small, setae on endopodite one-quarter length of endo-

podite, inner seta shorter; Exp2 lateral spine with well-developed process on

outer margin (Fig. 8D—-E); egg sac, unknown.

Male (Fig. 8F—M). Slender build (Fig. 8F); right Al slender (Fig. 8H—J);

right FR, three outer FS shorter than inner two (Fig. 8G); P5, left leg,

endopodite reduced, Exp2 spiniform process curved distally; right leg, Enp2

spinules sturdy and pointed; Exp2 lateral spine short, blunt; Exp3 claw not

strongly developed (Fig. 8K—M); spermatophore length 0,62.

Historical

Type material was not designated. Kiefer’s microscope slide numbers:

01154*, 01155*, 01156, 01165* (Dr Ulrich Franke, Landessammlungen fur

Naturkunde, Karlsruhe, Germany, pers. comm.).

Material for description was obtained from alkaline pans (Avenue,

Banagher 3, Blaauwater 1-5, Eilandspan, Liefgekosen, Magdalenasmeer North,

Rietkuil) in Lake Chrissie area, Eastern Transvaal, South Africa, collected May

1928, by Hutchinson et al. (1932).

Distribution (Fig. 2)

Lovenula excellens has been recorded only from pans in the Lake Chrissie

area and the upper Vaal River catchment. The Lake Chrissie pans are peculiar

in that they are relics of an ancient drainage system belonging to the Umpilusi

* with diagram.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 319

Fig. 8. Lovenula excellens. A-E. Adult female. A. Dorsal view. B. Thoracic ‘wings’

and GS, dorsal view. C. Furca, dorsal view. D. PS, posterior view. E. P5, Exp2 and

Exp3, posterior view. F-M. Adult male. F. Dorsal view. G. Furca, dorsal view.

H. Right Al. I. Al, segments 8-13. J. Al, end of terminal segment. K. P5, posterior

view. L. P5, left leg, Exp2, anterior view. M. PS, right leg, Enp2, posterior view. Bar

scalesinmm. A, F=1. C, H=0,5. B, D, G, I, K=0,25. E=0,1. J, L, M=0,05.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

River, from which the Vaal River has captured the headwaters (King 1951).

Lovenula excellens and Metadiaptomus transvaalensis were recorded as co-

existing in Lake Chrissie (Methuen 1910).

Remarks

This species posed a major problem in the resolving of the taxonomy of the

Lovenula species. It was described by Kiefer (1929) from material collected by

Hutchinson et al. (1932) in the Lake Chrissie area in the Eastern Transvaal and

was not mentioned again until its identity was queried by Kok (1974). Kiefer

(1929, 1934) did not emphasize the main taxonomic differences between

L. excellens and L. falcifera, although Hutchinson et al. (1932) stated clearly

that L. excellens was different enough from L. falcifera to be a good species.

More recently, a misidentification of L. falcifera as L. excellens from the Orange

River impoundments (Hart 1984) almost entrenched the supposition that

L. falcifera was a temporary pool-dweller and L. excellens a permanent-water

species, although the recent origin of man-made water bodies rules out this

possibility. The identity of L. excellens was confirmed by examination of

material collected by Chutter (1963) in his 1958-1960 survey of Lake Chrissie

and the Vaal River catchment.

Lovenula simplex Kiefer, 1929

Fig. 9

Broteas falcifer Sars, 1899: 6-24, pl. 1 (figs 1-15) (non Lovén, 1845).

Paradiaptomus falcifer Rihe, 1914: 8-9, 29-32, fig. 10a-f.

Lovenula falcifera Sars, 1927: 86, pl. 5 (figs 1-12). Harding & Smith, 1967: 518.

Lovenula simplex Kiefer, 1929: 309-310, figs 1-3; 1932a: 461, 487, figs 9-11; 1934: 105,

116-117, 123, figs 9-14. Gurney, 1929: 582. Brehm, 1958: 37. Dussart & Defaye, 1983: 56.

Gardiner, 1988: 181-229.

Paradiaptomus (Lovenula) simplex Dussart, 1989: 38, 129, figs 23B, 69.

Material examined (Figs 1-2, Gazetteer)

SAM-—A12440, ponds, Green Point Common, Cape Town, WFP, 1898,

labelled Lovenula falcifer. SAM-—A3788, Cape Flats, KHB, 1916. SAM—

A11534, Sars’ writing, Broteas falcifer. SAM—A11535, Sars’ writing Broteas

falcifer. SAM—A11549, Green Point Common, Paradiaptomus falcifer. SAM-

A11551, Touwsrivier, Paradiaptomus falcifer. SAM-—A11015, Diep River

Quarry, Milnerton, E. G. H. Oliver, undated. SAM-—1503, Faure, near Zee-

koevlei, ?1920. SAM-A12468, SAM-A12469, SAM-—A12470, very old, no

details, ?late 1800s. 5 Si 1T, 9 Si 1T, 9 Si 2T, 10 Si 2T, four different samples

from Sirkelsvlei, AJCG, August 1981. 14 GR 2T, 16 GR 2T, 17 GR 2T, three

different samples from Groot Rondevlei, AJCG, August 1981. Gi 1T, Gillidam,

AJCG, November 1981. SWT-19A, Sirkelsvlei, JAD, 1 September 1983.

SWT-27A, temporary pool, Noordhoek, JAD, August 1983. SWT-17H, 10 km

south-east of D. F. Malan Airport, Cape Town, JAD, 12 August 1983. De

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA Sys

Hoop Vlei, sites 1, 3, 4, 5, JMK, 1 November 1986. G501/18, Wiesdrift, JMK,

11 May 1989. G501/18, Soetendalsvlei ditch, JMK, 12 May 1989.

Description

Measurements are given in Table 1 and abbreviations on p. 302.

Female (Fig. 9A—E). Large, robust build (Fig. 9A); Al extends to posterior

border of GS; thoracic ‘wings’ slightly expanded, left longer than right, an

additional small spine on each ‘wing’; GS slightly expanded; AS twice length of

GS (Fig. 9B); furca (Fig. 9C); P5, setae on endopodite strong, about one-third

length of endopodite; Exp2 spiniform process well developed, directed back-

wards (Fig. 9D-E); egg sac, c. 87 eggs, L. 0,83 x B. 1,25 mm (SAM-—A11534);

c. 113 eggs (SAM-1503).

Male (Fig. 9F-N). Large (Fig. 9F); right Al strongly developed, segments

8-13 expanded and flattened (Fig. 9H—J); urosome sturdy, right FR with three

outer FS tapering distally (Fig. 9G); PS long, right leg extending to posterior

border of furca; left leg, endopodite reduced to a rounded process; Exp2

spiniform process straight and backwardly directed (Fig. 9L); right leg, Enp2

spinules sturdy and pointed (Fig. 9N); Exp2, outer spine slender, one-quarter

length of claw; Exp3 claw with slight S-shape (Fig. 9K, M); spermatophore

length 0,65 mm.

Historical

Type material was not designated. Kiefer’s microscope slide numbers:

01183, 01192*, 01193*, 01194* (Dr Ulrich Franke, Landessammlungen fir

Naturkunde, Karlsruhe, Germany, pers. comm.). Kiefer’s (1929) description

related to material collected from Simon’s Town, Cape Peninsula, by Ruthe

(1914) and housed in the Berlin Museum.

Distribution (Fig. 2)

Lovenula simplex is restricted to the coast of the Cape Province where it

has been recorded only from humic vleis (e.g. Gillidam, Sirkelsvlei and Groot

Rondevlei), pools on the Cape Peninsula and environs, and De Hoop Vlei on

the southern Cape coast. Gillidam is a small closed basin (35 m X 10 m) with

dark-brown water and large water-level changes, with pH 3,8—4,1, maximum

depth 1,4m, temperature range 10,5—29,0°C. Sirkelsvlei is a closed basin (475 m

x 90 m), with dark-brown water and large water-level changes, with pH 6,3-6,7,

maximum depth 1,4 m, temperature range 10,5—26,0 °C. Groot Rondevlei is an

open basin (420 m X 360 m), with brown water and moderate water-level

changes, with pH 5,7-6,4, maximum depth 1,6 m, temperature range 11,0-

26,0 °C (Gardiner 1988). In the Cape Province, Lovenula simplex co-occurs with

Metadiaptomus capensis (Green Point Common, De Hoop Vlei, Sirkelsvlei,

Varkensvlei, and Vermont Pan) and with Metadiaptomus purcelli (Cape Flats,

Gillidam, Groot Rondevlei, and Soetendalsvlei).

* with diagram.

sy)

ANNALS OF THE SOUTH AFRICAN MUSEUM

“ss

ee Toe j

SS Q

op)

Fig. 9. Lovenula simplex. A-E. Adult female. A. Dorsal view. B. Thoracic ‘wings’ and GS,

dorsal view. C. Furca, dorsal view. D. P5, posterior view. E. P5, Exp2 and Exp3, posterior

view. F-O. Adult male. F. Dorsal view. G. Furca, dorsal view. H. Right Al. I. Al, seg-

ments 8-13. J. Al, end of terminal segment. K. P5, posterior view. L. P5, left leg, Exp2,

anterior view. M. PS, right leg, Exp3 claw, medial view. N. P5, right leg, Enp2, posterior and

anterior views. Bar scalesinmm. A, F, H=1. C, K=0,5. B, D, G, I, L,M=0,25. E,N,

J =0,05.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 323

Remarks

Lovenula simplex has a very restricted distribution (Fig. 2), a fact that has

been overlooked because of the confusion of this species with L. falcifera by

Sars (1899, 1927), Rihe (1914), and Harding & Smith (1967). Kiefer (1929,

1934) pointed out that it was not the same species as L. falcifera but his opinion

was disregarded. Gurney (1929) noted that L. falcifera was ‘one variable

species’, not realizing in fact that there were two species involved.

KEY TO THE SPECIES OF LOVENULA

Figs 4—9

KEY TO FEMALES

1. Genital somite strongly expanded on left side. P5, Exp2 lateral spine with

Pee_Ociicw Mrocess ON OULEN Margin 265.2. 666. cee ee bee bee ee ee 2

— Genital somite slightly expanded on left side. P5, Exp2 lateral spine with

Mer asthitipEOCESS Of OULET MATEIN 6... os cc care. Ce tien ee Sa ee ee 3

2. Body length 3,0-3,2 mm. Genital somite with cone-shaped expansion on

left side, PS endopodite with paired terminal setae one-quarter of its length,

inner seta shorter than outer. P5, Exp2 lateral spine with pronounced

POScCSSOMOULEE MATOIM 2... eo. Chk eee te ene ee Lovenula excellens

— Body length 3,2—4,0 mm. Genital somite with rounded expansion on left

side. P5 endopodite with paired terminal setae more than half its length,

setae of equal length. P5, Exp2 lateral spine with small process on outer

SUEDE ccc ou 200 SNe eee ene oe eee aca Lovenula falcifera

3. Anal somite twice length of genital somite. Left thoracic ‘wing’ with two

MME UMGEOESSSCS 6. SS eo bn de da Sede eae ea eee Lovenula simplex

— Anal somite and genital somite of equal length. Left thoracic ‘wing’ with

BiBCCRNSSIMMlAT PTOCESSES .. 2. ce ee ee ee eee Lovenula africana

KEY TO MALES

1. Body length 3,0-4,0 mm. Geniculate (right) Al, combined length of last

two segments twice length of ante-penultimate segment. Left P5, Exp2

spiniform process curved; right P5, Exp2 lateral spine short ............ Zz

— Body length 3,2—4,0 mm. Geniculate (right) Al, combined length of last

two segments slightly exceeds length of ante-penultimate segment. Left P35,

Exp2 spiniform process straight; right P5, Exp2 lateral spine long ....... 3

2. Geniculate (right) Al strongly developed with segments 13-18 flattened

and expanded. Left P5, Exp2 spiniform process semi-circular; right P5,

ice spine Slender and pomted ...........606 50.0005 Lovenula falcifera

— Geniculate (right) Al less strongly developed with segments 13-18 slightly

expanded. Left P5, Exp2 spiniform process curved distally; right PS, Exp2

feces spine very Short and blunt 2... 5. a. coe nk ee Lovenula excellens

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

3. Right P5, Exp3 with blade-like claw with slight S-shape, Exp2 lateral spine

one-quarter length of claw. Left P5 endopodite reduced to rounded process

RM nice Vic Cee PLA NP einen Rta oil Seep a Eun Se Lovenula simplex

— Right P5 Exp3 with blade-like claw slightly curved, Exp2 lateral spine one-

sixth length of claw. Left P5 endopodite well developed with 2-3 terminal

SPU Ses Na oki) se Netra oa ate Nae Wig a Sea Lovenula africana

SPECIES REMOVED FROM THE GENUS LOVENULA

Two species, Paradiaptomus alluaudi and P. natalensis, have been removed

from the genus Lovenula.

Paradiaptomus alluaudi (Guerne & Richard, 1890)

Diaptomus alluaudi Guerne & Richard, 1890: 198-201; 1891: 213-217. Richard, 1893: 465-466,

figs 32-33. Poppe & Mrazek, 1895: 131. Schmeil, 1896: 177, pl. 14 (figs 7-9). Giesbrecht

& Schmeil, 1898: 67, 93. Daday, 1910b: 118. Tollinger, 1911: 52, fig. Pl. Gurney, 1929:

576, 581. Gauthier, 1931.

Diaptomus ungviculatus Daday, 1891: 48, pl. 4 (figs 4-9).

Diaptomus lorteti Barrois, 1891: 277, figs 6-11.

Lovenula (Neolovenula) alluaudi Kiefer, 1932a: 461, 486, figs 18-20. Gauthier, 1933b: 128.

Kiefer, 1934: 123; 1958: 158-160; 1978a: 74, 231, pl. 13; 1978b: 494. Damian-Georgescu,

1966: 44. Dussart, 1967: 89, fig. 23. Dumont & Decraemer, 1977: 258. Dussart & Defaye,

1983: 56.

Paradiaptomus lorteti Gauthier, 1933a: 64.

Neolovenula alluaudi Gauthier, 1938: 116. Loffler, 1961: 356, table 2. Dumont & Verheye,

1984: 319.

Paradiaptomus (Lovenula) alluaudi Dussart, 1989: 40—41, 130, fig. 24B.

Material examined

TM—uncatalogued slide of 2 males, labelled Neolovenula alluaudi,

Ostrovan-See, Mazedonien, H. W. Schafer, 9 December 1941.

BMNH 1961.6.28.1, wet specimens labelled Diaptomus alluaudi, Galatui

Lake, near Calarasi, Romania, A. D. Georgescu, undated.

Remarks

Kiefer (1932a) created the subgenus Neolovenula for Diaptomus alluaudi

and included it in the genus Lovenula as Lovenula (Neolovenula) alluaudi.

Characters that this species has in common with other Lovenula species are two

external spines on last segment of P1, asymmetrical thoracic ‘wings’ of female,

beak-like extension on last segment of male right Al and shape of exopodite of

male left P5. Characters not typical of Lovenula species are the small adult size

and the maxillipeds that do not have the very characteristic scythe-shape. Also,

in the female, the urosome has three somites, expansion of female genital

somite is dorso-ventral not lateral, endopodite of female P5 is reduced and

without terminal setae, outer Exp3 spine is shorter than inner, and the spines do

not project across Exp2. In the male, the right furcal setae are not modified,

spine on segment 11 of male A1 is shorter than spine on segment 13, whereas in

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA B75

all other Lovenula species it is longer, male PS has no endopodite on left leg,

and right P5 is atypical.

Paradiaptomus alluaudi is included in the subfamily Paradiaptominae on the

possession of two external spines on the last segment of the exopodite of the first

swimming leg, although the proximal spine is reduced. It cannot, however, be

included in the genus Lovenula even as a subgenus, as it lacks four of the most

highly weighted characters of the genus, namely, the massive raptorial maxil-

lipeds, the modification of the right furcal setae in the male and, in the female, a

urosome with two somites and the arrangement and projection of spines on the

P5. Paradiaptomus alluaudi is as tenuously related to the genus Paradiaptomus

as it is to the genus Lovenula. A solution to the problem would be to raise

Neolovenula to full generic status in the Paradiaptominae, and to include

‘Paradiaptomus’ alluaudi as a species in the monotypic genus Neolovenula.

Paradiaptomus natalensis (Cooper, 1906)

Adiaptomus natalensis Cooper, 1906: 97-103, pl. 12 (figs 1-14). Tollinger, 1911: 190, fig. YS.

Van Douwe, 19125: 29, 31.

Diaptomus pictus Brady, 1913: 464, pl. 34 (figs 1-6).

Paradiaptomus natalensis Gurney, 1929: 582.

Lovenula natalensis Kiefer, 1932a: 461; 1934: 122-123. Dussart & Defaye, 1983: 55.

Paradiaptomus (Lovenula) natalensis Dussart, 1989: 36.

Metadiaptomus natalensis Dussart, 1989: 128, fig. 22.

Material examined (Fig. 1, Gazetteer)

AM-VAL 372F, males and females, ground pool, near Lindeque’s drift,

8 km below Vaal River barrage, F. M. Chutter, 29 April 1958.

Remarks

Kiefer (1932), recognizing this species as a paradiaptomid, placed it in the

genus Lovenula although it is of small size and lacks both the large raptorial

maxillipeds and the three spinous setae on the male right furcal ramus—two

highly weighted Lovenula characters. Also, the female has a dorsal keel on the

last thoracic somite. The shape of the endopodite of the male right PS is a good

Paradiaptomus character and the absence of important Lovenula characters

justify placing this species in the genus Paradiaptomus. The shape and arrange-

ment of spines on the male geniculate Al is almost identical to that of

P. alluaudi, a fact that may indicate a relationship between these two species.

Gurney (1929) stated that A. natalensis was a typical Paradiaptomus and should

be included in that genus. Kiefer (1934) noted that the status of this species

could be decided only if the poor descriptions of Cooper (1906) and Brady

(1913) could be checked against new material, as no types were designated. It

was 45 years before this rare species was collected again by Chutter in 1958

(Chutter 1963), and its examination confirmed that it ranks as a species of

Paradiaptomus.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATIC STATUS OF LOVENULA AND PARADIAPTOMUS

Gurney (1929) assessed the status of South African diaptomids, but his

attempt to form two series, one for Lovenula and one for Paradiaptomus, was

flawed by poor descriptions and misidentifications. Gauthier (1951), in describ-

ing Paradiaptomus rex, discussed the affinities of the two genera. He stated

incorrectly that the only character that distinguishes the two genera is the

massive raptorial maxillipeds of Lovenula. More recently, Dussart & Defaye

(1983) indicated that Lovenula should be made a subgenus of Paradiaptomus, so

Defaye (1988) and Dussart (1989) established Paradiaptomus as the genus, with

subgenera Paradiaptomus s.s. and Lovenula. However, the four Lovenula

species have been shown to be closely related and current research leaves no

doubt that they have enough characters in common with one another, and dif-

ferent from Paradiaptomus, to retain generic status. These characters are the

large size of the adult, the massive raptorial maxillipeds, arrangement of spini-

form processes and shape of the terminal segment of geniculate Al, exopodite

of male left P5, the three outer spiny setae on the male right furcal ramus, and

the relative sizes and arrangement of Exp2 spines of female P5. A comparison of

the morphology of the mandible, maxillae 1 and 2 and maxilliped of the two

genera gives a good indication that Paradiaptomus has retained more of the

characters of their most recent common ancestor than has Lovenula. Feeding

appendage morphology is important in diaptomid systematics.

Many of the early taxonomists interchanged the generic names Lovenula

and Paradiaptomus, but Kiefer (1932a, 1934) never had any doubts as to which

species he placed in the genus Lovenula. He was the only taxonomist to study all

four Lovenula species and he described two of them. Confusion arose because

species of uncertain affinities such as L. mea, L. furcata and L. natalensis were

included in the genus. A major factor also has been the lack of any in-depth

study on the freshwater copepods of southern Africa and the tendency to think

of southern Africa in isolation from the rest of the continent. Ongoing studies

on the Diaptomidae have shown that the wealth and diversity of freshwater

diaptomids in southern Africa is such that their further study will have a major

impact on both the biogeography and systematics of the diaptomids in Africa.

Therefore, it seems pointless at this stage, to reduce a now known stable genus

(Lovenula) to subgeneric status in a genus (Paradiaptomus) that has as its type

Paradiaptomus lamellatus Sars, 1895, a species that lacks most of the important

Lovenula characters. Also, with the recent discovery of at least four new, as yet

undescribed, species of Paradiaptomus in southern Africa, that genus will need

to be reassessed.

ACKNOWLEDGEMENTS

My sincere thanks are due to collectors and museum curators who have so

generously provided me with material, in particular Mrs L. Brown, Transvaal

Museum, Pretoria, Ms B. A. Curtis, State Museum, Windhoek, Mrs M. G. van

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 327

der Merwe, South African Museum, Cape Town, Dr F. C. de Moor, Albany

Museum, Grahamstown, Dr G. A. Boxshall and Ms S. H. Halsey, Natural

History Museum, London, Professor G. Hartmann, Zoological Museum, Uni-

versity of Hamburg, Dr L. Forro, Hungarian Natural History Museum,

Budapest. I am especially grateful to Dr Boxshall for allowing me to examine

material during my 1987 visit and for his kindness to me then, and subsequently,

in authorizing loans and obtaining old literature. Many collectors sent material

direct, in particular, Drs J. A. Day, A. J. C. Gardiner, R. C. Hart, J. M. King

and M. T. Seaman. The University of Natal Geography and Audiovisual

Departments helped with reproduction of the illustrations, in particular, Mrs H.

Margeot and Mr M. Hunt.

I am honoured and deeply grateful to the late Professor G. E. Hutchinson

for his encouragement and for giving me his personal holdings of a large number

of Kiefer’s reprints. My colleagues, Prof. C. C. Appleton and Prof R. C. Hart

gave generously of their time and knowledge over many discussions; Professor

B. Dussart and Dr T. C. Walter showed me a new dimension in copepod tax-

onomy; Dr J. A. Day, University of Cape Town, gave immeasurable support

and advice. I wish to thank Miss E. Louw for her editorial advice and support.

My special thanks are due to Professor J. Heeg, my thesis supervisor, who pro-

vided expertise, advice and support, and whose enthusiasm for the subject has

been a sustaining influence on my work. Financial support from the FRD,

CSIR, and the University of Natal Research Fund, is gratefully acknowledged.

REFERENCES

*——not seen in the original

BARNARD, K. H. 1935. Scientific results of the Vernay-Lang Kalahari Expedition, March to

September, 1930. Annals of the Transvaal Museum 16 (3): 481-492.

Barrois, T. 1891. Notes préliminaires sur la faune des eaux de Orient. 1. Sur trois Diapto-

mus nouveaux des environs du Caire. Revue biologique du nord de la France 3: 230-234,

277-281, 316-318.

Brapy, G. S. 1913. On Entomostraca from various parts of South Africa. Annals of the Natal

Museum 2 (4): 459-474.

BreuM, V. 1958. Crustacea: Phyllopoda und Copepoda Calanoida. Jn: HANstTROM, B., BRINCK,

P. & RuDEBECK, G. eds. South African animal life 5: 26-39. (Swedish Expedition to south-

ern Africa, 1950-51.) Stockholm: Almqvist & Wiksell.

CuuTTer, F. M. 1963. Hydrobiological studies on the Vaal River in the Vereeniging area.

Part 1: Introduction, water chemistry and biological studies on the fauna of habitats other

than muddy bottom sediments. Hydrobiologia 21 (1): 1-65.

Cooper, A. W. 1906. Notes on a new species of Gymnoplea from Richmond, Natal, South

Africa; Adiaptomus natalensis (gen. et sp. nov.) Annals of the Natal Museum 2 (1):

97-103.

CunnincTon, W. A. 1920. The fauna of the African lakes: a study in comparative limnology

with special reference to Tanganyika. Proceedings of the Zoological Society of London

1920 (4): 507-622.

Dapay, E. 1891. Beitrage zur mikroskopischen Siisswasserfauna Ungarns. Conspectus Diapto-

morum faunae hungaricae. Természetrajzi fiizetek 13 (4): 48.

Dapay, E. 1908. Adatok Német-Kelet-Afrika édesvizi mikrofaunaja nak ismeretéhez. Mate-

matikai és természettudomanyi értesit6 26 (1): 43-57.

Dapay, E. 1910a. Species aliquot novae Entomostracorum Archivum zoologicum 1 (14):

187-195.

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dapay, E. 19106. Untersuchungen tber die Siisswasser Mikrofauna Deutsch-Ostafrikas. IX.

Copepoda. Zoologica 59 (1-5): 1-314.

DamiANn-GEorGESCU, A. 1966. A. Calanoida (forme de apa dulce). Fauna Republicii Socialiste

Romania 4 (8): 1-128.

DerFayE, D. 1988. Contribution a la connaissance des Crustacés Copépodes d’Ethiopie.

Hydrobiologia 164: 103-147.

DouweE, C. vAN. 1909. Copepoda, Ruderfusskrebse. 1. Eucopepoda, freilebende Copepo-

den. Jn: Brauer, A. ed. Die Stisswasserfauna Deutschlands 11: 1-136. Jena.

DouweE, C. vAN. 1912a. Ost-afrikanische Stisswasser-Copepoden. Zoologische Jahrbiticher

(Systematik) 33 (1): 1-8.

DouwE, C. vAN. 1912b. Neue Stisswasser-Copepoden aus Siidafrika. In: ScHULTZE, L. ed.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zen-

tralen Stid-Afrika ausgeftihrt in den Jahren 1903-1905 5 (1): 22-33. Denkschriften der

Medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 17 (1): 22-33.

DouweE, C. vAN. 1914. Copepoda. In: MICHAELSEN, W. ed. Beitrdge zur Kenntnis der Land-

und Stisswasserfauna Deutsch-Stidwestafrikas 1: 93-103. Hamburg: Friederichsen.

Dumont, H. J. & DECRAEMER, W. 1977. On the continental copepod fauna of Morocco.

Hydrobiologia 52 (2-3): 257-278.

Dumont, H. J. & VERHEYE, H. M. 1984. The nature and origin of the crustacean zooplankton

of Sahelian Africa, with a note on the Limnomedusa. Hydrobiologia 113: 313-325.

Dussart, B. H. 1967. Les Copépodes des eaux continentales d'Europe occidentale. 1. Cala-

noides et Harpacticoides. Paris: N. Boubée & Cie.

Dussart, B. H. 1980. Les crustacés copépodes d’ Afrique catalogue et biogéographie. Hydro-

biologia 72: 165-170.

Dussart, B. H. 1989. Crustacés copépodes calanoides des eaux intérieures africaines. Crusta-

ceana, Supplement 15: 1-205.

Dussart, B. H. & DeEFAyYE, D. 1983. Répertoire mondial des Crustacés Copépodes des eaux

intérieures. 1. Calanoides. Paris: C. N. R. S.

GARDINER, A. J. C. 1988. A study on the water chemistry and plankton in blackwater lakelets

of the south-western Cape. Ph.D thesis, University of Cape Town. 240 pp.

GauTHIER, H. 1931. Faune aquatique du Sahara Central. Récoltes de M. L. G. Seurat au

Hoggar en 1928. Bulletin de la Société dhistoire naturelle de Afrique du Nord 22 (8):

350-400.

GauTHiER, H. 1933a. Nouvelles recherches sur la faune des eaux continentales de l Algérie et

de la Tunisie. Bulletin de la Société d'histoire naturelle de l Afrique du Nord 25 (5): 63-68.

Gautier, H. 1933b. Faune aquatique du Sahara central. Récoltes de M. Th. Monod dans

’Emmidir et ’Ahnet. Bulletin de la Société d’histoire naturelle de l'Afrique du Nord 24

(5): 127-132.

GautTuieR, H. 1938. La vie aquatique dans les deserts sub-tropicaux. Jn: LECHEVALIER, P. ed.

La vie dans la Région Désertique Nord-tropicale de (Ancien Monde: 107-120. Paris:

Société de Biogéographie.

GautuigR, H. 1951. Contribution a l'étude de la faune des eaux douces au Sénégal (Entomo-

stracés). Alger: Imprimerie Minerva.

GIESBRECHT, W. 1892. Systematik und Faunistic der pelagischen Copepoden des Golfes von

Neapel. Fauna u. Flora des Golfes von Neapel und der angrenzenden Meeresabschnitte 19:

1 SSie

GIESBRECHT, W. & SCHMEIL, O. 1898. Copepoda I. Gymnoplea In: Deutsche Zoologische

Gesellschaft. Das Tierreich 6: 1-169. Berlin: Friedlander.

GrEEN, J. 1986. Zooplankton associations in some Ethiopian crater lakes. Freshwater Biology

16 (4): 495-499.

GrocHMALIcKI, J. 1913. Beitrage zur Kenntnis der Stisswasserfauna Ost-Afrikas. Copepoda

und Ostracoda. Bulletin de l’ Academie des Science de Cracovie (B) 1913: 517-537.

GUERNE J. DE & RICHARD, J. 1889. Revision des Calanides d’eau douce. Mémoires de la

Société zoologique de France 2: 53-231.

GuerNE J. DE & RicHarD, J. 1890. Déscription du Diaptomus alluaudi n. sp., recueilli par

M. Alluaud dans un réservoir d’eau douce a Lanzarote (Canaries). Bulletin de la Société

zoologique de France 15: 198-200.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA 329

GuerneE, J. DE & RICHARD, J. 1891. Synonymie et distribution géographique de Diaptomus

alluaudi. Bulletin de la Société zoologique de France 16: 213-217.

Gurney, R. 1904. Ona small collection of freshwater Entomostraca from South Africa. Pro-

ceedings of the Zoological Society of London 1904 (2): 298-301.

Gurney, R. 1929. Notes on the Diaptomidae of South Africa. Annals and Magazine of

Natural History (10) 4 (24): 572-584.

Gurney, R. 1931. British freshwater Copepoda. General part; Systematic part, Calanoida.

London: The Ray Society.

Harpinc, J. P. 1942. Cladocera and Copepoda collected from East African lakes by Miss

C. K. Ricardo and R. J. Owen. Annals and Magazine of Natural History 19) 11 (51):

174-191.

Harpine J. P. & SmitH, W. A. 1967. Some South African fresh water Copepoda. Annals of

the South African Museum 48 (21): 515-521.

Harr, R. C. 1984. Zooplankton community grazing in silt-laden Lake le Roux, Orange River,

South Africa. Verhandlungen der Internationalen Vereinigung Limnologie 22: 1602-1607.

Hart, R. C. 1985a. Seasonality of aquatic invertebrates in low-latitude and Southern Hemi-

sphere inland waters. Hydrobiologia 125: 151-178.

Hart, R. C. 19855. Embryonic development times of entomostracan zooplankton from Lake

le Roux (Orange River, South Africa), and their possible relationships to seasonal succes-

sion. Hydrobiologia 127: 17-26.

Hart, R. C. 1986. Zooplankton abundance, community structure and dynamics in relation to

inorganic turbidity, and their implications for a potential fishery in subtropical Lake le

Roux, South Africa. Freshwater Biology 16 (3): 351-371.

Hart, R. C. 1987. Population dynamics and production of five crustacean zooplankters in a

subtropical reservoir during years of contrasting turbidity. Freshwater Biology 18 (2):

287-318.

Hutcuinson, G. E., Pickrorp, G. E. & ScHUURMAN, J. F. M. 1932. A contribution to the

hydrobiology of pans and inland waters of South-Africa. Archiv ftir Hydrobiologie 24:

1-154.

KieFerR, F. 1928. 2. Beitrage zur Copepodenkunde (VIII). Zoologischer Anzeiger 76 (1-2):

5-18.

KrererR, F. 1929. Neue und wenig bekannte Siisswasser-Copepoden aus Sudafrika. Zoo-

logischer Anzeiger 80 (10-12): 309-316.

KiEFER, F. 1932a. Versuch eines Systems der Diaptomiden (Copepoda Calanoida). Zoo-

logische Jahrbticher (Systematik) 63 (4): 451-520.

KIEFER, F. 1932b. Versuch eines Systems der altweltlichen Diaptomiden (Copepoda Cala-

noida). Zoologischer Anzeiger 100 (7-8): 213-220.

Krerer, F. 1934. Die freilebenden Copepoden Siidafrikas. Zoologische Jahrbticher (Systema-

tik) 65 (2): 99-192.

Krerer, F. 1939. Crustacea. IV. Copepoda: Diaptomidae, Cyclopidae. Jn: Mission Scienti-

fique de ’Omo 5 (Zoologie). Mémoires du Muséum nationale dhistoire naturelle 9:

319-378.

Krerer, F. 1958. Uber einige Diaptomiden (Crustacea Copepoda) aus der Zentralsahara.

Zoologischer Anzeiger 160 (7-8): 158-160.

Krerer, F. 1960. Ruderfiisskrebse (Copepoda). (2nd edition, with addendum, 1973.) Stuttgart.

KieFeER, F. 1978a. Freilebende Copepoda. Jn: Das Zooplankton der Binnengewasser. Binnen-

gewdsser 26 (2): 1-343.

Krerer, F. 19785. Zur Kenntnis der Copepodenfauna agyptischer Binnengewasser. Archiv ftir

Hydrobiologie 84 (4): 480-499.

Kine, L. C. 1951. South African scenery. 2nd ed. London: Oliver & Boyd.

Kok, D. J. 1974. Aspects of the taxonomy and distribution of the genus Lovenula (Copepoda,

Calanoida) in the H. F. Verwoerd Dam. In: VAN ZINDEREN BAKKER, E. M. ed. The

Orange River Progress Report: 153-185. Bloemfontein: Institute of Environmental Sci-

ences, University of the Orange Free State.

LABArRBERA, M. C. & KituHam, P. 1974. The chemical ecology of copepod distribution in the

lakes of East and Central Africa. Limnology and Oceanography 19 (3): 459-465.

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

LOFFLER, H. 1961. Beitrage zur Kenntnis der iranischen Binnengewasser. II. Jnternationale

Revue der gesamten Hydrobiologie und Hydrographie 46 (3): 309-406.

LOFFLER, H. 1964. The limnology of tropical high-mountain lakes. Verhandlungen der Inter-

nationalen Vereinigung Limnologie 15: 176-193.

“LOFFLER, H. 1968. Die Crustaceenfauna der Binnengewasser Ost-afrikanischer Hochberge.

Hochgebirgsforschung 1: 107-170.

Loven, S. 1845. Fyra nya arter af Sétvattens-Crustacéer fran Sédra-Afrika. Kungliga Svenska

vetenskaps-akademiens handlingar 1845: 427-439.

Lownpes, A. G. 1930. Freshwater copepods from Abyssinia collected by Mr. J. Omer

Cooper. Proceedings of the Zoological Society of London 1930 (1): 161-179.

Lownpes, A. G. 1931. A small collection of Entomostraca from Uganda, collected by Mr.

G. L. R. Hancock. Proceedings of the Zoological Society of London 1931 (4): 1291-1299.

Lownpes, A. G. 1933. Reports on the Percy Sladen expedition to some Rift Valley lakes in

Kenya in 1929. V. Copepods from the Rift Valley lakes in Kenya. Annals and Magazine

of Natural History (10) 11 (63): 307-313.

Lownpes, A. G. 1936. Scientific results of the Cambridge Expedition to the East African

lakes, 1930-1. No. 16. The smaller Crustacea. Journal of the Linnean Society (Zoology)

40 (269): 1-31.

METHUEN, P. A. 1910. On a collection of freshwater Crustacea from the Transvaal. Proceed-

ings of the Zoological Society of London 1910 (1): 148-166.

Pesta, O. 1928. Krebstiere oder Crustacea. I. Ruderfiisser oder Copepoda (1. Calanoida,

2. Cyclopoida). In: DAHL, F. ed. Die Tierwelt Deutschlands und der angrenzenden Mee-

resteile 9: 1-136. Jena: Fischer.

Poppe, S. A. & MRrRAzeEK, A. 1895. Entomostraken des naturhistorischen Museums in

Hamburg. Siisswasser-Copepoden. Jahrbuch der Hamburgischen wissenschaftlichen

Anstalten 12: 125-134.

RICHARD, J. 1893. Copépodes recueillis par M. le Dr Th. Barrois en Egypte, en Syrie et Pale-

stine (Mars—Juin 1890). Revue biologique du Nord de la France 5: 465-466.

RUtueE, E. 1914. Die Siisswassercrustaceen der Deutschen Siidpolar-Expedition 1901-1903 mit

Ausschluss der Ostracoden. Deutsche Siidpolar-Expedition 1901-03 16 (8):5—66.

*RyLov, W. M. 1930. Keys to the determination of freshwater organisms of the U. S. S. R.

Part I. The freshwater Calanoida of the U. S. S. R. Leningrad.

*RyLov, W. M. 1935. Das Zooplankton der BinnengewAsser. Einfiirung in die Systematik und

Okologie des tierischen Limnoplanktons mit besonderer Beriicksichtigung der Gewasser

Mitteleuropas. Binnengewdsser 15: 1-272.

Sars, G. 1895. On some South-African Entomostraca raised from dried mud. Skrifter udg.

Videnskabsselskabet i Christiana (1, Mathematik-naturvidenskab Klasse) 1895 (8): 1-56.

Sars, G. O. 1899. On the genus Broteas of Lovén. Archiv for Mathematik og Naturvidenskab

21 (2): 1-27.

Sars, G. O. 1903. An account of the Crustacea of Norway 4. Copepoda Calanoida. Bergen:

Bergen Museum.

Sars, G. O. 1907. On two new species of the genus Diaptomus from South Africa. Archiv for

Mathematik og Naturvidenskap 28 (8): 3-17.

Sars, G. O. 1927. The fresh-water Entomostraca of the Cape Province (Union of South

Africa). Part 3: Copepoda. Annals of the South African Museum 25 (1): 85-149.

SCHMEIL, O. 1896. Deutschlands freilebende Siisswasser-Copepoden. 3. Teil: Centropagidae.

Bibliotheca zoologica, Stuttgart 8 (21): 1-144.

STEBBING, T. R. R. 1910. General catalogue of South African Crustacea. Annals of the South

African Museum 6 (4): 281-593.

THIEBAUD, M. 1915. Copépodes. Jn: Catalogue des Invertébrés de la Suisse 8: 1-125.

Geneva: Museum d’histoire naturelle.

TOLLINGER, M. A. 1911. Die geographische Verbreitung der Diaptomiden und anderer Suss-

und Brackwasser-Gattungen aus der Familie der Centropagiden. Zoologische Jahrbiicher

(Systematik) 30 (1): 1-302.

THE FRESHWATER DIAPTOMID GENUS LOVENULA IN AFRICA

GAZETTEER

Cape Province: CP; Natal: NTL; Orange Free State: OFS; Transvaal: TVL.

Aliwal North, CP

Apies and Crocodile rivers, TVL

Beaufort West, CP

Belfast, TVL

Brak River airfield

Bunyonyi, lake

Cape Flats, CP

Caprivi, Namibia

Chrissie, lake, TVL

De Hoop Vlei, CP

Dewetsdorp, OFS

D. F. Malan Airport, CP

Diep River, Milnerton, CP

Dodoma, Tanganyika

Faure, CP

Gautscha Pan, Boesmanland, Namibia

Gillidam, CP

Gouritzrivier, CP

Great Fish River, Elandsdrift, CP

Green Point Common, CP

Greyvenstein Dam, TVL

Grootfontein, CP

Groot Rondevlei, CP

Harrismith, OFS

Hora Keloli, Ethiopia

Kimberley, CP

Lobatsi, Bophuthatswana

Makgadikgadi Pan, Botswana

Marico River, TVL

Mazelspoort Dam, OFS

Middleburg, TVL

Molteno, near Beaufort West, CP

Neudamm, east of Windhoek, Namibia

Noordhoek, CP

Nuweleeurivierdam, OFS

Omatako Dam, Namibia

Ombika (Onambeke), Namibia

Ondangua, Namibia

Ongka, Namibia

P. K. le Roux Dam, CP

Quaggaplaat, CP

Richmond, NTL

Rukwa, lake, Tanzania

Sirkelsvlei, CP

Soetendalsvlei, CP

Sowa Pan, Makgadikgadi, Botswana

Spioenkop Dam, NTL

Sterkfontein, TVL

Stormberg, CP

Tamsu (Tamansu), Namibia

Tjokwe Pool, Boesmanland, Namibia

Touwsrivier, CP

21°00'S

25°10'S

29°02’S

25°48'S

31°23'S

22°30'S

34°07'S

2920'S

21S

19°20’S

ti5oS

1735'S

30°35’S

3232/9

229309

F30'S

34°16'S

34°44'S

20°10'S

28°40'S

28°35'S

3125'S

18°35'S

19°00'S

33°20'S

MW Ww

NM CO ON

c oO oO

2 bo

(1)

(a)

IWM eR I

(om) = 0O

fe) ) oO ) [e)

—

fa}

—

CO UN

[e) le} (e}

io)

(3)

C Oo» C (+ ) )

NERNSORBROKWDARNERE ROAR OW

QDNASNNSERSNSSISWONADSOHDBDASS

—"

(e)

sMesMesMesMesMescsMesMesMesMesMesMeoMesMesMesMesMesMeoMeshesMesMesies)

(=)

32 30'E

1825'E

19°58’E

26°10'E

29°30'E

29°04'E

26°40'E

20°36’E

20°00’E

20°03’E

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ukualonkathi, Ovamboland, Namibia PASTS 15°40’E

Ukualuthi, Ovamboland, Namibia 17°40'S 1593078

Vaal River catchment, TVL 26°45'S 27-3048

Welbedacht Dam, OFS 29°54'S 2652

White Hill Pan, Hwange, Zimbabwe 18°20’S 26335 E

Wiesdrift, CP 34°40'S 19°55’E

Windhoek, Namibia DIB SES Ifs05vE

Witbank Dam, TVL WS aS) 29°18’E

Witmos, CP 32733)S Sy) |)

" (ea = —~=i) etl! se

0 a, ae ©

] : - my : rt n :

” a iy r e a ed

: f : f a) . : i : rom

a : 7 : = 7 ; A R

wy 5

; - mn i ee) ot ' : y i

7 i n i

, - Oy

~ " u v i “

= z oe. ae

? i sy ,

i A ry ci

: z : ,

e '

\ are

. i

- ty

‘

“

Pk 3

———

= _——— a

s .

tte! 2

7 cs

: |

i i F a i) i

y oa Oe ihh J

=) 3 i

‘ = 4 . Z

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

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

Te SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

eye the Figure depicting C. namacolus ...’:“. ... nm /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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

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

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.