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. ANNALS OF THE

SOUTH AFRICAN MUSEUM

VOLUME 98

ANNALE VAN DIE

SUID-AFRIKAANSE MUSEUM

BAND 98

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ANN Mov A AY ay

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aan

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 98 BAND

0 NOVI ss

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1987-1989

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA

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

D2109

LIST OF CONTENTS

Page

ARNAUD, F. & CHILD, C. A.

The South African Museum’s Meiring Naude cruises. Part 17. Pycnogonida. (Pub-

lishedWPecemberlO Sse rer Seater or ey Cee eile an Ai aas lets mpd ret ae Al

Bircu, G. F. see DINGLE, R. V. et al.

BISESWAR, R.

Ochetostoma (Echiura) from southern Africa with a description of a new species.

(RitblishediMayslOSS s\n 1 eee eine nant oh on ane iee eek cas ou 29

BREMNER, J. M. see DINGLE, R. V. et al.

CuILD, C. A. see ARNAUD, F.

De Decker, R. H. see DINGLE, R. V. et al.

DINGLE, R. V. see WHATLEY, R. C.

DINGLE, R. V., BircH, G. F., BREMNER, J. M., DE DECKER, R. H., Du PLessis, A., ENGEL-

BRECHT, J. C., FINCHAM, M. J., Firron, T., FLEMMING, B. W., GENTLE, R. I., GOODLAD, S. W.,

Martin, A. K., MILts, E. G., Morr, G. J., PARKER, R. J., RoBson, S. H., RoGeErs, J., SALMON,

D. A., SIESSER, W. G., Simpson, E. S. W., SUMMERHAYES, C. P., WESTALL, F., WINTER, A. &

WoopDBORNE, M. W.

Deep-sea sedimentary environments around southern Africa (south-east Atlantic

and south-west Indian oceans). (Published October 1987.) ................-.. 1

Du PLEssis, A. see DINGLE, R. V. et al.

ENGELBRECHT, J. C. see DINGLE, R. V. et al.

FINCHAM, M. J. see DINGLE, R. V. et al.

Fitton, T. see DINGLE, R. V. et al.

FLEMMING, B. W. see DINGLE, R. V. et al.

GENTLE, R. I. see DINGLE, R. V. et al.

GOoDLAD, S. W. see DINGLE, R. V. et al.

Ho .ruulis, L. B. see MANNING, R. B.

Hu. ey, P. A.

Lanternfishes of the southern Benguela region. Part 2. Gymnoscopelus (Gymno-

scopelus) bolini Andriashev in South African waters, with comments on the

distribution of Subantarctic myctophids in the eastern South Atlantic. (Pub-

hisheGbhcbnuanyslo SO) ipernity Nc eee ert des Seas s pa)

Huttey, P. A. & LUTJEHARMS, J. R. E.

Lanternfishes of the southern Benguela region. Part 3. The pseudoceanic-oceanic

interfaces (Rublished October 1989.) 5.) ae Mae onan pew el abe Se dries: 409

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

KLINGER, H. C.

The ammonite subfamily Labeceratinae Spath, 1925: systematics, phylogeny, dimor-

phism and distribution (with a description of a new species). (Published

1 RSL OVATEN AY. TOIL) Saresscial a ie ba a a eae 189

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

Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family

Placenticeratidae Hyatt, 1900; with comments on the systematic position of the

genus Hypengonoceras Spath, 1924. (Published November 1989.) ............ 241

LUTJEHARMS, J. R. E. see HuLLEy, P. A.

LIST OF CONTENTS (Cont)

Page

MANNING, R. B. & Ho .tuuis, L. B.

South African species of the genus Geryon (Crustacea, Decapoda, Geryonidae).

(Published: May 1988:) ¢. ccc 52.6. 08 ss se encuetnenet oere tee sets ala teeters pata eee V7

MartTIN, A. K. see DINGLE, R. V. et al.

MILLs, E. G. see DINGLE, R. V. et al.

Mor, G. J. see DINGLE, R. V. et al.

ParKER, R. J. see DINGLE, R. V. et al.

Rosson, S. H. see DINGLE, R. V. et al.

Rocers, J. see DINGLE, R. V. et al.

SALMON, D. A. see DINGLE, R. V. et al.

SIESSER, W. G. see DINGLE, R. V. et al.

SIMPSON, E. S. W. see DINGLE, R. V. et al.

SUMMERHAYES, C. P. see DINGLE, R. V. et al.

WESTALL, F. see DINGLE, R. V. et al.

WHATLEY, R. C. & DINGLE, R. V.

First record of an extant, sighted, shallow-water species of the genus Poseidonami-

cus Benson (Ostracoda) from the continental margin of south-western Africa.

(Published November 19890): och aa entrar se 437

WINTER, A. see DINGLE, R. V. et al.

WoopDBorRNE, M. W. see DINGLE, R. V. et al.

WOOLDRIDGE, T. H.

A new species of Mysidopsis (Mysidacea) from coastal waters of southern Africa

and a key to the known species from the subcontinent. (Published May 1988.) 93

ZEIDLER, W.

A redescription of Afrochiltonia capensis (K. H. Barnard, 1916) with a review of the

genera of the family Ceinidae (Crustacea, Amphipoda). (Published July 1988.).. 105

Volume 98 is complete in 11 parts.

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Saopalliencr Ami auducc © hil (eG SS renee ee cee Sera ee nie GLa eee ele a e-citie:

ME 98 PART 1 OCTOBER 1987 ISSN 0303-2515

IAN <1 1988

er a ipa

ANNALS

OF THE SOUTH AFRICAN |

MUSEUM

CAPE ‘TOWN

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Bu ttouGu, 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 ausgeftihrt in den Jahren

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

(continued inside back cover)

eee ee ww ee we we we wee ee a es ee es ee

Annals of the South African Huseua

Volume 78 Part 1 DINGLE, R.V. £F Al.

Flease note that Figures 1 and 3 of this part comprise two large maps. |

These have been mailed separately from the text (Maps packed in a cylindrical

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ANNALS OF THE SOUTH AFRICAN MUSEUM

| ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

October 1987 October

Part 1 Deel

DEEP-SEA SEDIMENTARY ENVIRONMENTS

AROUND SOUTHERN AFRICA (SOUTH-EAST

| ATLANTIC AND SOUTH-WEST INDIAN OCEANS)

fy OINGLE, G. F. BIRCH, J. M. BREMNER, R. H. DE DECKER,

Peeeeeeeessis, J. C. ENGELBRECHT, M. J. FINCHAM, T.

FITTON, B. W. FLEMMING, R. I. GENTLE, S. W. GOODLAD, A. K.

MeanrIN, E.G. MILLS, G. J. MOIR, R. J. PARKER, S. H. ROBSON,

J. ROGERS, D. A. SALMON, W. G. SIESSER, E. S. W. SIMPSON,

fae SUMMERHAYES, F. WESTALL, A. WINTER & M. W.

WOODBORNE.

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

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

OUT OF PRINT/UIT DRUK

ly AGES SO), 0), 2S 8 ta), SC, 5, FD),

Go CED), ©, NOD 7), 1OGES),

11(1-2, 5,7; t—pii.), 14(1-2), 15(425), 24); 27, 31-3), 226), 33. 26@) 45)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 090 7

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

DEEP-SEA SEDIMENTARY ENVIRONMENTS AROUND

SOUTHERN AFRICA (SOUTH-EAST ATLANTIC AND SOUTH-WEST

INDIAN OCEANS)

R. V. Drincie'*, G. F. Bircu?, J. M. BREMNER’, R. H. DE DECKER’,

A. pu Ptesssis?, J. C. ENGELBRECHT*, M. J. FincHAm®, T. Fitton’,

W. FLEMMING’, R. I. GENTLE’, S. W. GoopLap’, A. K. Martin®,

. G. Mitts’, G. J. Morr’®, R. J. Parker’’, S. H. Rosson’, J. RoGeErs’,

. A. SALMON?2, W. G. Sresser??, E. S. W. Smmpson', C. P. SUMMERHAYES"*,

F. WESTALL’?, A. WINTER’ & M. W. WoopDBoRNE’

(With 7 figures and 3 tables)

[MS accepted 30 April 1987]

Oo mw

ABSTRACT

Continental shelf sediment provenances, deep ocean basin sediment patterns, tectono-

sedimentary features, and the bathymetry of approximately four million square kilometres of the

SE Atlantic and SW Indian oceans are summarized in two charts at a scale of 1 : 3 200 000.

Contrasting modern deep-sea sedimentary environments around southern Africa are explained in

terms of regional oceanography and terrigenous sediment supply, and their spatial development

is traced through Cenozoic time. The linkage of major changes in oceanic circulation to sediment

production, injection, and dispersal is emphasized, including the case for a direct relationship

between Neogene Antarctic Bottom Water scour and large-scale allochthonism along the west-

coast continental margin.

CONTENTS

PAGE

MTA GLUES UMN a ree ee eT ee ye Se rs re eS Ae 2 2

DISCUS SIO errr res et rey Mocicae Neo GAs eran derals Wench setace WRN cay 4

SEcmmEM SOUT CE Stuer) evel anny dake feve ie chess hale Siete ie inne 4

Allochthonous environments and feeders .............. 8

Steady-state accretion andierosiom 35-4525 0..5 055. 06>-- 12

Wermicenous and hemipelasic 3-34-4955) - 6 ene 1

Pelagic carbonates and authigenic sediments ....... Uy)

SVNO SISter ia ee worn, Pancras vrai eon Naga at ake xwrallar & ¢ A,

NCKMOWICUSEMICMUS ah te his Ske hry e meee ah ie ane ta ee 23

INCGKERCTICE SIs dream ee oem ha AUN UA ka Meare aie aks eee ass 24

*= Marine Geoscience Unit, University of Cape Town, South Africa; *» = ESSO Australia, Box

4047, Sydney, Australia; *= Geological Survey of South Africa, Pretoria, South Africa;

*= National Research Institute for Oceanology, Box 32, Stellenbosch, South Africa; ° = 184

Lower Road, London SE16, UK; *=address unknown; * = Senckenberg Institut, D2940

Wilhelmshaven, West Germany; ®*=Old Mill Road, Penmaenmawr, LL34 6TN, UK;

°= SOEKOR, Parow, Cape Town, South Africa; 1° = EXXON Research Center, Houston, USA;

*1 = Department of Geology, University of Auckland, New Zealand; 1* = Amcoal, Box 1575,

Vereeniging 1930, South Africa; ** = Department of Geology, Vanderbilt University, Nashville,

USA; ** = BP Research Centre, Sunbury-on-Thames, UK; '* = Alfred Wegener Institut for Polar

Research, D2850 Bremerhaven, West Germany; * = deceased.

* = Present address: South African Museum, Cape Town, South Africa.

Ann. S. Afr. Mus. 98 (1), 1987: 1-27, 7 figs, 3 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

Since mid-Cenozoic time (Oligocene), sedimentation in the deep ocean

basins around southern Africa has been effected by three major processes:

(a) episodic injection of allochthonous material from the basin margins by a

combination of tectono-sedimentary mass movements and benthic currents;

(b) steady-state benthic accretion and erosion; and (c) steady-state pelagic

accretion (Table 1). This contribution describes the main ocean _ basin

sedimentary features around southern Africa, and interprets their distribution

and development in the context of these processes.

TABLE 1

Sedimentary processes, environments, and features

PROCESSES SEDIMENTARY ENVIRONMENTS ASSOCIATED FEATURES

Episodic

slope wasting and conti- (a) proximal basin: slope and slumps, canyons, chutes,

nental rise and ocean basin’ rise allochthonous structures lobes, fans

accretion and feeders

(b) distal basin: fill abyssal plains

Steady-state benthic

targeted accretion and ero-_ distal basin: zones of drift and drifts, waves, billows,

sion (incl. authigenic min- scour omission surfaces, Mn

eralization) pavement, basement out-

crops

Steady-state pelagic

accretion (incl. ice rafting) low energy proximal and distal drapes, billows

pelagic regions, especially on

ridges and plateaux

Figure 1 summarizes the sedimentary environments of the deep-sea basins

around southern Africa and has been compiled from numerous studies, but

particularly the regional surveys of Emery et al. (1975), Kolla et al. (1980), and

Tucholke & Embley (1984), more local accounts (e.g. Bornhold & Summerhayes

1977; Dingle et al. 1978; Summerhayes et al. 1979; Dingle & Camden-Smith 1979;

Embley & Morley 1980; Westall 1984; Martin 1984; Goodlad 1986), and the

continental shelf sediment maps produced by Birch et al. (in press) and Bremner

Fig. 1 (see accompanying map). Deep-sea sedimentary environments around southern Africa

(south-east Atlantic and south-west Indian oceans). Compiled from published and unpublished

data: Bang (1968); Birch (1975, 1980); Birch er al. (in press); Bornhold & Summerhayes (1977);

Bremner (1978); Bremner et al. (in press); Dingle (1977, 1980); Dingle & Camden-Smith (1979);

Dingle et al. (1978); Dingle & Robson (1985); Embley & Morley (1980); Emery et al. (1975);

Flemming (1978, 1981); Goodlad (1986); Kolla et al. (1980); Martin (1984); Robson & Dingle

(1986); Rogers (1977, 1986); Shannon (1985); Summerhayes et al. (1979); Tucholke & Carpenter

(1977); Tucholke & Embley (1984); Westall (1984).

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 3

et al. (in press). There are strong contrasts in the nature and style of sediments

between the south-east Atlantic and south-west Indian oceans, which result

primarily from a combination of two phenomena: climate (terrestrial and

oceanographic), and ocean basin physiography. Climate dictates the quantity and

relative proportions of terrigenous, biogenic, and authigenic material potentially

available for injection into the deep basins, and is controlled by the two quasi-

resident high-pressure atmospheric circulation cells (anticyclones) that lie on

either side of southern Africa. One over the south-west Indian Ocean and the

south-eastern seaboard of Africa supports a high-energy, relatively low-

productivity western boundary surface current (Agulhas Current) adjacent to the

upwellin perceoecce 4 Kin — Benguela Current

cells e AABW —-— Agulhas Current

— —-= NADW — --— Angola Current

Fig. 2. Oceanographic setting of southern Africa. Water-mass flow paths are schematic, and

based on numerous authors cited in the text. Coastal upwelling sites are after Shannon (1985).

Abbreviations: rivers—KR = Kunene, LR = Limpopo, OR = Orange, TR = Tugela, ZR =

Zambezi; physiographic features—AB = Agulhas Basin, ABK = Agulhas Bank, AP = Agulhas

Plateau, AR = Agulhas Ridge, CB = Cape Basin, MB = Mozambique Basin, MR =

Mozambique Ridge, NV = Natal Valley, TB = Transkei Basin, WR = Walvis Ridge.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

humid hinterland of south-east Africa, while the other over the south-east

Atlantic is associated with a low-velocity, high-productivity eastern boundary

surface current (Benguela Current) adjacent to the arid hinterland of south-west

Africa (Fig. 2). The former sustains shelf sedimentary environments charac-

terized by high terrigenous input and high-energy benthic boundary conditions;

the latter sustains low terrigenous input, low-energy benthic boundary conditions.

Complex sea-floor morphology in the south-west Indian Ocean has its origins

in a more complicated plate tectonic history than the less cluttered south-east

Atlantic Ocean. This, coupled with a high sediment accumulation rate results in

physiographic differences between the two basins that lead to the partitioning, at

about 34,5°S in the Natal Valley, of the two deep current regimes that effect the

dispersal and/or erosion of bottom sediments in the area (Antarctic Bottom

Water, AABW;; and North Atlantic Deep Water, NADW: Figs 2, 3). The results

are gradients in sediment thickness and sea-floor current velocity, which are

generally high in the south-west Indian and low in the south-east Atlantic basins.

Finally, it will be suggested that the foregoing pattern of deep-sea

sedimentary environments has persisted at least for most of late Cenozoic time

(i.e. Neogene and Quaternary, 24,6 m.y.), although the effectiveness of the

various processes controlling sedimentation has fluctuated.

DISCUSSION

SEDIMENT SOURCES

The single most significant factor in understanding the differences shown in

Figure 1 between the eastern Cape—northern Agulhas basins and the Natal

Valley—Transkei Basin is an appreciation of the amount of terrigenous sediment

that they receive. The former area is sediment-starved, whereas the latter is not

(Embley & Morley 1980; Tucholke & Embley 1984), and the reason for this is

vividly illustrated by considering the proportion of modern terrigenous input

supplied to the southern African continental shelves via rivers: west coast, 4,1 per

cent; south coast, 17,9 per cent; east coast, 78,0 per cent (Table 2a, Fig. 4).

These figures become even more skewed when the fate of the detritus is

considered. On the west coast (south of 22°S) only three rivers are perennial and

contribute to the shelf sediment flux (Orange, Olifants and Berg), and it is

suspected that only a small proportion of their load crosses the shelf. Large

quantities of mud are trapped in a nearshore strip (Fig. 1) (Birch 1975; Rogers

1977; Bremner 1978), although small quantities of clay-sized detritus are

entrained in the Benguela Current and carried from the Orange River at least as

far north as the eastern end of the Walvis Ridge (Diester-Haass & Rothe 1987).

Fig. 3 (see accompanying map). Bathymetry around southern Africa (south-east Atlantic and

south-west Indian oceans). Compiled from: Dingle (1977, 1980, 1986); Dingle er al. (1977, 1978);

Dingle & Camden-Smith (1979); Kolla et al. (1980); Rabinowitz et al. (1980); Simpson (1982);

Tucholke & Embley (1984).

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 5

TABLE 2

Terrigenous sediment input

(a) MODERN RIVER INPUT (X 10° m*/yr)

Input Percentage End point

west coast: Orange, 6,52 41 Cape Basin

Olifants, Berg 0,5°

south coast (to Cape Padrone) 30,598 17,9 Transkei Basin,

Agulhas Basin

east coast (C. Padrone to Limpopo) 133,0” 78,0 Natal Valley

Tou 170,59

(b) AEOLIAN INPUT

West coast (18-30°S)—perhaps as much as 6,5 x 10° m? per large katabatic event (‘berg wind’)

(see Shannon & Anderson 1982; Chapman & Shannon 1985).

Vol* Area* Rate” Rate/area*

(km?) (km?)

Orange Basin® 15 14 2,3 | 16

Natal Valley‘ 32 21 4,9 23

“es Se oS nem § = mela

@ = Milliman & Meade (1983) ‘ = recalculated from Dingle et al. (1983)

“= Flemming & Hay (1983) * = Dingle & Hendey (1984)

$ = A. K. Martin (personal data)

Sand-sized particles (mostly from the Orange, which contributes 93% of the total

sediment input) move northwards by longshore drift and are blown onshore into

the Namib Desert (Rogers 1977). In a similar fashion, along the south coast,

muds are trapped in nearshore belts and some sand at least is moved along shore

and incorporated into small coastal dune fields (Birch 1980).

In addition to fluvial input, extensive aerosol plumes of fine sediment are

periodically swept off the continental interior and injected for up to 200 km

across the west-coast continental shelf (Figs 1, 4). These are generated by strong

katabatic winds (up to 15 m/sec) from high-pressure systems over southern

Africa, which force hot, dry air across the arid coastal plains (‘berg winds’:

Shannon & Anderson 1982; Shannon 1985; Chapman & Shannon 1985). There

are no quantitative data on the volumes of sediment transported during—nor on

the frequency of—these berg winds, but Shannon & Anderson (1982) have

suggested that one large katabatic event operating between 18° and 28°S may

involve as much particulate material as the annual sediment input from the

Orange River. South of 23°S there are three zones in which large south-westerly

trending plumes have been consistently identified: the largest is centred on the

Orange River valley; the second between 10 and 20 km north of Liideritz; and the

third in the vicinity of Walvis Bay. These zones are thought to coincide with

topographic depressions that focus the winds (Shannon & Anderson 1982).

Deposition appears to be confined to the continental shelf and, in comparison

with fluvial input, is well dispersed, fine grained, and unlikely to be lost to the

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

=— =

\ --s

Mag

-——

— =

—e-—”

run off =7.0

4.1% total

B _-’ run off = 30.59,

17.9% total 3

Cyast— 16

A 7 7

ey Pa 7

os Z 7

py = A gulhas 7

SF Plateau of

—~_as as == Fracture zone

————3km 3km_ isobath

ra terior cg se Extent of drainage area

Area of major aerosols

Fig. 4. Drainage areas and fluvial sediment yields in southern Africa.

1 = between Kunene and Orange rivers—no significant runoff; 2 = west coast: Orange, Olifants

and Berg rivers; 3 = southern Cape; 4 = east coast, including Tugela and Limpopo rivers. Runoff

yield is in X 10° m*/yr.

Abbreviations: c.a.r. = Cenozoic sediment accumulation rate/area within the 0,5 km isopachyte

in m*/km?/yr; AFFZ = Agulhas—Falkland Fracture Zone; MFR = marginal fracture ridge.

Rivers—K = Kunene, OR = Orange, OL = Olifants, B = Berg, T = Tugela, L = Limpopo.

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 7

marine environment by transport back onshore (although entry into food and

chemical cycles will reduce the flux to the sea-floor). Depending on the frequency

of these winds (and the veracity of Shannon & Anderson’s 1982 suggestion), the

total annual terrigenous supply to the west-coast shelf is realistically likely to

fluctuate from 7,0 x 10° m® (no katabatic events) to 20,0 x 10° m* (two events).

Diester-Haass et al. (1986) and Diester-Haass & Rothe (1987) have shown

that during Neogene glacial, low sea-level episodes, there were increases of

between two and six times the non-glacial rates in the terrigenous sediment flux to

the eastern Walvis Ridge. This was probably achieved through a combination of

aeolian and Benguela Current activity, and included the injection of shelf-derived

glauconite and phosphorite grains.

The foregoing situations are in strong contrast to those off the east coast,

where the very much greater fluvial terrigenous input is available for injection

into the Natal Valley. Sands are entrained in the Agulhas Current (which reaches

velocities above 250 cm/sec—Flemming 1978; Flemming & Hay in press), and

together with relict Pleistocene carbonate gravel lags, form large underwater

dunes that leave the shelf via at least four canyon-head complexes (Flemming

1981; Hay 1984; Dingle & Robson 1985). The precise fate of the east-coast mud is

not known, but little or none resides on the shelf (Fig. 1). Presumably it all passes

into the deep-sea environment (probably via the canyon systems), but the route

appears to involve an initial phase in surface waters in the inshore shear zone of

the Agulhas Current before the muds cross the shelf (Walters & Schumann 1985).

Modern terrigenous supply is grossly inflated by agricultural malpractices (e.g.

Dingle et al. 1983; Dingle & Hendey 1984), and may distort predictions of

regional trends, but calculations based on sediment basin volumes indicate a

mean Cenozoic accumulation ratio for east coast : west coast of 2: 1, at rates/

area of 23 and 16 m*/km/?/yr, respectively (Table 2c, Fig. 4).

These figures raise the question of possible variations in either rate of

sediment supply, or provenance during Cenozoic time, because they contrast

strongly with the ratio of modern terrigenous input for the east coast : west coast

of at least 10 : 1 (depending on the frequency of west-coast berg winds). If the

east-coast terrigenous supply remained constant through Cenozoic time, then

alternative sources of sediment must have made up the shortfall. Candidates

include large increases in authigenic (i.e. phosphatic, ferro-manganese, glauco-

nitic minerals) and biogenic (carbonates and opaline silicia) components, and

possibly ice-rafted detritus. With our present data, which are essentially two-

dimensional, it is not possible to assess the volumes of these individual

components, but we suspect that the problem of a significant shortfall may be

more apparent than real. Dingle & Hendey (1984) briefly addressed the problem

of variation in mean annual sediment input rates off the west coast (off area 2 in

Fig. 4), and concluded that there has been a drastic reduction from Upper

Cretaceous, through Palaeogene to Neogene (excluding modern) time:

16 : 3: 0,5 x 10° m®. We do not have similar figures for the east coast area, but if

Neogene rates there are similar to the mean Cenozoic rates (there was no

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

comparable increase in the aridity in this region), then the Neogene east : west

coast ratios of rates/area would be of the order of 7 : 1. Clearly, this is an aspect

of the regional sediment budget that warrants further study.

These terrigenous supply rates reflect arid hinterlands for the west-coast

rivers, and humid hinterlands along the south- and east-coast mountain ranges,

which in turn are sustained by the atmospheric and oceanic circulation adjacent to

the coasts (Fig. 2). Along most of the west coast intense upwelling of cold,

nutrient-rich water is associated with the Benguela Current system (see Shannon

1985; Chapman & Shannon 1985; Shannon & Pillar 1986, for extensive reviews of

physical, chemical, and biological aspects of the Benguela Current). This has

been a feature of the oceanic climate since at least Upper Miocene times (Siesser

1978; Diester-Haass & Rothe 1987) and, coupled with the low terrigenous

dilution rate, has promoted extensive production of authigenic sediment

(glauconite and carbonate apatite), pelagic carbonate sands, and more locally

diatomaceous muds (Fig. 1). Although evidence for modern production of

phosphatic sediments is limited to the Walvis Bay area (e.g. Bremner 1980),

Mio—Plio—Pleistocene deposits occur as far south as 37°S and across the Agulhas

Bank to the southern part of the east-coast sand stream (28°E). Pleistocene

sediments on most of the Agulhas Bank consist of relict quartz and benthic

carbonate sands (Birch et al. in press).

A recently described feature of modern sedimentation on the south and east

coasts is the development of prograding ‘submerged spit-bars’ (Martin &

Flemming 1986). Along the east coast these form where the large transverse

Agulhas Current and counter-current-driven dunes converge with littoral drift

paths, whereas off the south coast similar structures form where swell-driven

littoral sediments moving eastward cannot bypass capes. Large volumes of sand

have been incorporated in these structures (for example, individual submerged

spit-bars on the south coast contain up to 5 km*), which themselves may form the

boundaries of inshore sediment traps.

ALLOCHTHONOUS ENVIRONMENTS AND FEEDERS

Episodic injection of sediment into the deep ocean basins around southern

Africa has occurred on a massive scale during later Cenozoic time, and Table 3

summarizes the main sediment types available for injection into the ocean basins

from the shelf source areas. On the west coast this has primarily been in the form

of slumps and slides, whereas off the east coast a wide variety of mass-wasting and

accretionary mechanisms and processes has operated—e.g. slumps, slides, debris

flows, and turbidity currents. These differences are a direct result of the contrasts

in the low energy-low terrigenous input versus high energy-high terrigenous input

shelf regimes of the two coasts, respectively.

Fight discrete allochthonous masses have been identified between the Walvis

Ridge and the southern tip of the Agulhas Bank (Summerhayes et al. 1979;

Dingle 1980, 1983). These affect an area of about 260 000 km?, and fall into three

categories: north of 27°S; peripheral to the main Orange Shelf depocentre; and

south of the Cape Canyon.

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 9

TABLE 3

Shelf sediment types and minerals available for injection into adjacent ocean basins

Sediments/minerals Potential destination

West coast glauconite CAPE BASIN

phosphorite

pelagic carbonate sand

opal

fine terrigenous debris

South coast glauconite N. AGULHAS BASIN

phosphorite AGULHAS PASSAGE

relict quartz and carbonate TRANSKEI BASIN

sand (including oncolites) EXTREME S. NATAL VALLEY

East coast terrigenous sands NATAL VALLEY

terrigenous muds TRANSKEI BASIN

relict carbonate sands (including

oncolites)

(a) Three thin (up to 75 m) slides lie off the Walvis shelf. In this region the slope

is smooth, regular and canyon-free. The slides’ head regions lie in about 1 900 m

water depths.

(b) A major slump (up to 720 m thick) lies on the north-west face of the Orange

Shelf (Chamais Slump). There is an extensive fissured head zone (lying between

200-1 500 m water depth) and locally irregular sea-floor, but no evidence of

canyons. The Orange River probably debouched directly into this region during

Neogene and Quaternary low sea-levels (Dingle & Hendey 1984), and the slump

head structures locally intersect the zone of authigenic sediments on the outer

shelf. At least two, thinner, and tectonically less-complex slides lie south of the

Tripp Seamount on the south-east lower face of the Orange Shelf. These occur

along a smooth, regular continental slope with no evidence of canyons. Their

head regions lie deeper than 2 km water depth, but there is a wide, sub-parallel

fissured zone that extends shallower than 1 km in the vicinity of Childs Bank,

where it may intersect the outer edge of the shelf authigenic sediment zone.

steep, rugged (probably canyoned) slope adjacent to a relatively narrow

continental slope. The slumps are thick (up to 720 m) but relatively narrow and

tectonically complex. Large glide-plane scars incise the slope, and are deflected

across the margin when they intersect the Cape Canyon, whose thalweg is littered

with detached blocks where it crosses the lower slope (Dingle 1986). The head

region of the canyon and associated glide planes intersect a major shelf zone of

authigenic sediments west of Cape Town. Tucholke & Embley (1984) show that

the outer edge of this slumped sector has been scoured by Cape Basin AABW

flow.

All the west-coast allochthonous zones (with the exception of the

southernmost part of the Cape Town sector) have wide distal lobes that are

thought to represent more mobile debris-like deposits associated with the main

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

slide activity. None (with the notable exception of the Cape Canyon) appear to

have feeders or chutes, presumably because there is either insufficient sediment,

or no mechanism to divert significant quantities of shelf sediment on to the

continental slope. Significant slope wasting processes probably operated south of

the Cape Canyon immediately following slumping (creating the relatively rugged

slope topography), and may reactivate during low sea-levels when the small rivers

in the south-western corner of the continent discharge directly on to the outer

shelf.

In summary, on a regional scale, the heads of all the west-coast allochtho-

nous masses lie in zones of outer shelf—upper slope biogenic carbonate sands and

muds, and locally intersect areas of authigenic-rich sediments.

East-coast allochthonous features present a more complex picture than their

west-coast counterparts because they formed along a margin with a large, steady-

state sediment supply that has allowed them to progress to maturity (in terms of

slope wasting), and locally, to have produced large-scale continental rise

accretion. There are three sectors, each with distinctive structures: the south-

eastern side of the Agulhas Bank; 26°E to Durban; Tugela to Limpopo cones

(submarine fans).

(a) The whole south-eastern side of the Agulhas Bank, to water depths as

shallow as 200 m, has been involved in a long slump that causes a distinct

concavity in the bathymetry (Agulhas Slump: Dingle 1977). Its head region

contains a fissured zone and several large glide-plane scars that intersect Pliocene

limestones and outer-shelf patches of authigenic sediments.

Downslope movement was impeded by the Marginal Fracture Ridge (MFR)

that crops out intermittently along the outer margin (e.g. Robson & Dingle 1986),

so that the toe region of the slide failed to penetrate into the Agulhas Passage.

The ‘fresh’ state of sea-floor microtopography suggests that scour by the Agulhas

Current has prevented Holocene sedimentation over the slumped zone.

(b) The MFR can be traced to 35°S 26°E, north-east of which it did not form a

dam to downslope mass movement. Between this point and 31°S (just south of

Durban) the shelf is very narrow (c. 20 km) and locally rugged. (Flemming 1981

identified 85 canyon-like features along this sector.) For this area Figure 1 has

been compiled from Flemming (1981), Birch (1981), Dingle & Robson (1985),

and Robson & Dingle (1986), and shows that slope canyons are concentrated in

two complexes: in the north between Port Shepstone and Port St Johns, and in

the south between Kei River and Cape Padrone. Both these regions are loci of

slumping and large-scale mass wasting of the slope (Dingle & Robson 1985), and

are centres through which large quantities of detritus are siphoned from the shelf

and injected into the Natal Valley. Flemming (1981) recognized their role as exits

via which the Agulhas Current-driven shelf dune fields passed on to the slope, and

Dingle & Robson (1985) compared the sequence of slumping, canyon formation,

slope wasting and thalweg incision with that described by Farre et al. (1983) from

the New England slope. Large coalescing continental rise lobes emanate from

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 11

these two centres, and are crossed by canyon extensions and debris flow chutes

that spread aprons of allochthonous material over much of the southern Natal

Valley floor. In the extreme south, the canyon/glide-plane heads intersect

authigenic mineral-rich sediments as far north as the Fish River (c. 33°30'S), but

otherwise the bulk of the injected shelf debris is derived from the river-borne

terrigenous input and the carbonate lag gravels on the outer shelf (Flemming

1978, 1981; Hay 1984; Flemming & Hay in press). The southern edge of the main

apron is dammed against the east-west trending Agulhas Drift that forms the

northern edge of the Transkei Basin. However, the distal lithofacies of this

allochthonous input probably penetrates into the Transkei Basin, where it forms

small abyssal plains and infilled channels floored by turbidites (Westall 1984).

rivers, whose modern contributions are 5,9 and 65,0 x 10° m*/yr, respectively

(53,3% total east-coast supply—Flemming & Hay 1983). Both rivers have

constructed large submarine fans that prograde across the continental shelf into

deep water and, although each has a history that extends into the Cretaceous,

they were both active in Neogene and Quaternary time (Dingle et al. 1978;

Martin 1984; Goodlad 1986). This has been achieved through the sheer volume of

supply, and the fact that both rivers discharge south-west of coastal offsets that

create semi-permanent clockwise gyres inshore of the Agulhas Current (Flem-

ming 1981). Small pockets of muddy sediment occur in these gyres (e.g. Martin

1984; Felhaber 1985). It is significant that other east-coast rivers that have large

modern sediment yields have not constructed similar submarine fans (e.g. Fish:

13,5; Kei: 9,2; and Mzimvubu: 5,6 x 10° m*/yr—Flemming & Hay 1983). This

must be attributed either to unsuitable coastal geometry (which permits the

Agulhas Current to disperse sediment rapidly), or to the fact that the high yields

merely reflect modern agricultural practice. A further constructional feature lies

to the east of the Limpopo Cone, where localized upwelling of the Agulhas

Current has generated a small area of authigenic sediments on the Inharrime

Terrace (Dingle et al. 1978).

Canyons are known from two areas: the Tugela Cone and the Zululand

coast. The two large canyons on the Tugela Cone probably represent alternative

pathways for Tugela River and shelf sediments to the Natal Valley (Goodlad

1986), and are comparable in size to the Cape Canyon, although neither appears

to be structurally controlled. The southernmost (Tugela Canyon) debouches on

to the deep sea-floor on the southern flank of the cone, and may generate a chute

system that connects with that emanating from the Port St Johns canyons,

whereas the Goodlad Canyon runs north-east and links with a system tapping the

Zululand shelf. Several small deep canyons cross this very narrow (less than

5 km) shelf and head almost to the modern coast (Bang 1968; Flemming 1981;

Martin 1984), where they were presumably linked to rivers of the Zululand

coastal plain during low sea-level stands. Flemming (1981) reported submarine

dunes on this narrow shelf, which acts as a zone of bedload parting related to the

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

shelf gyre off Maputo. He postulated that the canyons siphon off the shelf load,

which must ultimately move down the broad valley on the western side of the

Central Terrace and link with the northern Tugela Cone input.

Martin (1984) has described a slump complex along the steep Zululand slope,

which in the north appears to have encroached on to the Central Terrace. Its

relationship with the southern end of the Limpopo Cone is unclear. Other smaller

slumps have been partially mapped off Maputo, and the Inharrime and Central

terraces (Martin 1984). Also in the northern part of the Natal Valley, the distal

edge of the Limpopo Cone coincides with the Almirante Leite Ridge and the

western pathway of the Agulhas Current scour zone. There are two main

branches to the current in this area (Fig. 1) (Martin 1981, 1984; Martin et al.

1982), which between them have kept the Central Terrace a region of low or zero

sedimentation through most of Neogene time.

STEADY-STATE ACCRETION AND EROSION

Terrigenous and hemipelagic

Dynamic, steady-state sedimentation on the ocean floors around southern

Africa is principally effected by two surface currents (Benguela and Agulhas),

and two deep-water masses (AABW and NADW) (Fig. 2). Of these, the

Benguela plays a major role as a sediment generator but, in areas south of the

Walvis Ridge, a minimal role in sediment movement, because surface velocities

are generally not greater than 30 cm/sec at 25°S and 50 cm/sec at 33°S (Stander

1964; see Shannon 1985, fig. 17, for summary), and 5—10 cm/sec at 200-300 m

depth (Moroshkin ef al. 1970; see Shannon 1985, fig. 18, for trajectories). In

contrast, there is abundant documentation (cited below) that the other three

currents are major vehicles in transporting suspensate, and that during the course

of late Cenozoic time they have targeted erosion and deposition on a large scale.

Antarctic Bottom Water probably enters the area across sills in the

Atlantic-Indian mid-ocean ridge between 20° and 30°E before circulating

clockwise in the eastern and western parts of the Agulhas Basin (e.g. Tucholke &

Embley 1984). Because it is confined to water depths below about 4 km, its

effects are restricted to the Cape Basin, Agulhas Passage, perimeter of the

Agulhas Plateau, and Transkei and Mozambique basins. Tucholke & Embley

(1984) have predicted a clockwise circulation in the Cape Basin, with AABW

entering through the seamount province at the north-east end of the Agulhas

Ridge (Figs 1-3). Evidence of bottom sediment scour along the foot of the

northern Agulhas Ridge and the eastern Walvis Ridge, and the lower continental

rise Off south-western Africa, has been provided by numerous workers (e.g.

Connary 1972; Emery et al. 1975; Bornhold & Summerhayes 1977; Embley &

Morley 1980; Emery & Uchupi 1984; Siesser et al. in press) that supports the

hypothesis of Tucholke & Embley (1984) of a scour zone under the core of the

AABW mass. Data on benthic velocities are sparse, but some values have been

predicted from photographic, sedimentologic and oceanographic evidence: less

than 10 cm/sec (Wust 1955; Bornhold & Summerhayes 1977); 15 cm/sec

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 13

(Connary 1972). These are lower than required to erode semi-consolidated fine-

grained sediments (40-100 cm/sec— Flemming 1977), so that the existence of a

benthic nepheloid layer up to 1 km thick (Connary 1972) indicates that they are

not representative of episodic bursting velocities.

The scour zone associated with the AABW circulation is shown in Figure 1

along the lower part of the west-coast continental rise. It generally lies between 4

and 4,5 km, and whilst Tucholke & Embley (1984) suggest that it probably is not

actively eroding the sea-floor at present, the widespread phenomenon of

truncated bedding—as seen on seismic records—is unequivocal evidence for

large-scale erosion in the upper Cenozoic (e.g. Emery et al. 1975; Tucholke &

Embley 1984). Palaeogene sediments have been proved at the sea-floor in two

deep-water areas: at DSDP site 361 (4 549 m) within the scour zone south-west of

Cape Town they are over-compacted early Eocene chalks and oozes (Bolli et al.

1978); and in the scour moat on the northern side of the Agulhas Ridge (4 943 m)

they are muddy, glauconitic Oligocene sands (Siesser et al. in press). Tucholke &

Embley (1984) suggest that a short episode of intense erosion occurred in the late

Miocene, and that since then the ability of the AABW to sweep most of the new

input away from the core zone has maintained a dynamic equilibrium, with very

low sediment accumulation rates (0,4 m/m.y.—Embley & Morley 1980). If this is

the case, then the area shown on Figure | must represent the site of late Miocene

to Holocene omission surfaces (condensed sequences or hardgrounds; e.g.

Neogene to Quaternary ferro-manganese nodule pavements— Rogers 1986). In

fact, evidence suggests that, at least in the region off Cape Town, there has been a

later erosional event because the distal parts of the slumps in this area have been

eroded (Tucholke & Embley 1984). It follows from a dating of the slumps that

this younger AABW erosive episode was not older than Pliocene, and is possibly

as young as Pleistocene (Dingle 1980).

In the extreme south-east of the Cape Basin, Tucholke & Embley (1984)

predict that the AABW pathway leads through the seamount province at the

southern tip of Africa and into the Agulhas Passage. There have been numerous

reports of ferro-manganese nodules from this area (e.g. Tucholke & Embley

1984), and a recent photographic and sampling survey by Rogers (1986) shows

examples of fused nodule pavements, as well as partly buried nodule fields and

ripples in glauconitic sand. The latter observation suggests allochthonous input

from the adjacent shelf where feeders (e.g. Cape Canyon) have tapped the

authigenic shelf sediments.

AABW routes east of 20°E cannot be readily identified directly from

geological evidence because of the complex sea-floor morphology. Dingle &

Camden-Smith (1979) suggested that steady-state, relatively strong, westward-

flowing bottom currents were responsible for a narrow moat in the sediments

along the northern foot of the Agulhas Plateau. This developed across a strongly

asymmetric velocity gradient during late Cenozoic time (rather than by erosion),

but in the most recent phase of sedimentation the moat has been partially filled

with turbidites (Westall 1984). We link the latter event to the youngest period of

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

slope wasting and allochthonous injection from the margin of south-east Africa.

Tucholke & Embley (1984) suggest that modern AABW continues to follow a

similar path: entering along the northern side of the Agulhas Passage, circulating

clockwise in the Transkei Basin, and leaving along the southern side of the

Agulhas Passage. In contrast, Westall (1984) predicts that AABW moves north-

east into the Passage, but that it passes eastwards out of the Transkei Basin. In

support of this hypothesis, she cites evidence from sediment bedforms, and

current-meter data from the central and northern flanks of the passage (Dingle &

Camden-Smith 1979; Camden-Smith et al. 1981; Westall 1984). The current-

meter data of Camden-Smith et al. (1981), which indicated an average south-west

velocity of 22 cm/sec at a site close to the foot of the Agulhas Plateau, Westall

(1984) interprets as recording large eddies from the main AABW north-east flow.

Although there may be a small, irregular return flow of AABW along the

southern Agulhas Passage, we prefer Westall’s overall model, because the data

do not indicate a steady south-west current out of the passage, as required by

Tucholke & Embley’s (1984) interpretation. Erosional features along the western

flanks of the Agulhas Plateau (e.g. Tucholke & Carpenter 1977) can be attributed

to the most southerly portions of the AABW mass that do not enter the Agulhas

Passage but peel off southwards (Tucholke & Embley 1984). It is probable,

therefore, that a return AABW flowed along the northern Agulhas Plateau—

southern Agulhas Passage in late Tertiary and perhaps Pleistocene time, but

current-meter evidence does not suggest a similar modern regime.

Drapes of thin Quaternary sediment over Eocene and Miocene strata

(Salmon 1979), together with billows and elongate sediment drift, and numerous

basement outcrops attest to long-term, steady-state targeted erosion and

deposition in the Agulhas Passage. This is particularly so along its northern side,

where AABW velocities with a range 2—49 cm/sec and a north-north-east

progressive velocity vector have been measured at one site at the foot of the

continental slope (at 4 590 m—Camden-Smith et al. 1981). Along the northern

side of the Transkei Basin bathymetric contours shallower than 4,6 km swing

sharply east—west and partition the two deep-water masses that move through the

Agulhas Passage (Westall 1984); AABW is forced to its right, whilst the NADW

continues north-eastwards. The sudden change in direction of AABW, where it

encounters a rising sea-floor formed by the terrigenous apron from the east-coast

slumps and canyon feeders, originally targeted deposition along a narrow zone

and resulted in the formation of the 480 km long Agulhas Drift (Figs 1, 5) (E-W

Ridge’ of Westall 1984). Truncated bedding planes along its southern flank

suggest that modern AABW may be eroding the Agulhas Drift (Dingle &

Camden-Smith 1979).

A model for steady-state targeted sedimentation under the influence of the

NADW is presented in Figure 5. Bottom-water potential temperatures (Kolla

et al. 1980, fig. 7) indicate that along the continental slope and rise off south-east

Africa, NADW flows north-east beneath the south-west flowing Agulhas

Current. After eastward deflection around the southern face of the Tugela Cone

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS

“uIseg

raysuely = gL ‘AoqeA [RJWN =.LN ‘9Spry anbiquiezow = YW ‘ulseg onbiquiezoy] = gq ‘odessed seynsy = SdV ‘neared seujnsy = dV ‘Mosul ul suoneiAciqqy ‘(p86L) 1eISoM pue “(r86T) Agiquiq

R ax[OyoN], ‘(OS6T) ‘72 12 TOM {(986T) PEIPCOH <(SR6T) LOSGoY *F BUI ‘(6L61) HWS-uapuED 7% e[SuIq Aq poysi[qnd seep! pue vlep UO Ajieuntd poseg ‘MaVW Aq pepore Sutoq st Jo “Burjor00R

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16 ANNALS OF THE SOUTH AFRICAN MUSEUM

(which here is banked behind the buried South Tugela Ridge), further northward

progress of the NADW is stopped by the southern face of the Naude Ridge

(Goodlad 1986). This barrier rises from 2,5 to 1,8 km water depth, and deflects

the water mass southward down the eastern side of the Natal Valley along the

western flank of the Mozambique Ridge, at the southern end of which the AABW

and NADW masses reunite and flow eastwards and then northwards into the

Mozambique Basin (Kolla et al. 1980). Two long (at least 300 km) sediment drifts

occur in the eastern Natal Valley adjacent to the Mozambique Ridge, and both

indicate long-term, steady-state, targeted deposition (Dingle & Robson 1985).

We postulate that the East Transkei Drift forms from fine suspensate that has

circulated round the Natal Valley (much of it probably acquired from the canyon

feeders off East London, Port St Johns, Port Shepstone and the Tugela), with

deposition along the locus of the shear zone between the NADW core and eddies

developed against the irregular steep walls of the Mozambique Ridge. Fine

suspensate, carried eastwards across the Natal Valley by eddies peeling off from

the NADW and by turbidity currents discharging directly via the canyon feeders

on to the rise prism, are deposited along a second shear zone. This is located

where eastward moving bottom water interferes with the NADW moving down

the eastern Natal Valley. The locus of convergence of these suspensate-carrying

water masses coincides with the West Transkei Drift. Dingle & Robson (1985)

show that the initiation of the East Transkei Drift (in Layer A sediments) pre-

dates the West Transkei Drift (Layer B: Fig. 6). Southward-travelling turbidites

accumulate in the depression between the two drifts, with those in the east

probably having a Mozambique Ridge provenance.

A further consequence of the passage of NADW along the lower continental

margin of south-east Africa, together with the periodic eastward flushing of

channelled and sheet-flow suspensates, is seen in low (or negative) sedimentation

rates in the western part of the Natal Valley south of 31°S. Here, coccolith and

stable isotope data indicate sub-Recent sediments at the sea-floor, which were

deposited at water temperatures lower than those obtaining today (relatively low

percentage of Emeliania huxleyi at less than 20,5°C—Fincham & Winter 1986).

The West Transkei and Agulhas drifts can be traced to within 70 km of each

other, and it is possible that they join. Because the southern end of the former is

buried, however, it seems more likely that they both shrink in size and that the

region between them is the main pathway for Natal Valley turbidity currents into

the Transkei Basin.

Exit routes for AABW and NADW from the Natal Valley have not yet been

identified, but a narrow ‘barren erosional zone’ that has been mapped for at least

700 km between 4 and 5 km water depth along the foot of the Mozambique

Ridge, and the large sediment wave field adjacent to it have been attributed to

north-east flowing AABW (Kolla et al. 1980; Tucholke & Embley 1984). Bottom

water entering the Mozambique Basin from the Agulhas Basin is estimated to

contain about 10 wgm/litre of suspensate (Kolla et al. 1980) and, with the ensuing

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 17

low sediment accumulation rates, manganese nodules and thin sediment veneers

cover the sea-floor. North of 37°S resuspension of muds from Quaternary

(probably mostly Pleistocene) turbidites raises the suspensate concentration to

around 20 wgm/litre and a long 200—20 km wide zone of mud waves has formed.

The waves are northward migrating, 35-100 m high, and have a wavelength of

5 km. Kolla et al. (1980) estimate, from sea-floor photography, the AABW

velocity at 8-10 cm/sec. The ultimate provenance of the detritus is thought to be

the Zambezi River, and runoff from Madagascar.

Pelagic carbonates and authigenic sediments

Extensive areas of modern deep-sea pelagic carbonates are confined to the

crests of the Agulhas Plateau and the Mozambique Ridge. Both are topographi-

cally isolated from significant terrigenous input, and lie above the carbonate

lysocline. The Mozambique Ridge is covered by sands that have a carbonate

content that is generally greater than 75 per cent (Kolla et al. 1980) and are

composed of planktonic foraminifera, with smaller areas of pteropods (Vincent

1972; UCT unpublished data). Current control of this deposition is indicated by

large-scale billow-like bed forms, locally coarse grain textures, and manganese

nodules and encrustations that have been reported from the Ridge’s crest and

flanks south of 35°S (Vincent 1972; Summerhayes & Willis 1975; Tucholke &

Embley 1984; Robson 1985, pers. comm. 1985). Similarly, high carbonate values

have been reported from the northern Agulhas Plateau (66% — Westall 1984),

where large erosional and depositional bedforms have been identified from

seismic records (Emery eft al. 1975; Tucholke & Carpenter 1977; Dingle &

Camden-Smith 1979; Westall 1984), and manganese nodules and encrustations

have been widely reported (Summerhayes & Willis 1975; Tucholke & Embley

1984).

Rogers (1986) has shown that in the south-east Cape Basin, in the vicinity of

the Agulhas Ridge, ferro-manganese nodule pavements are locally draped with

modern sediment. In addition, the outer layers of nodules in the area have a

relatively high content of terrigenous detritus (including ice-rafted debris). This

evidence suggests that since some time in the Pleistocene, nodule growth has been

smothered by increased sediment supply caused by a combination of reduced

AABW scour and influx of ice-rafted detritus (Rogers 1986).

SYNOPSIS

Regional patterns of sedimentation during Neogene time in the south-east

Atlantic and south-west Indian oceans were, and are, controlled by the complex

interaction of large-scale allochthonous and synsedimentary processes, and

steady-state water-mass circulation. Fluctuations in their relative importance

have resulted in significant spatial and temporal changes in the Neogene

sedimentary record, so that, for instance, some features at present are either

dormant or relict (e.g. some ferro-manganese nodule fields, the Cape Canyon,

ANNALS OF THE SOUTH AFRICAN MUSEUM

us00q

ausd091|9

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Agulhas Current influence

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yualing seylnsy

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eene

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LSVd

uolyisodeap-uou 10 MO[S

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Ww aUadOIY

snoudsisi9} o1deoW

jUdTIND ejanduog

qouan|jul jUalIND ejensuag

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS

“SUOT}e}IO Beep

pue uoneurldxo 10j x0} 90g “JIC loysueI] ISOM = CLA ‘WUC leysuery isegq = GLA “Yu seynsy = Cy “s0vlay, JeNUa_ oy} Uo ATJeINedsa ‘uaiinD

sey[nsy 4} Jopun uonisodep-uou = ¢ ‘uoIsoID MEW YW 2U900)sSIa1q Aq POAOUII [BIID\VU = Z ‘UOISOIN MWY eUs.0IJ 218] Aq poAoulad eL9}eU = il

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“ulseg ToysueL] oY} 0} AaB A [eIeN OY) UMOP Joos] e UT somMeay smoys ASBOD 1Seq, OY} (q) ‘s}UsAo AreJUOUIpes poyeuTUOp oAeY AAGVY pure JuoIND

Pjonsuog oy) Yyorym UT a[Yord uIseq-0}-Jfays & smoys (III seUNSY ou} se see Ie} SB SoIN}e9J SOpNIoUI YOIYyM) .JSeOD 3soM, OY} (e) :sj7ed OM} UT SI WeISeIp

SUL “BOLSVW ULSYINOs punose sulseq UeId0 oY} UT seouaNbes AIejUSUNIpes UIapoul 0} Arena] -prw jedrourd Jo sdiysuonejas s1ydersnens onewayos -9 “S17

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EAST COAST

abundant terrigenous fluvial supply and biogenic debris, meagre authigenic input

WEST COAST

meagre terrigenous fluyial supply and authigenic & aeolian input

S &E N&W

non-=deposition south

slow or non=deposition pon fon

seas pee increased turbidite

\ b input

| SS

Wey J NS Neogene

Ys =) slumps

\ Fee

incision of E. coast

canyons

Benguela Current influence

K = onset of ‘modern’

Agulhas Current

————

———— ene

Eee sediments

onset of

Benguela Current

lees mid-4

EF Tertiary3{\\\p

ALES |) pee

i mid= Tertiary slumps of Zululand

= increased terrigenous supply:

onset of palaeo-Agulhas Clirrent

allochthonous major events

sediments

erosion surface

autochthonous e

sediments major slump events

drifts

sediment removed

non = deposition

Holocene =

B io)

Pleistocene 5

Pliocene eo

Miocene

Oligocene

Eocene

Vv ¥

Fig. 6. Schematic stratigraphic relationships of principal mid-Tertiary to modern sedimentary sequences in the ocean basins around southern Africa. The

diagram is in two parts: (a) the ‘West Coast’ (which includes features as far east as the Agulhas Drift) shows a shelf-to-basin profile in which the Benguela

Current and AABW have dominated sedimentary events; (b) the ‘East Coast’ shows features in a transect down the Natal Valley to the Transkei Basin,

where the Agulhas Current and NADW have dominated sedimentary events. Major erosion events commence at horizons marked by a thick line,

whereas the material removed by them (outlined by wavy lines) extends down in time, truncating earlier surfaces and features.

1 = material removed by late Miocene AABW erosion, 2 = material removed by Pleistocene AABW erosion, 3 = non-deposition under the Agulhas

Current, especially on the Central Terrace. AD = Agulhas Drift, ETD = East Transkei Drift, WID = West Transkei Drift. See text for explanation and

data citations.

WAASAW NVOIddV HLNOS FHL JO STIVNNV

SLNSWNOUWIANS AUVINAWIGSS VAS-dadd NVOIAVY NUYAHLNOS

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

west-coast slumps and slides, and the Agulhas Drift). Figure 6 is an attempt to

identify and date these changes and to relate them to environmental phenomena.

The key factor, which switches other processes on and off, appears to be

alterations in oceanic circulation.

Local mid-Tertiary glaciation of Antarctica was probably triggered by the

isolation of the continent at shallow water levels (Hag 1981). This resulted in the

creation of steady-state, cold, corrosive, bottom currents in the world ocean, and

the establishment of AABW pathways by earliest Oligocene time (33 m.y .—

Shackleton & Kennett 1975). Tucholke & Embley (1984) recognized an early

Oligocene erosion surface in the Cape Basin and on the flanks of the Agulhas

Plateau, and related this to AABW penetration into the south-east Atlantic.

Martin et al. (1982) and Martin (1984) date the initiation of the palaeo-Agulhas

Current to this period, and link the two phenomena. Martin & Flemming (in

press) argue that the establishment of this major surface current would have had a

significant effect on the climate of the hinterland, and it follows that the deep-

water sedimentary environments in the southern part of the Natal Valley would

reflect the change. The major lithologic change identified on seismic records

occurs between Layers A and B (Dingle & Camden-Smith 1979; Dingle &

Robson 1985). Westall (1984) dated this event (P.E. Horizon: Fig. 5) as

Palaeocene by analogy (but not direct correlation) with Horizon D in the south-

east Atlantic (Emery et al. 1975; Bolli et al. 1978), but in the absence of more

substantive evidence, the younger early Oligocene date suggested by the

reasoning of Martin & Flemming (in press) fits the available data more

comfortably. Consequently, Figure 6 shows earliest Oligocene sediments record-

ing the commencement of two sediment drifts: the Agulhas Drift (AABW), and

the West Transkei Drift (NADW or equivalent), which are related to the

initiation of the Agulhas Current and resultant increase in terrigenous input from

south-east Africa. The East Transkei Drift is older (Layer A), and we postulate

Fig. 7 (see facing page). Development of west coast allochthonous structures.

Key: X=intersection of erosion surface II and depositional surface I. Y = intersection of

erosion surface II and basal glide plane. Z = intersection of sea-floor and upper level of effective

scour (i.e. its shallowest depth) of the post-‘late Miocene’ AABW. I: Structures prior to the ‘late

Miocene’ erosion event. Sediment wedge to be removed by ‘late Miocene’ AABW erosion is

enveloped by the heavy dashed line. Arbitrary stratigraphic horizons (one is blacked) are used in

subsequent sketches to illustrate the developing structural and erosional relationships. II: Lower

slope support for outer margin sediments has been removed by ‘late Miocene’ AABW erosion.

X is the shallowest depth to which erosion operated. Erosion surface II (i.e. new sea-floor) is

marked by a heavy line. Sediment wedge to slump in phase III is enveloped by light dashed line.

This line is the incipient basal glide plane. Following erosion, the shallowest depth of effective

AABW scour is Z, upslope of which post-‘late Miocene’ sediment continues to accumulate

(upper thin blacked layer). III: Plio—Pleistocene slumping has translated slope sediments into

deeper water as rotated blocks. Point X over-rides point Y. Sediment wedge to be removed by

‘Pleistocene’ AABW erosion is enveloped by the heavy dotted line. IV: The outer edge of the

distal slump block is trimmed by ‘Pleistocene’ AABW erosion. Previous sea-floor configurations

are shown for comparison by light dot-dash lines.

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS

ee m= == trace of erosion surface ||

ecceeeeceeece trace of erosion surface lV

—— erosion surfaces (II & IV)

—-— —-s— incipient basal glide plane

— ee — original seafloor in phase |

—_—-—-—- original seafloor in phase II

—-:-—---=— original seafloor in phase II]

& Original seafloor

sediment deposited between

late Miocene erosion and

slumping in phase III

late Miocene

Plio- Pleistocene

Ill

Pleistocene

7p) ANNALS OF THE SOUTH AFRICAN MUSEUM

that it was sustained by detritus carried clockwise around the Natal Valley (in

contrast to predominantly across it: see Fig. 5).

Tucholke & Embley (1984) detected no major regional Upper Oligocene to

Upper Miocene hiatuses in the south-east Atlantic, but dated the initiation of the

modern deep-sea scour zones to massive ‘late Miocene’ erosion by AABW. There

is a certain amount of ambiguity in their dating (Tucholke & Embley 1984,

fig. 10), and it is unclear whether this is a composite erosional event covering

NHS5-NH7 (planktonic foram zones N16 to N18; late Miocene to earliest

Pliocene) or relates to one Neogene hiatus in particular. Keller & Barron (1983)

dated the three events as: NHS—9 m.y.; NH6—7 m.y.; and NH7—5 m.y.; the

youngest of which has recently been correlated with widespread AABW erosion

in the Atlantic sector of the Southern Ocean (Ledbetter & Ciesielski 1986). This

event (or events) stripped at least 500 m of middle and late Tertiary cover from

the continental rise and basin floor along the eastern Cape Basin (Bolli et al.

1978; Tucholke & Embley 1984; Siesser et al. in press). In the Southern Ocean,

widespread AABW erosion also occurred at 3,5—4 m.y. and 1,0—2,5 m.y. but,

according to Tucholke & Embley (1984), active erosion in the Cape Basin did not

persist significantly beyond Miocene time, since when it has maintained a wide

swathe of very slow sedimentation or non-deposition (Fig. 1). Off the east coast,

the late Miocene surge in AABW activity may have a slightly delayed action

analogue, where Martin etal. (1982) and Martin (1984) recognized the

establishment of the modern flow path of the Agulhas Current just after 5 m.y.

(i.e. earliest Pliocene). This resulted in active scouring of the Central Terrace,

and must have enhanced the Agulhas Current’s ability to transport detritus. One

effect of the invigorated AABW flow was to start to dismantle the Agulhas Drift,

which it had been constructing since early Oligocene time (Fig. 6).

At present we do not have an accurate timetable for the formation of the

massive allochthonous structures that occur on the outer continental margin

around southern Africa, but there does seem to be a synchroneity between east

and west coasts (Fig. 6). The youngest phase has been dated as Pleistocene and/or

Pliocene by Summerhayes et al. (1979), and by Dingle (1980) on the west coast,

and post-5 m.y. (i.e. Pliocene or younger) by Dingle (1977), Martin (1984), and

Dingle & Robson (1985) on the south and east coasts. In addition, ‘mid-Tertiary’

episodes have been postulated for the west coast (Dingle 1980), and off Zululand

(Martin 1984).

Because the outer edges of the allochthonous masses have been eroded by

the Cape Basin scour core (see seismic profiles in Tucholke & Embley 1984), it

follows that in this region at least, the slumps pre-date the youngest episode of

scouring. Tucholke & Embley (1984) consider this scouring to be ‘late Miocene’,

which is older than geological evidence suggests for the youngest slumping.

Figure 7 attempts to reconcile these dates and to incorporate the phenomenon of

scouring into a genetic model for the allochthonous masses. Stage I is pre-‘late

Miocene’ and shows the deep-water slab that the pulse of corrosive AABW

removed in ‘late Miocene’ time. In Stage II, support for mid-slope sediments has

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 23

been weakened by removal of this slab, whilst sedimentation continued into

Pliocene time in water depths shallower than point ‘Y’ (which represents a retreat

from the high point of erosion at ‘X’ following the cessation of active AABW

erosion). This additional loading during Stage II eventually triggered down-slope

rotation of blocks in Plio—Pleistocene time (Stage III). A post-Stage III

(?Pleistocene) episode of erosion by AABW is indicated by the trimmed outer

edges of some of the allochthonous units south-west of Cape Town (Stage IV).

This event may correlate with part of the 1,0-2,5 m.y. (Matuyama Chron)

AABW erosion event identified by Ledbetter & Ciesielski (1986) in the Southern

Ocean. Ancillary processes, which may have precipitated or facilitated alloch-

thonism, but whose relative effects cannot be assessed at this stage, possibly

include: base of slope sapping by ground water during periods of low sea-levels

(Oligocene, early Pliocene, and Pleistocene) (Robb 1984); seismic shocks; high

interstitial methane content in sediments (particularly along the west coast—

Emery et al. 1975; Summerhayes et al. 1979).

The ruggedness of the west-coast continental slope south of 34°S can

probably be attributed to small canyon and gulley formation on the slump-

affected slope that was accentuated by the discharge of rivers from the

mountainous Cape Town—Agulhas hinterland across the narrow coastal plain

during Pleistocene sea-level lows. This is a direct analogue to the narrow East

London margin and contrasts strongly with the non-dissected continental slope

farther north (where the coastal plains were much wider). Dingle & Hendey

(1984) date the initial cutting of the Cape Canyon as ‘mid-Tertiary’, when the

Oligocene Orange River discharged through the modern Olifants estuary. The

presence of a deep incision across the slope may explain the ‘focusing’ of Plio-—

Pleistocene slump glide planes along its lower eastern wall.

ACKNOWLEDGEMENTS

This compilation is based on work carried out in the Marine Geoscience Unit

at the University of Cape Town since 1967. Sea-time and research facilities were

funded by the University of Cape Town, Geological Survey of South Africa,

South African National Committee for Oceanographic Research, National

Research Institute for Oceanology, Sea Fisheries Research Institute, and the

Foundation for Research Development. Over the years, the officers and crew of

the University of Cape Town research vessel Thomas B. Davie bore the brunt of

the work at sea and we gratefully acknowledge their skill, dedication and

comradeship. We thank Judy Woodford for doing the cartography and also the

editorial staff of the South African Museum for guidance with the coloured maps.

Publication costs were generously met by the South African Museum, Foundation

for Research Development, and the Editorial Board of the University of Cape

Town.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Agulhas Passage, southwest Indian Ocean. Marine Geology 39 (3/4): M81—M88.

CHAPMAN, P. & SHANNON, L. V. 1985. The Benguela ecosystem. Part II. Chemistry and related

processes. In: BARNES, E. ed. Oceanography and Marine Biology 23: 183-251.

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thesis, Columbia University.

Diester-Haass, L. & RoOTHE, P. 1987. Plio—Pleistocene sedimentation on the Walvis Ridge,

SE Atlantic (DSDP Leg 75, site 532): influence of surface currents, carbonate dissolution

and climate. Marine Geology 77 (1/2): 53-85.

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HPC 532B (IPOD Leg 75) on the Walvis Ridge Terrace (SW Africa). Marine Geology 73

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DINGLE, R. V. 1977. The anatomy of a large submarine slump on a sheared continental margin

(SE Africa). Journal of the Geological Society of London 134 (3): 293-310.

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the continental slope and rise off southwestern Africa. Marine Geology 37 (3/4): 333-354.

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Petroleum Geologists. Studies in Geology Series 15: 1-24—-1-30.

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Geological Survey/University of Cape Town Marine Geoscience Unit 16: 20-25.

DiINnGLE, R. V. & CampeEN-SmiTH, F. 1979. Acoustic stratigraphy and current-generated

bedforms in deep ocean basins off southeastern Africa. Marine Geology 33 (3/4): 239-260.

DINGLE, R. V., Goopiapb, S. W. & Martin, A. K. 1978. Bathymetry and stratigraphy of the

northern Natal Valley (SW Indian Ocean): a preliminary account. Marine Geology 28 (1/2):

89-106.

DINGLE, R. V. & HENDEY, Q. B. 1984. Late Mesozoic and Tertiary sediment supply to the

eastern Cape Basin (SE Atlantic) and palaeo-drainage systems in southwestern Africa.

Marine Geology 56 (1/4): 13-26.

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS 25

DinGciE, R. V., Morr, G. J., BREMNER, J. M. & Rocers, J. 1977. Bathymetry of the

continental shelf off the Republic of South Africa and South West Africa. Geological

Survey of South Africa, Marine Geoscience Series Map 2. Pretoria.

DINGLE, R. V. & Rosson, S. 1985. Slumps, canyon and related features on the continental

margin off East London, SE Africa (SW Indian Ocean). Marine Geology 67 (1/2): 37-54.

DINGLE, R. V., SIESSER, W. G., & Newton, A. R. 1983. Mesozoic and Tertiary geology of

southern Africa. Rotterdam: Balkema.

EmBLEY, R. W. & Mor ey, J. J. 1980. Quaternary sedimentation and paleo-environmental

studies off Namibia (South-West Africa). Marine Geology 36 (3/4): 183-204.

Emery, K. O. & Ucuupl, E. 1984. The geology of the Atlantic Ocean. New York: Springer-

Verlag.

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margin off western Africa: Cape St Francis (South Africa) to Walvis Ridge (South West

Africa). Bulletin of the American Association of Petroleum Geologists 59 (1): 3-59.

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break: passage from youthful to mature phase in submarine canyon evolution. Special

Publication, Society of Economic Paleontologists and Mineralogists 33 (1): 25-39.

FELHABER, T. A. 1985. The sedimentology and mineralogy of Quaternary shelf sediments off

the Tugela River, southeast African continental margin. Technical Report. Joint Geological

Survey/University of Cape Town Marine Geoscience Unit 15: 230-247.

FincHaM, M. J. & WINTER, A. 1986. Agulhas Current imprint on oxygen isotopes of southern

Indian Ocean surface sediments. Technical Report. Joint Geological Survey/University of

Cape Town Marine Geoscience Unit 16: 208-216.

FLEMMING, B. W. 1977. Depositional processes in Saldanha Bay and Langebaan Lagoon.

Bulletin. Joint Geological Survey/University of Cape Town Marine Geoscience Unit 8: 1-215.

FLEMMING, B. W. 1978. Underwater sand dunes along the southeast African continental

margin—observations and implications. Marine Geology 26 (3/4): 177-198.

FLEMMING, B. W. 1981. Factors controlling shelf sediment dispersal along the southeast

African continental margin. Marine Geology 42 (1/4): 259-277.

FLEMMING, B. W. & Hay, E. R. 1983. On the bulk density of South African marine sands.

Technical Report. Joint Geological Survey/University of Cape Town Marine Geoscience Unit

14: 171-176.

FLEMMING, B. W. & Hay, E. R. In press. Sediment distribution and dynamics on the Natal

continental shelf. Jn: SCHUMANN, E. H. ed. Coastal ocean studies off Natal, South Africa.

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Indian Ocean). Bulletin. Joint Geological Survey/University of Cape Town Marine

Geoscience Unit 15: 1-415.

GRUNDLINGH, M. L. & LUTJEHARMS, J. R. E. 1979. Large scale flow patterns of the Agulhas

Current System. South African Journal of Science 75 (6): 269-270.

Haq, B. U. 1981. Paleogene paleoceanography: early Cenozoic oceans revisited. Oceanolo-

gica Acta 4 (supplement): 71-82.

Hay, E. R. 1984. Sediment dynamics on the continental shelf between Durban and Port St

Johns (southeast African continental margin). Bulletin. Joint Geological Survey/University

of Cape Town Marine Geoscience Unit 13: 1-238.

KELLER, G. & BARRON, J. A. 1983. Paleoceanographic implications of Miocene deep-sea

hiatuses. Bulletin of the Geological Society of America 94 (5): 590-613.

KOLLA, V., EITTREIM, S., SULLIVAN, L., Kostecki, J. J. & Burkie, L. H. 1980. Current-

controlled, abyssal microtopography and sedimentation in Mozambique Basin, southwest

Indian Ocean. Marine Geology 34 (3/4): 171-206.

LEDBETTER, M. T. & CrEsIELSKI, P. F. 1986. Post-Miocene disconformities and paleoceanogra-

phy in the Atlantic sector of the Southern Ocean. Palaeogeography, Palaeoclimatology,

Palaeoecology 52 (3/4): 185-214.

Martin, A. K. 1981. The influence of the Agulhas Current on the physiographic development

of the northernmost Natal Valley (SW Indian Ocean). Marine Geology 39 (3/4): 259-276.

Martin, A. K. 1984. Plate tectonic status and sedimenatry basin in-fill of the Natal Valley

(S.W. Indian Ocean). Bulletin. Joint Geological Survey/University of Cape Town Marine

Geoscience Unit 14: 1-209.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Martin, A. K. & FLEMMING, B. W. 1986. The Holocene shelf sediment wedge off the south

and east coast of South Africa. In: Knicut, R. J. & McLean, J. R. eds. Shelf sands and

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shelf and margin. In: SCHUMANN, E. H. ed. Coastal ocean studies off Natal, South Africa.

Lecture notes on coastal and estuarine studies 17. New York: Springer.

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northernmost Natal Valley, hiatus development and Agulhas Current palaeo-oceanography.

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MILLIMAN, J. D. & MEADE, R. H. 1983. World-wide delivery of river sediment to the oceans.

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southern Africa. Chart 526A. Cape Town: Department of Oceanography, University of

Cape Town.

Ross, J. M. 1984. Spring sapping on the lower continental slope offshore New Jersey. Geology

12 (5): 278-282.

Rosson, S. 1985. Ferromanganese nodules from the western flank of the Mozambique Ridge,

southwest Indian Ocean. Technical Report. Joint Geological Survey/University of Cape

Town Marine Geoscience Unit 15: 268-278.

Rosson, S. & DINGLE, R. V. 1986. A GLORIA traverse over the marginal fracture zone and

slump and canyon province off SE Africa. Marine and Petroleum Geology 3 (1): 31-36.

Rocers, J. 1977. Sedimentation on the continental margin off the Orange River and the Namib

Desert. Bulletin. Joint Geological Survey/University of Cape Town Marine Geoscience Unit

mle

Rocers, J. 1986. Seismic, bathymetric and photographic evidence of widespread erosion and a

manganese-nodule pavement along the continental rise of the southeast Cape Basin.

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16: 121-142. :

SALMON, D. A. 1979. Some Tertiary foraminifers in deep sea piston cores from the Central

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features and processes. Jn: BARNES, M. ed. Oceanography and Marine Biology 23:

105-182.

SHANNON, L. V. & ANDERSON, F. P. 1982. Applications of satellite ocean colour imagery in the

study of the Benguela Current system. South African Journal of Photogrammetry, Remote

Sensing and Cartography 13 (3): 153-169.

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on the continental slope and rise of southwest Africa: a reconnaissance study.

Marine Geology 31 (3/4): 265-277.

SOUTHERN AFRICAN DEEP-SEA SEDIMENTARY ENVIRONMENTS Dif

SUMMERHAYES, C. P. & WILLIS, J. P. 1975. Geochemistry of manganese deposits in relation to

environments on the sea floor around southern Africa. Marine Geology 18 (3): 159-173.

TUCHOLKE, B. E. & CarPENTER, G. B. 1977. Sediment distribution and Cenozoic

sedimentation patterns on the Agulhas Plateau. Bulletin of the Geological Society America

88 (9): 1337-1346.

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South Africa. Memoir. American Association Petroleum Geologists 36: 145-163.

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western Indian Ocean. Unpubl. Ph.D. thesis, University of Southern California.

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Ocean. Bulletin. Joint Geological Survey/University of Cape Town Marine Geoscience Unit

12: 1-276.

Wust, G. 1955. Stromgeschindigkeiten im Tiefen und Bodenwasser des Atlantischen Ozeans

auf Grund dynamischer Berechnung der Meteor-Profile der Deutschen Atlantischen

Expedition 1925/27. Deep-Sea Research 3 (supplement): 373-397.

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OCHETOSTOMA (ECHIURA) FROM

SOUTHERN AFRICA WITH A

DESCRIPTION OF A NEW SPECIES

R. BISESWAR

Cape Town Kaapstad

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OCHETOSTOMA (ECHIURA) FROM SOUTHERN AFRICA

WITH A DESCRIPTION OF A NEW SPECIES

R. BISESWAR

Zoology Department, University of Durban-Westville, Durban

and Zoology Department, University of Cape Town, Rondebosch

(With 23 figures and 1 table)

[MS accepted 17 August 1987]

ABSTRACT

In southern Africa, the genus Ochetostoma is currently represented by ten species:

Ochetostoma natalense is described as new; O. baronii, O. kempi, O. palense, O. decameron and

O. formosulum are redescribed and figured (the first four are new records for the region);

O. capense and O. arkati are briefly diagnosed; O. caudex and O. erythrogrammon, recorded

from several localities, are fairly well known. The status of one species from Park Rynie Beach,

on the Natal coast, remains to be resolved. A key for the identification of all the species in the

genus is provided, and the distribution of the southern African forms is briefly discussed.

CONTENTS

PAGE

intHOGUCH OME rete Sota ort enn ia ee seaeastcialine See aus 29

ID CSC UPC OMS pyre ce tae eet tin eA nines ek AS Ra oe, GAYS eG eG 30

IXKV 1O UNS SOLSES OUDGVAOMOTUE 0620000000 00b05000d0000006 30

PNCKMOWICCS SITES yar trys etd ceo seer eid tat ale ee ee lk ais, Bis 73

INCIETCIICCS Heyer cere eee eye eee nee male kn A ole aly a 1

PND DREVIAN OMS ne woe ene San yn wie Nee a ee a ae aes 1D

INTRODUCTION

At present, knowledge of the southern African echiuran fauna is rather

scanty. This is the fourth in a series of papers on the systematics and distribution

of the echiurans of Africa south of 20°S (Biseswar 1983, 1984, 1985). Biseswar

(1985) dealt with the genera and species of echiurans from southern Africa and

mapped their distribution, partly from the published works and partly from

surveys made along the coast.

The present paper is a comprehensive report on the species of Ochetostoma

from these waters. In their monograph, Stephen & Edmonds (1972) have

outlined some of the taxonomic problems encountered in the group. It is evident

from the literature that many of the species listed in this genus need re-

examination and redescription.

Collecting trips, specifically in search of echiuroids, were undertaken to

several localities along the Natal and Cape coasts. Most of the shallow-water

Ann. S. Afr. Mus. 98 (2), 1988: 29-75, 23 figs, 1 table.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

material used in this study was, however, obtained from the rocky shores of the

Natal coast. Some species were also obtained as a result of requests made to

several universities and institutions in South Africa. Unless otherwise stated in

the text specimens were collected by the author. Abbreviations used in the figures

are explained on p. 75.

Living specimens were photographed and then narcotized by adding small

quantities of magnesium sulphate to sea-water. Measurements of the proboscis

and trunk were taken after preservation in 10 per cent formalin. All the

specimens were subsequently dissected and, where possible, variations were

noted. Gross morphology was studied with the aid of a camera lucida.

DESCRIPTIONS

Ochetostoma Leuckart & Ruppell, 1828

Type-species: Ochetostoma erythrogrammon Leuckart & Rippell, 1828.

Diagnosis

Longitudinal muscle layer of body gathered into distinct bundles. Inner

oblique muscles between the longitudinal bands form fascicles. No anal or

posterior setae. Gonoducts are paired, ranging from one to seven pairs.

Gonostomal lips elongate and usually coiled.

Remarks

The genus is the largest of all the echiuran genera and comprises 24 species.

It was only after the publications of Spengel (1912) and Wharton (1913) that the

taxonomic importance of the condition of the oblique muscles was realized

(Stephen & Edmonds 1972). Unfortunately, some of the earlier descriptions

contain no information about the oblique muscles and hence the generic position

of a number of species that have been assigned to the genus Ochetostoma is

doubtful. They may belong to the genus Listriolobus. More detailed redescrip-

tions of some of the species, based on additional material, are required before

their taxonomic positions can be satisfactorily resolved.

Distribution

The genus Ochetostoma is very widely distributed in tropical and subtropical

waters of the Indian, Atlantic and Pacific oceans. The majority of the species is

confined to shallow waters of the intertidal zone but a few have been reported

from considerable depths. In their monograph Stephen & Edmonds (1972) gave

the distribution range of each species.

KEY TO THE SPECIES OF THE GENUS OCHETOSTOMA

The keys provided by Datta Gupta & Menon (1971) and Stephen &

Edmonds (1972) for the species of Ochetostoma have been modified to

accommodate O. natalense sp. nov. An attempt has been made to update the

existing keys in the light of additional information obtained from the redescrip-

ECHIURA FROM SOUTHERN AFRICA 31

tions of some of the species. In the present key, additional less rigorous characters

have been included to assist in the identification of single individuals. A

discussion of the structure and terminology of the group is included in a previous

paper (Biseswar 1983).

COMOAMCISM alse eres ek ene rey tre a hatte anes. ite foes 3s Z,

GonmOduciss pairs. Or lESSes 256 cee ae ee ee ee ee en es 3

First pair of gonoducts presetal; 12 longitudinal muscle bands ..............

3 tev Le lOReNE oT sree aan aera O. zanzibarense Stephen & Robertson, 1952

All gonoducts postsetal; 7 longitudinal muscle bands......................

Ee ee cig ea ee ne ee ante O. senegalense Stephen, 1960

GOMOGUCESPS MO AIS Meee Cee te a eae Ue an en MM Ca Baws a ah, Qe ned 4

Gonoducts: LOWE WEeMrpPaltSenc «soe. tase e ee Ee heed ee bee ele saws 5

First 3 pairs of gonoducts presetal; 19 longitudinal muscle bands............

WEE eae a a aie ee nee ans O. hornelli (Prashad, 1921)

First 2 pairs of gonoducts presetal; 10-11 longitudinal muscle bands.........

UA ee sana das % O. bombayense (Prashad & Awati, 1929)

GonoducisAipainss withsiisipait presetall 2. .-45 . 9a ate eres oes 6

GOnOduCtS:s OMMEWCR PAlNSins ethos. ee wee ue He wlne cih eee hee < Geb Mees 7

10 longitudinal muscle bands ........... O. decameron (Lanchester, 1905)

17-20 longitudinal muscle bands............... O. kempi (Prashad, 1919)

Conoducissipains only fisepainpresetalea. 4) 4eees- see e eae a o- 8

COnOdMcisy Pallsemm nine as see ee Se eas eed a weed 12

Proboscis slightly bifurcated with ventral and terminal rims crenated;

18 longitudinal muscle bands; colour of trunk pink .......................

5'i5 8 SRA US ee Se eae ae O. indosinense Wesenberg-Lund, 1939

Broboscismor bitincated ymanemsismootn 498.4447. 425005.459+650 45. 9

Small flat papillae confined to anterior and posterior surfaces of trunk; living

specimens red in colour; usually 12-13 longitudinal muscle bands, occa-

Sioned yal rear te. Wa Bisrsacs « Sovies.s su Mo's 6 ae O. australiense Edmonds, 1960

Papillae distributed over entire surface of trunk, much larger posteriorly... 10

. Alimentary canal short, about three times trunk length; 15 longitudinal

muscle bands; bright green imlife~...°..5.... 05... O. palense Ikeda, 1924

Alimentary canal long, about five times trunk length ................... iL

. Integument thin and transparent in middle region of trunk; 12—18 longitudi-

nal muscle bands .......... O. erythrogrammon Leuckart & Ruppell, 1828

Integument thick and opaque throughout; usually 16—22 longitudinal muscle

bands, very occasionally 14-16 ............... O. caudex (Lampert, 1883)

mmlinteniasalammUscleraDSemte ne ass as eh occ sete ne ee eee e lene ee 18)

ImntenbasaltmuscletpresemM@ame ae oe ee ota eee e wwe wee been 20

meaterabedgesiol proboscisimodified =). 62.4.8. 6Ge bere eos eee wee 14

Bateraliedges Onuproboscismotmodified 22. .5...4-....- 550252 sees: 17

. Gill-like processes along basal edges of proboscis; 8 longitudinal muscle

WEINGS on aco hs bia cle Cele Sle oe See O. arkati (Prashad, 1935)

ROVDOSCIS WITHOUT eIllelike StmUCtHURES. 56.60.46 o5¢eee0 1. 5sce- saree eee 15

dpa

Wa}é

ANNALS OF THE SOUTH AFRICAN MUSEUM

. Proboscis with dendritic outgrowths; 7 longitudinal muscle bands...........

BS Lies iors Arent ce O. septemyotum Datta Gupta, Menon & Johnson, 1963

Proboscis without dendritic outgrowths but proximal border with processes

with scalloped €dges ic... Sceds eee eo on sotto 16

. Oesophageal diverticulum present; 13 longitudinal muscle bands; trunk with

30-35 rings of large oblong papillae ..._ O. mercator Wesenberg-Lund, 1954

Oesophageal diverticulum absent; 12-13 longitudinal muscle bands; skin

smooth with only a few rings of papillae at posterior end of trunk ...........

Re eC Me a ty ar O. glaucum (Wesenberg-Lund, 1957)

. 8 longitudinal muscle bands; proboscis one-third trunk length; integument

transhicents 2 iets. 2 ee are ene eye ee ere O. octomyotum Fisher, 1946

Muscle bands 10'oF more... 6.3 6. ee es ee ee ee 18

. Trunk yellow in colour, covered with small uniformly distributed papillae;

10=1 muscle: bands... 32 eee O. hupferi (Fischer, 1895)

Trunk not yellow, papillae more densely packed at posteriorend ........ 19

. Trunk light green to bluish in colour, up to 25 mm long and transparent in

middle region; 13 longitudinal muscle bands.. O. pellucidum (Fischer, 1895)

Trunk reddish-purple, up to 94 mm in length; integument thick and opaque;

i7—20ilonentudinalimuscleibandsie saan ane eee O. natalense sp. nov.

. sMruscle bands less than 1003.30 I eee 21

Muscle bands more thamlQ ac a ee eee Dip

. Posterior end of trunk with a ring of elongate papillae; trunk up to 35 mm in

length; 7-8 muscle bands ............... O. formosulum (Lampert, 1883)

Posterior end of trunk without elongate papillae; trunk up to 120 mm in

length;7—S muscle bands o ae O. capense Jones & Stephen, 1955

Interbasal muscle extends over pharynx; trunk greyish-green; 14 longitudinal

muscle bands’... aeons eee O. manjuyodense (Ikeda, 1905)

Interbasalimuscle passes below planymxe....- 244250 oe oe eee 23

Proboscis readily deciduous; oblique muscles between longitudinal bands well

developed: 17-19 muscle bands) 3 5.4 O. baronii (Greeff, 1879)

Proboscis not readily deciduous; oblique muscles between longitudinal bands

weakly developed; 18-21 muscle bands........ O. myersae Edmonds, 1963

Ochetostoma natalense sp. nov.

Figs 1-4, Table 1

Material

Holotype, SAM—A21924, in the South African Museum, Cape Town. Adult

male, Park Rynie Beach, Natal (31°19’S 30°44’E); collected by K. S. Ganga,

18 October 1982.

Paratypes, SAM—A21925, in the South African Museum, Cape Town. Seven

specimens from Park Rynie Beach, Natal (31°19’S 30°44’E) and Isipingo Beach,

Natal (29°05’S 30°56’E); collected 1983, 1984.

ECHIURA FROM SOUTHERN AFRICA 33

Habitat

The specimens from Park Rynie Beach occurred in coarse sand under a

projecting ledge of rock facing the shore, while those from Isipingo Beach were

found under loose rocks in a rock tunnel. At both localities, the specimens were

found in the intertidal area, close to the high-water mark.

Description

Size. Trunk length (holotype) 80 mm, proboscis length 58 mm. Of the

paratypes, five sexually mature since gametes present in gonoducts. Trunk length

of sexually mature specimens ranges from 54 mm to 94 mm and proboscis from

30 mm to 58 mm (Table 1); hence proboscis ranges fron about one-third to

three-quarters of trunk length.

TABLE 1

Measurements (in mm) of preserved specimens of Ochetostoma natalense sp. nov.

Specimen Total Length* Trunk length Ratio

proboscis:trunk

*Holotype 138 80 0,73

_ Paratype 126 94 0,34

Paratype 120 85 0,41

*Paratype alat 71 0,56

*Paratype 110 69 0,59

Paratype 86 50 0,72

Paratype 82 47 0,74

*Paratype 69 54 0,56

* Trunk length and proboscis length

* Gametes present in gonoducts

Colour. In living specimens, proboscis is pale yellow. Trunk with reddish-

purple longitudinal stripes marking longitudinal muscle bands, interspaces

bluish-grey. Posterior extremity of trunk white, tinged with green in a few

specimens. In preserved specimens colour changes to pale pink.

External features

Proboscis. Proboscis fleshy, non-deciduous and of uniform diameter

throughout (Fig. 1A). Spatula-shaped in live animals but in preserved specimens

lateral edges curl inwards to form a tubular structure. Lateral margins of

proboscis smooth and free at base.

Trunk. Trunk sausage-shaped, more or less of uniform diameter (Fig. 1A).

Papillae minute, densely arranged over most of trunk except at posterior end

where they are very prominent, round to ovoid in shape. Size and distribution of

papillae uniform in all specimens. A white mucous cap encases posterior

extremity of trunk in formalin-preserved specimens. Entire integument thick and

opaque. Longitudinal muscle bands range from 17 to 20, inner oblique muscle

layer between longitudinal bands forming distinct fascicles. Ventral setae one

ANNALS OF THE SOUTH AFRICAN MUSEUM

A: TANG SE Bhan a nrane

Pea ROM Te

0,5 mm

10 mm

1 seta.

10na

A. Ventral aspect. B. Right funct

Fig. 1. Ochetostoma natalense sp. nov

ECHIURA FROM SOUTHERN AFRICA 35)

pair, located at anterior end of trunk, close behind mouth. Genital pores two

pairs, opening posterior to setae.

Setae. Each seta consists of a cylindrical shaft with curved terminal end

tapering towards tip (Fig. 1B). Much narrower distal third of seta golden-yellow

in colour while proximal two-thirds dark brown. Concentric markings present

mainly on middle two-thirds of shaft. Each seta is invested in connective tissue

and supported by radiating muscle strands as in other related species. There is no

interbasal muscle between setae.

Internal anatomy

Alimentary canal. Alimentary canal is considerably longer than trunk and

intricately coiled (Fig. 2). In holotype, numerous closely arranged mesenteric

strands fasten gut to body wall (Fig. 2). Mesenteric strands fewer in paratypes

and more sparsely arranged. Foregut comparatively small, terminating at ring

vessel. After ring vessel, intestine pursues complicated course through body

cavity, forming several ascending and descending limbs. A double sheet of

mesentery fastens oesophagus to anterior part of body wall.

s r int

10 mm

Fig. 2. Ochetostoma natalense sp. nov. Anterior part of the

alimentary canal, showing the mesenteric strands.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Intestine proper commences soon after ring vessel and is differentiated into

presiphonal, siphonal and postsiphonal regions. Presiphonal region marked by

presence of ciliated groove, which in holotype is about 36 mm long. Intestine

considerably dilated and extremely thin-walled in region of siphon, whereas in

postsiphonal region it is narrow with fairly thick walls. Last part of intestine of

uniform diameter, leading into more or less straight rectum. Small, spherical

rectal caecum present in all specimens.

Anal vesicles. Anal vesicles elongate, distensible tubes lying in coelom and

opening into rectum. In holotype, anal vesicles are abnormal (Fig. 3). Left anal

vesicle about three-quarters length of trunk, while right one about one-fifth as

long. In addition to these, a third anal vesicle present, which, instead of

terminating blindly, forms a loop and reopens into rectum (Fig. 3). All three

vesicles with numerous, tiny ciliated funnels. In paratypes, as in specimens of

related species, anal vesicles are normal, and are almost equal to or longer than

trunk. Numerous thin, mesenteric strands fasten proximal end of vesicle to body

wall, whereas distally they loosely interweave through coils of intestine.

Gonoducts. Two pairs of elongate, tubular gonoducts are located behind

setae; posterior pair much larger (Fig. 4A). In holotype, gonoducts are swollen at

base. Gonostomal lips elongate and spirally coiled. Opening of gonostome basal

in position. In sexually mature individuals, gonoducts considerably distended due

to presence of gametes.

Blood system. Intestinal ring sinus is an incomplete vascular ring located at

end of foregut (Fig. 4B). Neuro-intestinal vessel elongate, double for most of its

length. Dorsal vessel, after arising from ring sinus, continues anteriorly, dorsal to

oesophagus and pharynx, and enters proboscis.

Remarks

Distinctive features of the present species from southern Africa include the

number and location of the gonoducts in relation to the setae, the presence of

17—20 longitudinal muscle bands, the nature of the integument, the characteristic

shape and distribution of the dermal papillae, and the absence of an interbasal

muscle. The interbasal muscle is a useful taxonomic character in echiurans. Other

features of lesser significance are the colour and size of the animals and the ratio

of the proboscis length to trunk length.

In the genus Ochetostoma there are 13 species that possess two pairs of

gonoducts (see key p. 30).

The species O. arkati, described originally from Calcutta, India, differs from

O. natalense in several respects. In the former species the longitudinal muscles

are gathered into 7-8 bands and the proboscis is short, about a third to a quarter

of the trunk length. However, the most important distinguishing feature of

O. arkati is the presence of short, branched, gill-like processes along the edges of

the posterior half of the proboscis.

The species O. septemyotum differs from other known echiurans in the genus

in possessing a proboscis where the lateral margins are produced into a series of

ECHIURA FROM SOUTHERN AFRICA

lav

rec

10 mm

Fig. 3. Ochetostoma natalense sp. nov. Dorsal dissection of the

trunk showing the abnormal anal vesicles of the holotype.

ANNALS OF THE SOUTH AFRICAN MUSEUM

gon

10 mm

irs

10 mm

Fig. 4. Ochetostoma natalense sp. nov. Anterior end of the trunk

cavity. A. Gonoducts. B. Blood vessels.

ECHIURA FROM SOUTHERN AFRICA 39

folds with dendritic outgrowths that are bent inwards into the proboscis

groove.

Ochetostoma baronii, O. capense, O. myersae and O. formosulum are

distinct from O. natalense in possessing an interbasal muscle that passes through a

loop of the neuro-intestinal vessel. In addition, there are several other differences

in the proboscis, integument and dermal papillae that separate the above species

from O. natalense.

Ochetostoma hupferi from Nyango, West Africa, is based on a single

damaged specimen recorded from a depth of 11 m. In this species, the trunk is

small, light yellow in colour and covered with small, uniformly distributed

papillae. The proboscis is 5 mm in length and the ventral setae are large, golden-

yellow in colour. Fischer’s description (1895) mentions that the longitudinal

muscle layer is gathered into 10-11 bands. The most important differences

between O. hupferi and O. natalense lie in the number of longitudinal muscle

bands, in the size of the ventral setae and in the distribution of the dermal

papillae. Differences are also present in size as well as in the colour of live

animals.

The original description of O. manjuyodense is based on a single specimen

from Manjuyodi, Philippines, in which the proboscis was missing. Fisher (1948)

merely recorded this species from Hawaii. The trunk is 18 mm in length and has a

greyish-green tint. The dermal papillae are minute, almost invisible to the naked

eye, except at the posterior end where they are somewhat larger and arranged

almost in circular rows. A very distinctive feature in this species is the interbasal

muscle that extends over the pharynx. The longitudinal muscles are gathered

into 14 broad bands. The anal vesicles are covered with numerous short-

stalked funnels, which are visible to the naked eye. In addition to the interbasal

muscle, important differences also occur in the size and distribution of the

dermal papillae, in the number of muscle bands and in the colour of the

animals.

Ochetostoma mercator, described by Wesenberg-Lund (1954), is known only

from the holotype. In this species the trunk is 25 mm long and the proboscis,

which is broad, fleshy and non-deciduous, is one-fifth its length. An important

feature that distinguishes this species from O. natalense is the presence of

30-35 rings of large, oblong papillae on the trunk. Another difference lies in the

longitudinal muscle layer, which in O. mercator is gathered into 13 bands. The

proboscis of O. mercator is also much smaller in comparison with the length of

the trunk.

Fisher (1946) erected the species O. octomyotum from several specimens

collected from the Californian coast. His description mentions that the trunk of

full-grown specimens is 110 mm long and the proboscis ranges from one-third to

the full length of the body. The longitudinal muscle layer is gathered into eight

bands, but in the posterior third of the trunk it forms a continuous sheet. The

integument of O. octomyotum is translucent. Thus important differences between

the two species lie mainly in the number of longitudinal muscle bands and in the

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

nature of the integument. As in O. natalense, the ventral setae have no interbasal

muscle.

Ochetostoma pellucidum was described originally from several specimens

from the West African coast at depths of 10 m. In this species the trunk, which is

light green to bluish in colour, is 25 mm long and the proboscis is about one-

quarter its length. The longitudinal muscles are gathered into 13 bands. Fischer’s

(1895) description mentions that the integument is opaque anteriorly and

posteriorly but transparent in the middle region. Hence differences exist between

O. pellucidum and O. natalense in the number of longitudinal muscle bands and

in the nature of the integument. Other differences are present in the colour and

size of the specimens. Furthermore, the anal vesicles of O. pellucidum are only

about one-quarter as long as the trunk and the ventral setae are large.

The species O. glaucum is known from two rather badly preserved

specimens. According to the description provided by Wesenberg-Lund (1957),

the trunk is about 22 mm in length and the proboscis is 6 mm long. An interesting

feature in this species is the presence of two rows of processes on the proximal

part of the proboscis. The outer row consists of rather long lobes with slightly

scalloped edges. The integument is smooth with only a few rings of papillae on the

posterior part of the trunk. There are 12—13 longitudinal muscle bands. This

species differs significantly from O. natalense in the structure of the proboscis, in

the distribution of the papillae and in the number of longitudinal muscle bands.

Thus it is evident that O. natalense differs significantly from the other known

species of Ochetostoma that possess two pairs of gonoducts, thereby justifying the

establishment of a new species.

Etymology

This species is named after the province of Natal, along which coast it was

found.

Ochetostoma baronii (Greeff, 1879)

Figs 5-7

Thalassema baronii Greeff, 1879: 151, pl. 6 (figs 62-67). Selenka, 1885: 8. Shipley, 1899a: 55;

1899b: 336, 345, pl. 33 (figs 1, 7). Sluiter, 1902: 47. Augener, 1903: 348. Verrill, 1904: 40.

Lanchester, 1905: 34. Fischer, 1922: 15. Hérubel, 1924: 108.

Thalassema kefersteini A. ten Broeke, 1925: 94.

Ochetostoma baronii Fisher, 1946: 241. Stephen, 1960: 513. Mackie, 1961: 247. Stephen &

Edmonds, 1972: 429. Amor, 1976: 123.

Ochetostoma edax Fisher, 1946: 245, fig. 14.

Type-locality

Arrecife, Canary Islands (Atlantic Ocean).

Material

One sexually mature female, Ponté Torres, west coast of Inhaca Island;

collected by Zoology Department, Witwatersrand University, period 1956-1963.

ECHIURA FROM SOUTHERN AFRICA 41

Two juvenile specimens, Park Rynie Beach, Natal; collected 28 May 1983 and

8 September 1983.

Distribution

Numerous localities in tropical and subtropical waters of the Indian, Atlantic

and Pacific oceans (Stephen & Edmonds 1972).

Habitat

At both localities the specimens were found under rocks in the intertidal

zone. While the specimen from Inhaca occurred in mud in a permanent pool,

those from Park Rynie were found in relatively coarse sand.

Description

Size. Trunk of preserved specimen from Inhaca 66 mm long, greatest

diameter 17 mm; proboscis 18 mm long, detached from specimen. Trunk length

of Park Rynie specimens 21 mm and 30 mm and proboscis length 6 mm and

8 mm, respectively. Hence proboscis ranges from about a quarter to one-third

trunk length.

Colour. In living specimen from Inhaca Island, trunk purple, proboscis pale

yellow with fine green stripes. Colour of trunk and proboscis of Park Rynie

specimens rich dark green; green coloration due to symbiotic algae. In preserved

specimens, colour ranges from pale pink to dull greyish-brown.

External features

Proboscis. Proboscis markedly deciduous. In living specimens may exceed

half length of trunk. Proboscis spatula-shaped in living specimens but tubular

when preserved (Fig. 5A). Anterior and lateral margins smooth and free at base

of proboscis. In smaller specimen from Park Rynie, proboscis detached during

narcotization.

Trunk. Trunk sausage-shaped, more or less of uniform diameter in both

specimens from Park Rynie. Trunk of Inhaca specimen broad anteriorly but

tapering at posterior end (Fig. 5B).

Entire integument of trunk densely covered with minute, rounded papillae.

Papillae irregularly distributed, more closely arranged at extremities of trunk and

larger posteriorly. Much smaller papillae interspersed among larger ones.

Integument translucent, parts of alimentary canal and nerve cord visible.

Longitudinal muscles gathered into 18-20 bands. Inner oblique muscles between

longitudinal bands distinctly fasciculated.

Setae. Setae one pair, located on antero-ventral surface of trunk, just

posterior to mouth (Fig. 5B). Genital pores two pairs, opening behind setae.

Each seta consisting of a cylindrical shaft with curved terminal end (Fig. 5C).

Distal bent end slightly flattened. Cylindrical part of shaft with faint concentric

markings. Setae embedded in connective tissue and located in cone-shaped setal

sacs as in other species. Cylindrical interbasal muscle present.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

0,5 mm

Fig. 5. Ochetostoma baronii. A. Detached proboscis. B. Ventral view of trunk.

C. Left functional seta.

Internal anatomy

Alimentary canal. Alimentary canal long, coiled, about five times trunk

length (Fig. 6). Foregut extremely small, ending at intestinal ring vessel. Gizzard

and crop not distinguishable as no distinct boundaries present externally. Fine

mesenteric strands fasten alimentary canal to body wall at several points.

Numerous dilatations present in intestine due to presence of sand grains and shell

fragments. Contents of intestine not in the form of pellets. Small precloacal

caecum Opening ventrally into rectum.

ECHIURA FROM SOUTHERN AFRICA 43

> gon

int

aE eeeenel

10 mm

Fig. 6. Ochetostoma baronii. Dorsal dissection of the trunk, illustrating the

alimentary canal and anal vesicles.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anal vesicles. Anal vesicles thin-walled, unbranched, blind-ending tubes

opening into rectum (Fig. 6). Right anal vesicle of Inhaca specimen two-thirds

length of trunk, without ciliated funnels. Left vesicle approximately one-quarter

length of trunk with tiny unstalked funnels on distal third. Right anal vesicle of

smaller specimen from Park Rynie almost as long as trunk; left one less than one-

third trunk length. Both vesicles in larger specimen almost equal to trunk length;

ciliated funnels present on vesicles of both specimens.

Gonoducts. Gonoducts two pairs, located posterior to ventral setae (Fig. 7).

Gonoducts of Inhaca specimen considerably distended due to presence of

numerous ova. Gonostomal lips long, spirally coiled (Fig. 7). Openings of

gonostomes basal in position. Both specimens from Park Rynie immature.

Gonoducts oval in smaller specimen, somewhat elongate in larger one.

‘3 Ke:

p \

\ 0)

‘

;

gon

5 mm

Fig. 7. Ochetostoma baronii. Anterior end of the trunk cavity, showing

the blood system and gonoducts.

ECHIURA FROM SOUTHERN AFRICA 45

Blood system. Neuro-intestinal vessels paired, elongate, connecting dorsal

and ventral vessels by means of intestinal ring sinus (Fig. 7). Ring sinus located at

end of foregut. Interbasal muscle passes through loop of neuro-intestinal vessel.

Dorsal vessel weakly coiled with numerous small dilatations, passes anteriorly

and enters proboscis. Ventral vessel continues posteriorly, beside nerve cord,

terminating in region of rectal caecum.

Remarks

The present specimens of O. baronii represent new records for the southern

African region, extending the range of distribution southwards from the Zanzibar

coast.

According to Greeff (1879), the trunk is up to 80 mm in length, dark green in

colour with violet longitudinal muscle bands and white papillae scattered over the

surface. The longitudinal muscle layer is gathered into 17-19 bands. Selenka’s

description (1885), however, mentioned the presence of 20—23 muscle bands. An

interesting feature mentioned by Greeff (1879), Selenka (1885) and Shipley

(1899a) is the presence of short branching outgrowths on the anal vesicles.

The present specimens of O. baronii from the east coast of southern Africa

closely approach the description given by Greeff (1879), except for differences in

the structure of the anal vesicles. All the specimens lack the branched outgrowths

and conform with the descriptions provided by Fisher (1946), Lanchester (1905)

and Amor (1976).

Furthermore, the specimens from southern Africa as well as those examined

by Amor (1976) have a markedly deciduous proboscis, a feature that seems to

have gone unnoticed by previous authors.

Amor (1976) examined the type specimens of O. kefersteini and O. edax and

compared them with numerous specimens of O. baronii collected from the

Canary Islands. From this investigation she concluded that both O. edax and

O. kefersteini were synonymous with the species of Greeff. The present study

supports Amor’s conclusion.

The species of O. myersae Edmonds, 1963, based on four specimens from

Long Reef, New South Wales, appears to be closely related to O. baronii and,

according to Edmonds, the colour of living specimens is chlorophyll green. The

length of the trunk ranges from 20 mm to 35 mm and the proboscis, which is not

readily deciduous, is about a quarter to half the length of the trunk. The

longitudinal muscles are gathered into 18-21 bands. An important difference

between O. myersae and O. baronii lies in the oblique muscles, which in the

former species are weakly developed. The integument is covered with soft, white,

wart-like papillae, which are large and most conspicuous on the posterior third of

the trunk. Edmonds (1963) reported the presence of two pairs of postsetal

gonoducts in three of his specimens and only one pair in his largest specimen. As

in O. baronii, an interbasal muscle is present that passes through a loop of the

neuro-intestinal vessel. The anal vesicles are thin, elongate tubes that lack

branched outgrowths. The ciliated funnels are borne on very short stalks. Hence

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

differences between O. baronii and O. myersae are apparent in the integument,

the proboscis and in the size and arrangement of the dermal papillae. The

differences appear to be sufficiently distinctive to retain these as two separate

species.

Ochetostoma formosulum (Lampert, 1883)

Figs 8-10

Thalassema formosulum Lampert, 1883: 339. Shipley, 1899b: 348. Sluiter, 1902: 48, fig. 13.

Wharton, 1913: 248. Prashad, 1921: 35.

Ochetostoma formosulum Fisher, 1946: 241. Wesenberg-Lund, 1963: 140. Datta Gupta et al.,

1963: 57, figs la—c. Stephen & Edmonds, 1972: 433.

Type-locality

Manila, Philippines.

Material

Two specimens, one sexually mature, from Durban Bay; collected by

University of Cape Town Ecological Survey, 1963. One of these specimens

(dissected) was described by Wesenberg-Lund (1963).

Distribution

Indo-West-Pacific, from Japan, the Philippines and Indonesia, and the Natal

coast.

Habitat

Both specimens occurred in sand at a depth of 8-9 m.

Description

Size. Trunk of intact sexually mature specimen 22 mm long, greatest

diameter 11 mm. Proboscis 5 mm long, less than one-quarter trunk length.

Colour. Dissected specimen cream, intact specimen pinkish-brown.

External features

Proboscis. Proboscis fleshy, highly contracted, non-deciduous and more or

less spherical in outline (Fig. 8A). A series of tight ridges on lateral edges,

margins frilled. Dorsal surface with small, scattered papillae. Proboscis with

broad incision at anterior end. Lateral margins fused posteriorly, forming a

narrow lower lip.

Trunk. Trunk broad at anterior end, tapering posteriorly (Fig. 8A). White

papillae sparsely distributed over middle two-thirds or more of trunk, more

closely arranged at extremities. Papillae arranged in rings at posterior end

(Fig. 8A). Anus at tip of small, smooth conical projection. A ring of large,

elongate papillae around base of conical projection about 3 mm away from

posterior tip of trunk. Integument very thin and transparent, somewhat thicker

ECHIURA FROM SOUTHERN AFRICA 47

-———_______

5 mm

Fig. 8. Ochetostoma formosulum. A. Ventral aspect. B. Left functional seta.

and opaque posteriorly. Nerve cord, gonoducts and parts of alimentary canal

visible through transparent body wall. Genital pores two pairs, postsetal. Muscle

layers continuous over most of trunk. Longitudinal muscle bands indistinct except

in posterior third of trunk where narrow bands faintly discernible. Inner oblique

muscle layer fasciculated, visible only in posterior end of trunk.

Setae. Setae large, golden-brown in colour, hook-like, about 4 mm long

(Fig. 8B). Terminal bent end flattened and clearly demarcated from straight,

cylindrical part of shaft by a conspicuous ridge. Both setae located in special sacs,

supported by numerous radiating muscle strands. Interbasal muscle cylindrical,

passing through small loop of neuro-intestinal vessel.

Internal anatomy

Alimentary system. Alimentary canal long, highly coiled tube, fastened to

body wall at several points by thin mesenteric strands. Pharynx club-shaped with

ANNALS OF THE SOUTH AFRICAN MUSEUM

a Pav

Pr int

SS gon

TFT €)

Fig. 9. Ochetostoma formosulum. A. Posterior end of trunk cavity, showing the

anal vesicles and posterior part of the alimentary canal. B. Anterior part of the

trunk cavity, showing the gonoducts.

ECHIURA FROM SOUTHERN AFRICA 49

thick muscular walls. Oesophagus narrow, more or less of uniform diameter.

Foregut long, terminating at intestinal ring sinus. Presiphonal region of intestine

with a ciliated groove. Intestine considerably dilated, extremely thin-walled and

transparent in region of siphon. Entire intestine compactly filled with small,

sausage-shaped faecal pellets consisting of fine sand particles (Fig. 9A). Rectal

caecum small, spherical.

Anal vesicles. Anal vesicles small, about one-third trunk length, unbranched

and transparent (Fig. 9A). Narrow anteriorly but considerably dilated and sac-

like posteriorly. Ciliated funnels minute, stalked, sparsely distributed over

surface of both vesicles.

Gonoducts. Gonoducts two pairs (Fig. 9B), posterior to ventral setae, oval in

juvenile specimen but elongate, tubular and considerably distended in sexually

mature individual due to presence of numerous ova. Gonostomal lips thin,

spirally coiled; gonostomal openings basal in position.

Blood system. Intestinal ring sinus is an incomplete vascular ring at posterior

end of foregut (Fig. 10). Dorsal vessel prominent, continuing anteriorly and

entering proboscis. Paired neuro-intestinal vessels short, stout, forming small

loop around interbasal muscle (Fig. 10). Heart conspicuous at posterior end of

dorsal vessel. Ventral vessel terminating posteriorly in rectal caecum.

+—_—_____—_______4

3 mm

Fig. 10. Ochetostoma formosulum. Anterior part of the trunk cavity,

showing the blood vessels.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Ochetostoma formosulum (Lampert, 1883) was described from a single

specimen from Manila, Philippines. This species was later recorded and described

from several other localities in the Indo-West-Pacific Ocean. Wesenberg-Lund

(1963) recorded and described a single specimen collected from a central

sandbank in Durban Bay. Although this species is fairly well known, some

features require further discussion.

The most important distinguishing features of O. formosulum are the

presence of two pairs of postsetal gonoducts with spirally coiled gonostomal lips,

a very thin and transparent integument, 6—8 longitudinal muscle bands and an

interbasal muscle that passes through a loop of the neuro-intestinal vessel. Other

taxonomic features include the proboscis which is less than one-quarter of the

trunk length, the frilled lateral margins of the proboscis, and the characteristic

shape and distribution of the dermal papillae.

It is rather surprising to note that, with the exception of Wharton (1913),

none of the other authors mentioned the presence of an interbasal muscle in this

species. In their key to species of Ochetostoma, Datta Gupta & Menon (1971)

were incorrect in including O. formosulum with those species that lack an

interbasal muscle.

The description provided by Lampert (1883) mentioned a trunk length of

30 mm and proboscis length of 8 mm. With the exception of the dermal papillae,

the present specimens from Durban Bay closely correspond with Lampert’s

description. According to Lampert (1883) and Wharton (1913), the papillae are

not arranged in transverse or longitudinal rows on any part of the trunk.

Sluiter’s description (1902) was based on 21 well-preserved specimens from

Siboga. The trunk ranged from 15 mm to 23 mm in length and the proboscis from

4 mm to 6 mm. Sluiter also mentioned the presence of a conical projection and a

ring of large papillae at the posterior extremity of the trunk.

Wharton’s description (1913) was based on a single individual from Samar.

The trunk is 35 mm long and the proboscis is less than one-third its length.

The trunk of the single specimen described by Wesenberg-Lund (1963) is

17 mm long and the proboscis 6 mm. Hence from the description provided by the

above authors, it is apparent that the proboscis ranges from a quarter to one-third

the trunk length. Regarding the dermal papillae, Wesenberg-Lund (1963: 140)

stated: “The dermal organs show a slight tendency to an annular arrangement;

they are nowhere especially crowded, and a small area round the posterior end

seems almost smooth.’

The anal vesicles are broad and sac-like in the intact specimen from Durban

Bay and hence similar to those described by Lampert (1883), Wharton (1913) and

Datta Gupta et al. (1963). Wesenberg-Lund (1963), however, described these

organs as white, transparent tubes.

Ochetostoma formosulum appears to be related to O. septemyotum and

O. arkati in possessing two pairs of gonoducts and 7-8 longitudinal muscle bands.

The latter two species, however, besides lacking an interbasal muscle, also differ

ECHIURA FROM SOUTHERN AFRICA 51

significantly in the structure of their proboscises. In O. septemyotum the lateral

margins of the proboscis are produced into a series of folds with dendritic

outgrowths; in O. arkati gill-like structures are present along the basal edge of the

proboscis.

Ochetostoma decameron (Lanchester, 1905)

Figs 11-13

Thalassema decameron Lanchester, 1905: 35, pl. 1 (fig. 5).

Ochetostoma decameron Fisher, 1946: 241. Stephen & Edmonds, 1972: 432.

Holotype

From Ohwaka Bay, Zanzibar; deposited by Lanchester in the British

Museum (Natural History).

Material

One sexually mature specimen, Park Rynie Beach (31°19'S 30°44’E), Natal

coast; collected 8 September 1983.

Distribution

Ohwaka Bay, Zanzibar, and Natal coast, South Africa.

Habitat

The Park Rynie specimen occurred in sand, under rocks in the intertidal

area, close to the high-water mark.

Description

Size. Trunk of preserved specimen 18 mm long, greatest diameter 6 mm.

Proboscis half trunk length.

Colour. In living specimen, proboscis pale yellow, trunk reddish-purple.

Colour of preserved specimen pale pink.

External features

Proboscis. Proboscis fleshy, non-deciduous, spatula-shaped in living speci-

mens but in preserved condition lateral margins roll inwards forming a tube

(Fig. 11A). Anterior and lateral margins smooth. Small, rounded papillae visible

on dorsal surface under dissecting microscope. Lateral margins of proboscis

united at base, forming a narrow lower lip ventral to mouth.

Trunk. Trunk broad in middle region, tapering more towards posterior end

(Fig. 11A). Integument thin and transparent. Small, rounded papillae irregularly

distributed over entire surface of trunk, more closely arranged at extremities.

Trunk papillae larger than those on proboscis and not of uniform size. Much

smaller papillae interspersed among larger ones.

Longitudinal muscle bands not visible from external surface but, under

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

dissecting microscope, 10 inconspicuous bands apparent only in middle region of

trunk. Longitudinal and inner oblique muscles form a continuous sheet in mid-

dorsal region of trunk. Oblique muscles weakly developed, arranged in fascicles

between longitudinal bands.

Setae. Setae one pair, golden-yellow, located about 3 mm away from anterior

end of trunk (Fig. 11A). Each seta consisting of a cylindrical shaft with a curved

terminal end tapering in a sharp point (Fig. 11B). Distal third of seta much

narrower. Interbasal muscle absent.

0,3 mm

Fig. 11. Ochetostoma decameron. A. Ventral aspect. B. Left functional seta.

ECHIURA FROM SOUTHERN AFRICA 33)

Internal anatomy

Alimentary canal. Alimentary canal comparatively short, consisting of a few

irregular coils (Fig. 12A). Foregut short, narrow tube of uniform diameter,

terminating at ring sinus. Thin-walled presiphonal and siphonal regions of

intestine compactly filled with coarse sand grains and shell fragments. Intestinal

contents not in the form of pellets. Few thin mesenteric strands fastening

alimentary canal to body wall. A small spherical rectal caecum present.

Anal vesicles. Anal vesicles thin-walled transparent tubes, about half length

of trunk (Fig. 12A). Ciliated funnels few, unstalked over surface of both vesicles.

Gonoducts. Gonoducts four pairs, white in colour; first pair anterior to

ventral setae while remaining three pairs postsetal in position (Fig. 12B). Presetal

gonoducts smallest of all, situated just anterior to cone-shaped setal sacs and

partially obscured by muscle strands radiating from base of setae. Gonoducts

spherical to oval and equidistant from each other. Postsetal gonoducts distended

due to presence of gametes. Gonostomal lips broad, weakly coiled and large in

comparison to size of gonoducts (Fig. 12C). Gonostomal openings basal in

position.

Blood system. Neuro-intestinal vessels short, connecting dorsal and ventral

vessels by means of intestinal ring sinus. Paired neuro-intestinals unite anteriorly

before opening into ventral vessel (Fig. 13). Dorsal vessel prominent, passing

anteriorly and entering proboscis. Ventral vessel continuing posteriorly, along-

side nerve cord and terminating in rectal caecum.

Remarks

Ochetostoma decameron is based on a single specimen from Zanzibar, in

which the proboscis was missing. Its discovery at Park Rynie is a new record for

southern Africa. Lanchester’s (1905) description was extremely brief and,

according to Stephen & Edmonds (1972), the taxonomic position of this species is

still uncertain. Nothing is known about the size of the specimen, the structure of

the proboscis, the disposition of the oblique muscles, and the distribution of the

dermal papillae. Lanchester mentioned the presence of four pairs of gonoducts,

with the first two pairs presetal in position. The body wall is extremely thin and

the longitudinal muscles are aggregated into 10 fairly broad but inconspicuous

bands. According to Stephen & Edmonds (1972), the holotype in the British

Museum (Natural History) is damaged and the muscle systems are not clearly

evident.

Important distinguishing features of the present specimen from Park Rynie

include the four pairs of gonoducts, 10 longitudinal muscle bands, a thin and

transparent integument, the narrow lower lip of the proboscis and the

characteristic shape and arrangement of the dermal papillae. Other features of

lesser taxonomic importance are the colour and size of the specimen.

The present specimen closely approaches Lanchester’s (1905) description in

the number of gonoducts and longitudinal muscle bands, as well as in the nature

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

prs gon

Fig. 12. Ochetostoma decameron. A. Dorsal dissection of the trunk, showing the alimentary

system and anal vesicles. B. Anterior part of the trunk cavity showing the gonoducts. C. First

postsetal gonoduct from left.

ECHIURA FROM SOUTHERN AFRICA a)

Fig. 13. Ochetostoma decameron. Anterior part of

trunk cavity, showing blood vessels.

of the integument. An important difference, however, lies in the location of the

gonoducts in relation to the ventral setae.

Ochetostoma decameron appears to be related to O. kempi in possessing four

pairs of gonoducts. According to Prashad (1919), the trunk of O. kempiis 77 mm

long and the longitudinal muscles are gathered into 20 bands. The integument of

the latter species is thick and opaque. Hence, significant differences are present in

the size of the animals, in the number of longitudinal muscle bands, and in the

nature of the integument. Another distinguishing feature lies in the oblique

muscles, which in O. decameron are weakly developed.

Additional material in the future will give a better understanding of the

variations within O. decameron and its taxonomic position.

Ochetostoma kempi (Prashad, 1919)

Figs 14-16B

Thalassema kempi Prashad, 1919: 336, fig. 2; 1935: 41.

Ochetostoma kempi Fisher, 1946: 241. Datta Gupta & Menon, 1971: 176, fig. 2. Stephen &

Edmonds, 1972: 437.

Holotype

From the Andaman Islands; deposited by Prashad in the Indian Museum,

Calcutta (cat. no. W194/4).

Material

One specimen, Isipingo Beach (29°05'S 30°56’E), Natal coast; collected

12 May 1983.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Andaman Islands and Natal coast, South Africa.

Habitat

The specimen from Isipingo Beach occurred in sand under rocks in a rock

tunnel, in the intertidal zone close to the high-water mark.

Description

Size. Trunk of preserved specimen 69 mm in length, greatest diameter

16 mm. Proboscis 39 mm long, slightly exceeding half length of trunk.

Colour. In live animal, proboscis pale yellow, trunk with reddish-purple

stripes marking longitudinal muscle bands. Interspaces narrow, bluish-grey in

colour. In preserved condition the colour changed to light brown.

External features

Proboscis. Proboscis fleshy, non-deciduous, slightly truncated at anterior

end. Spatula-shaped in live specimen but in preserved state lateral margins curl

inwards forming a narrow tube (Fig. 14A). Lateral margins of proboscis smooth

and free at base.

Trunk. Trunk sausage-shaped, tapering posteriorly (Fig. 14A). Small,

rounded papillae irregularly distributed over entire surface of integument.

Papillae minute on anterior two-thirds or more of trunk but larger posteriorly.

Entire integument thick and opaque. Longitudinal muscle layer gathered into

17-19 bands; inner oblique muscles between longitudinal bands distinctly

fasciculated. A pair of golden-yellow setae located about 8mm away from

anterior end of trunk.

Setae. Each seta consisting of a cylindrical shaft with curved terminal end,

tapering in a pointed tip (Fig. 14B). Proximal end of shaft slightly bent. Non-

functional or replacement seta small, in close association with right functional one

(Fig. 14C). Fine concentric ridges on surface of both setae. Interbasal muscle

absent.

Internal anatomy

Alimentary canal. Alimentary canal slightly coiled, several times length of

trunk. Intestine attached to body wall at several points by very thin and elongate

mesenteric strands. Ventral sheet of mesentery attaching oesophagus to body

wall. Foregut small, ending at ring sinus. Intestine subdivided into presiphonal,

siphonal and postsiphonal regions as in related species. Presiphonal region with

ciliated groove. Intestine considerably dilated and extremely thin-walled in region

of siphon. Gut compactly filled with fine sand grains and shell fragments, not

moulded into pellets. Rectal caecum large and spherical (Fig. 15).

Anal vesicles. Paired anal vesicles long, slender tubes exceeding length of

trunk (Fig. 15). Numerous, tiny, unstalked ciliated funnels on surface of both

vesicles.

ECHIURA FROM SOUTHERN AFRICA

] seta.

10na

Ventral aspect. B. Right

non-funct

functional seta. C. Much smaller,

Fig. 14. Ochetostoma kempi.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

10 mm

Fig. 15. Ochetostoma kempi. Dorsal dissection of the trunk, showing the anal vesicles.

Gonoducts. Gonoducts four pairs, first pair presetal (Fig. 16A). Arrange-

ment of gonoducts asymmetrical; right presetal gonoduct slightly anterior to

corresponding one on left. Second and third pairs on both sides located very close

to each other, with second pair anterolateral in position to third pair. First and

second pairs of gonoducts small, more or less oval while fourth pair elongate,

tubular and largest of all (Fig. 16A). Gonostomal lips small, spirally coiled,

Opening into base of gonoduct.

Blood system. Ring sinus at end of foregut is an incomplete vascular ring

(Fig. 16B). Paired neuro-intestinal vessels unite prior to opening into ventral

ECHIURA FROM SOUTHERN AFRICA

10 mm

Fig. 16. Ochetostoma kempi. Anterior part of trunk cavity. A. Gonoducts.

B. Blood vessels.

Sy)

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

vessel. Dorsal vessel prominent, arising from ring sinus and passing anteriorly

into proboscis. Ventral vessel terminating near rectal caecum.

Remarks

The discovery of O. kempi from Isipingo 1s a new record for southern Africa.

However, it is not unexpected for this species to be found as far west as the east

coast of southern Africa.

According to Prashad (1919), the trunk, which is 77 mm in length, is wider in

the middle region but gradually tapers at both ends. The proboscis is short and

stumpy and slightly truncated at the anterior end. It is 17 mm long and hence less

than a quarter the trunk length. Prashad stated that both the proboscis and trunk

are covered with papillae. On the proboscis, the papillae are minute and just

visible, while on the trunk they are small anteriorly but gradually increase in size

at the posterior end. A few large papillae are also scattered among the smaller

ones over the whole surface. The longitudinal muscle layer is gathered into

20 bands. According to Prashad (1919), there are four pairs of gonoducts located

posterior to the ventral setae. The first two pairs are poorly developed but the

fourth pair is the best developed of all. The anal vesicles are very much contracted

and about one-third the length of the trunk. Prashad (1935), however, re-

examined the holotype of O. kempi and found that the first two pairs of

gonoducts were presetal in position.

The present specimen from Isipingo differs from the type specimen in several

respects. The most important differences lie in the position of the gonoducts in

relation to the ventral setae, the size and arrangement of the dermal papillae, and

the length of the anal vesicles. The papillae on the anterior two-thirds of the trunk

are small and evenly distributed and are not interrupted by larger papillae. The

differences in the size of the proboscis and anal vesicles could be due to different

degrees of contraction of these organs during narcotization and fixation. It is very

likely that Prashad’s (1919) description was based on a specimen that was highly

contracted.

Datta Gupta & Menon (1971) identified a specimen, also collected from the

Andaman Islands, as O. kempi. According to these authors, the proboscis is

nearly tubular with the lateral margins slightly indented. The body is covered with

rounded papillae that are prominent towards the posterior end. An interesting

feature about their specimen is that the extreme posterior end of the trunk is

devoid of papillae and is made up of concentric fleshy rings. The longitudinal

muscle layer is gathered into 20 bands. These authors also mentioned the

presence of four pairs of gonoducts, with the first pair presetal in position. Hence

the arrangement of the gonoducts, in relation to the ventral setae, conforms with

that of the specimen from Isipingo. The size and distribution of the dermal

papillae and the anatomy of the blood system are also very similar. The specimen

described by Datta Gupta & Menon (1971), however, differs in lacking a rectal

caecum. The posterior end of the trunk of their specimen also appears to be

rather unusual. Unfortunately, these authors did not give any details regarding

ECHIURA FROM SOUTHERN AFRICA 61

the size of their specimen. However, from their illustration it is apparent that the

proboscis is about half the length of the trunk.

The specimen from Isipingo is ascribed to O. kempi on the basis of the

number of gonoducts and longitudinal muscle bands. According to Datta Gupta

(1976) and Saxena (1983), the number of gonoducts is constant within a species

and is an important taxonomic character in echiurans.

Ochetostoma kempi appears to be related to O. decameron in possessing four

pairs of gonoducts. Significant differences, however, occur in the size of the

animals, in the number of longitudinal muscle bands and in the nature of their

integuments. Additional material in the future should give a better understanding

of the variations within O. kempi.

Ochetostoma palense (Ikeda, 1924)

Figs 17-19B

Thalassema palense Ikeda, 1924: 39, figs 13-15.

Ochetostoma palense Stephen & Edmonds, 1972: 440.

Material

One specimen, Isipingo Beach (29°05’S 30°56’E), Natal coast; collected

13 June 1983.

Distribution

Known only from the holotype from Palau Islands, Japan, and Natal coast,

South Africa.

Habitat

The Isipingo specimen occurred in coarse sand under rocks in a rock tunnel,

in the intertidal area, close to the high-water mark.

Description

Size. Trunk of preserved specimen 22 mm long, greatest diameter of trunk

7 mm. Proboscis 7 mm long, about one-third trunk length.

Colour. In living specimen, proboscis pale yellow, trunk reddish-purple.

Preserved specimen white.

External features

Proboscis. Proboscis spatulate, non-deciduous; lateral margins smooth and

free at base. When preserved, lateral edges of proboscis curl inwards forming a

tube (Fig. 17A).

Trunk. Trunk cylindrical, tapering gradually towards posterior end

(Fig. 17A). Anus at tip of small conical projection. Papillae small, rounded,

irregularly distributed over entire surface of trunk, more closely arranged at

posterior end. Integument extremely thin and transparent. Nerve cord and other

internal organs visible through transparent body wall. Longitudinal muscle bands

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. Ochetostoma palense. A. Ventral aspect. B. Right functional seta.

not visible externally. In dissected specimen, 15 inconspicuous muscle bands

apparent only in posterior half of trunk. Interspaces narrow. In anterior half of

trunk longitudinal and oblique muscles form a continuous sheet. Oblique muscles

between longitudinal bands weakly fasciculated.

Setae. Setae minute, golden-yellow, about 3 mm away from anterior end of

trunk. When dissected out and viewed under compound microscope, each seta

consists of a straight shaft with a curved terminal end (Fig. 17B). Proximal half of

shaft broader with a few concentric markings at base. Interbasal muscle absent.

Internal anatomy

Alimentary canal. Alimentary canal comparatively short, about three times

trunk length and with relatively few coils (Fig. 18). Intestine attached to body

ECHIURA FROM SOUTHERN AFRICA 63

wall by a few thin mesenteric strands. Foregut is a narrow tube, about 4 mm long,

ending at ring sinus. Presiphonal region of intestine 6 mm in length, marked by

presence of a ciliated groove. Ciliated groove commences soon after ring sinus

and leads into intestinal siphon. Siphonal region of intestine considerably dilated

and extremely thin-walled. Intestine narrows in postsiphonal region and leads

into more or less straight rectum. A small, spherical rectal caecum present.

4 mm

Fig. 18. Ochetostoma palense. Dorsal dissection of the trunk,

showing the alimentary canal.

Anal vesicles. Anal vesicles (Fig. 18) small, less than one-quarter trunk

length. Ciliated funnels minute, unstalked, on surface of both vesicles.

Gonoducts. Gonoducts three pairs, spherical to somewhat oval in shape with

weakly coiled gonostomal tips (Fig. 19A). First pair of gonoducts presetal.

Gonostomal lips broad, unlike condition in several other species of Ochetostoma.

Second and third pairs of gonoducts much larger than first pair.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. Ochetostoma palense. Anterior part of the trunk cavity.

A. Gonoducts. B. Blood vessels.

ECHIURA FROM SOUTHERN AFRICA 65

Blood system. Intestinal ring sinus is an incomplete vascular ring at end of

foregut (Fig. 19B). Paired neuro-intestinal vessels unite before opening into

ventral vessel. Dorsal vessel prominent, arising from ring sinus and entering

proboscis.

Remarks

Ochetostoma palense (Ikeda, 1924) is based on a single specimen from Palau

Islands, Japan. The holotype is small, about 22 mm in its entire length, of which

about one-quarter belongs to the proboscis. The pear-shaped trunk is 8 mm at its

widest part. Ikeda mentioned that the animal is uniformly bright green in life. The

thin and transparent integument is covered with minute papillae, which are larger

and more closely arranged at the extremities of the trunk. The longitudinal

muscles are gathered into 15 bands that are apparent only on a small portion of

the anterior part of the trunk. An interbasal muscle is absent. There are three

pairs of spherical gonoducts with slightly coiled gonostomal lips. The first pair of

gonoducts are presetal. The anal vesicles are thin-walled tubes, over half the

length of the trunk, with few unstalked funnels. A rectal caecum is absent.

According to Ikeda (1924) the alimentary canal is short as compared with the size

of the animal, with a few irregular coils.

The present specimen from Isipingo is similar to the Japanese specimen in

size, distribution of the dermal papillae, the nature of the integument, as well as

in the number of gonoducts and longitudinal muscle bands. In both specimens the

gonostomal lips are weakly coiled. The specimen from Isipingo, however, differs

in possessing a rectal caecum. Differences are also present in the colour of the

animals as well as in the lengths of the anal vesicles.

Ochetostoma palense appears to be closely related to O. erythrogrammon

Leuckart & Riippell, 1828, and Wesenberg-Lund (1939) considered the species to

be synonymous. However, there are several features that tend to separate the two

species. An interesting feature of the specimen from Isipingo is that the

alimentary canal is comparatively short, about three times the trunk length. In

O. erythrogrammon tt is long and highly coiled, and measurements made in a few

specimens have shown that it is five times as long as the trunk. The gonostomal

lips of O. palense, unlike those of O. erythrogrammon, are not thread-like and

are only weakly coiled. Another distinguishing feature is that the longitudinal

muscle bands of O. palense are not conspicuous throughout the trunk. Further-

more, the oblique muscles between the longitudinal bands are very weakly

developed.

Stephen & Edmonds (1972) refrained from placing O. palense in the

synonymy of O. erythrogrammon mainly because of the differences in the

dimensions of the alimentary canal and in the disposition of the gonostomal lips.

In view of the above differences, it has been decided to retain O. palense as a

species distinct from O. erythrogrammon. Any further decisions will have to

await the collection of more, preferably adult, material.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ochetostoma arkati (Prashad, 1935)

Thalassema arkati Prashad, 1935: 41, figs 1-4.

Ochetostoma arkati: Wesenberg-Lund, 1959: 203, figs 13-14; 1963: 141.

Type-locality

Sandheads off the mouth of the River Hooghly, Ganges Delta, Calcutta.

Diagnosis

Proboscis short and stumpy, a third to a quarter of total length of animal;

edges of posterior half form short, branched, gill-like processes. Ventral margins

of proboscis free at base. Dorsal surface of proboscis smooth; ventral side

trough-shaped, deepest at proximal end. Trunk oval, up to 38 mm in length,

covered with minute, round to oval papillae, which are very sparse or absent in

middle region. Borders of anus deeply crenulated and scalloped. Area around

anus smooth, thin-skinned and devoid of papillae. Longitudinal muscle layer

gathered into 7-8 bands, separated by broad interspaces. Two pairs of postsetal.,

tubular gonoducts almost as long as trunk; gonostomal lips elongate and spirally

coiled. Anal vesicles yellowish, very thin and about one-third length of animal.

Distribution

Calcutta, India (Prashad 1935); African coast, Nigeria and Cape Town

(Wesenberg-Lund 1959), and Port Elizabeth (Wesenberg-Lund 1963). It is very

likely that this species is widely distributed through the western part of the Indo-

Pacific, and the southern part of the eastern Atlantic.

Remarks

Wesenberg-Lund’s (1959, 1963) material agreed with Prashad’s (1935)

description, except that the number of longitudinal muscle bands was 7-8 instead

of eight. Ochetostoma arkati is adequately described and is distinct from all the

other species in the genus in possessing branched, gill-like processes along the

edges of the posterior half of the proboscis.

Ochetostoma capense Jones & Stephen, 1955

Ochetostoma capensis Jones & Stephen, 1955: 273, figs 1-3. Wesenberg-Lund, 1963: 142.

Diagnosis

Proboscis small, in preserved specimens about one-eighth of trunk length

and in living specimens one-twelfth to one-third; ventral edges crenate when

proboscis is extended, but thrown into a series of tight ridges when contracted.

Colour varies from cream, through different shades of cream-yellow, to light

orange. Trunk up to 120 mm when fully extended, thickly covered with papillae

on anterior ventral surface but fewer at posterior end. Papillae dispersed in rows

in middle region of trunk. Colour of trunk variable, ranging from dark grey-

brown to purple-brown. Trunk cylindrical or sausage-shaped and typically widest

ECHIURA FROM SOUTHERN AFRICA 67

at posterior end. Longitudinal muscles gathered into seven main bands, most

conspicuous at ends of trunk. A few weakly developed bands occur in between

the main bands. Two pairs of tubular, postsetal gonoducts, usually half to three-

quarters as long as trunk, but sometimes quite short. Gonostomal lips extended

into long spiral filaments. Interbasal muscle present, passes through a loop of

neuro-intestinal vessel. Anal vesicles long, thin, brown tubes of variable length,

from one- to three-quarters of trunk length and bearing minute, ciliated funnels,

which are most numerous towards the free ends of the vesicles. Rectal caecum

present.

Distribution

Zwartkops River, Langebaan Lagoon and Durban Bay (Jones & Stephen

1555)

Remarks

Ochetostoma capense is based on a large number of specimens. It is the only

estuarine species recorded from southern Africa, and is probably endemic. This

species appears to be related to O. septemyotum Datta Gupta, Menon &

Johnson, 1963, but differs significantly in the structure of the proboscis. This

species has been illustrated and fully described (Jones & Stephen 1955).

Ochetostoma sp.

Figs 20—23

Material

One sexually mature female, Park Rynie Beach, Natal; collected 7 June

1982.

Habitat

The specimen occurred in the intertidal zone in relatively coarse sand under a

projecting ledge of rock facing the shore.

Description

Size. Trunk of preserved specimen 40 mm in length, greatest diameter

8 mm. Proboscis 11 mm, about one-quarter as long as trunk.

Colour. Proboscis pale yellow in living specimen. Longitudinal stripes

marking longitudinal muscle bands dark red, interspaces bluish-grey. Colour of

preserved specimen pale pink.

External features

Proboscis. Proboscis non-deciduous, shovel-shaped with smooth lateral and

terminal margins (Fig. 20). Lateral margins of proboscis fuse at base forming a

narrow lower lip ventral to mouth. Ventral groove of proboscis smooth but dorsal

surface covered with minute, densely arranged, rounded papillae.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 20. Ochetostoma sp. Ventral aspect.

ECHIURA FROM SOUTHERN AFRICA 69

Trunk. Trunk sausage-shaped, tapering more towards posterior end

(Fig. 20). Entire integument densely covered with small, rounded papillae.

Papillae much larger and white in colour at posterior end of trunk. Longitudinal

muscles aggregated into 20 bands; inner oblique muscles between longitudinal

bands distinctly fasciculated. Entire integument thick and opaque. Ventral setae

missing. Two tiny scars on integument indicate that setae have fallen off.

Interbasal muscle absent.

Internal anatomy

Alimentary canal. Alimentary canal long, consisting of several ascending and

descending limbs. An interesting feature is the presence of numerous, closely

arranged, white mesenteric strands along entire length of gut (Fig. 21). Oesophagus

comparatively short, terminating at ring sinus. A ventral sheet of mesentery fastens

oesophagus to body wall. Intestine shows usual divisions into presiphonal, siphonal

and postsiphonal regions. Presiphonal region of intestine with a ciliated groove

commencing soon after ring vessel. In region of siphon, intestine considerably

dilated and thin-walled. A small, spherical rectal caecum present.

Anal vesicles. Anal vesicles paired distensible tubes exceeding length of

trunk (Fig. 21). Both vesicles covered with tiny, unstalked ciliated funnels.

Funnels more numerous at distal ends of vesicles.

Gonoducts. Gonoducts three pairs, tubular, light brown in colour. First pair

smallest, located anterior to ventral setae (Fig. 22A). Gonostomal lips small,

weakly coiled (Fig. 22B), unlike condition in several other species of Ocheto-

stoma. Gonostomes robust, and not thread-like in appearance. Proximal end of

gonostome broad and sac-like. All gonoducts compactly filled with eggs, visible

through transparent wall.

Blood system. Ring sinus is an incomplete vascular ring located at end of

foregut (Fig. 23). Paired neuro-intestinal vessels unite before opening into

ventral vessel. As in other species, dorsal vessel passes anteriorly and enters

proboscis.

Remarks

Important features of the present specimen include: the presence of an

unusually large number of mesenteric strands attaching the gut to the body wall,

the structure of the gonostomes, and the 1 : 4 ratio of the proboscis to the trunk

length. Another distinguishing feature seems to be the small size of the sexually

mature specimen.

Ochetostoma erythrogrammon, O. caudex, O. palense, O. australiense and

O. indosinense possess three pairs of gonoducts and 12-22 longitudinal muscle

bands and hence appear to be related to this specimen from Park Rynie.

The trunk length of sexually mature specimens of O. caudex ranges from

72 mm to 105 mm (Biseswar 1983), and in O. erythrogrammon from 80 mm to

160 mm (Stephen & Edmonds 1972). In both these species, the proboscis is one-

third to three-quarters as long as the trunk. This sexually mature specimen of

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 21. Ochetostoma sp. Dorsal dissection of the trunk, showing the

alimentary system and anal vesicles.

Ochetostoma sp., however, is small, with a trunk length of only 40 mm and the

proboscis is about one-quarter as long.

In addition, this specimen has a proboscis in which the lateral margins unite

posteriorly to form a narrow lower lip.

Another distinguishing feature of the present specimen lies in the structure of

the gonostomes. The gonostomal lips, unlike those of O. caudex and O. erythro-

grammon, are comparatively small and are not spirally coiled. Furthermore, in

ECHIURA FROM SOUTHERN AFRICA Gl

pr s gon

po s gon

1 mm

Fig. 22. Ochetostoma sp. A. Anterior part of trunk cavity, showing the

gonoducts. B. First postsetal gonoduct.

the latter two species, the mesenteric strands that fasten the gut to the body wall

are relatively few.

In O. palense the trunk is 22 mm long and the integument is thin and

transparent. Furthermore, the longitudinal muscle layer is gathered into

15 bands, which are apparent only on a small portion of the anterior part of the

trunk. Hence O. palense differs from the present specimen in size as well as in the

nature of the integument. Another important feature that distinguishes

WA ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. Ochetostoma sp. Anterior part of the trunk cavity, showing blood vessels.

O. palense is the presence of a comparatively short alimentary canal with a few

irregular coils. Ikeda (1924) also mentioned the absence of a rectal caecum.

In O. australiense the proboscis is up to 41 mm long and the trunk is up to

102 mm in length. The Australian specimens usually have 12-13 longitudinal

muscle bands, very occasionally 11-14. The gonostomal lips are long and spirally

coiled. According to Edmonds (1960), the anterior and posterior surfaces of the

trunk bear small flat papillae. Differences between O. australiense and the

present specimen are evident in the size of the animals, the number of

longitudinal muscle bands, the structure of the gonostomal lips and in the nature

and distribution of the dermal papillae.

Ochetostoma indosinense Wesenberg-Lund, 1939, from South Annam, is

known only from the holotype. In this species, the proboscis is 9 mm long and the

trunk length is 50 mm. According to Wesenberg-Lund (1939) the proboscis is

slightly bifurcated with the ventral and terminal rims crenated. The proboscis is

light green in life and the trunk is pink. There are 18 longitudinal muscle bands

with narrow interspaces. The gonostomal lips are long but not coiled.

Ochetostoma indosinense differs from the present specimen in several respects.

The proboscis of O. indosinense is not typical of Ochetostoma. Furthermore, the

inner oblique muscle layer is very thin and continuous. Ochetostoma indosinense

also differs in lacking a rectal caecum. Differences are also present in the colour

of live animals.

There seems little doubt, therefore, that this specimen from Park Rynie

differs in several respects from other known species of Ochetostoma that possess

three pairs of gonoducts. A closer study of additional specimens in the future may

well indicate that the present specimen represents an undescribed species.

ECHIURA FROM SOUTHERN AFRICA 73

ACKNOWLEDGEMENTS

I wish to thank Dr Jennifer A. Day of the Department of Zoology,

University of Cape Town, for her constructive criticism of the manuscript.

Financial support for this research was provided by the Council for Scientific and

Industrial Research. Thanks are also due to the University of Durban-Westville

for contributing to publication costs and for facilities. The assistance given by my

colleagues, Messrs K. S. Ganga and G. K. Moodley, during collecting trips is

gratefully acknowledged.

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M. E. & Toporovic, M. eds. Proceedings of the international symposium on the biology of

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Biological Research, Sinisa Stankovic.

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Ochetostoma Leuckart & Rippell (Echiura) together with a list of the species of the genus.

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with the description of two new species. Annals and Magazine of Natural History (13) 6 (1):

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FISCHER, W. 1922. Westindische Gephyreen. Zoologischer Anzeiger 55: 10-18.

FISHER, W. K. 1946. Echiuroid worms of the North Pacific Ocean. Proceedings of the United

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ABBREVIATIONS

av anal vesicle niv neuro-intestinal vessel

dv dorsal vessel oes oesophagus

gl gonostomal lip (D Wit pellet-like faeces

gon gonoduct p r int postsiphonal region of intestine

h heart po s gon postsetal gonoduct

ibm interbasal muscle pr s gon presetal gonoduct

int intestine rav right anal vesicle

irs intestinal ring sinus ir © rectal caecum

lav left anal vesicle rec rectum

mc mucous cap SS setal sac

ms mesenteric strand $ r int siphonal region of intestine

nc nerve cord VV ventral vessel

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

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Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, 1861: 87.

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

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R. BISESWAR

OCHETOSTOMA (ECHIURA) FROM

SOUTHERN AFRICA WITH A

DESCRIPTION OF A NEW SPECIES

As «

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

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

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

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

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SIBRARIES

SOUTH AFRICAN SPECIES OF THE GENUS

GERYON (CRUSTACEA, DECAPODA,

GERYONIDAE)

RAYMOND B. MANNING

L. B. HOLTHUIS

Cape Town Kaapstad

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SOUTH AFRICAN SPECIES OF THE GENUS GERYON

(CRUSTACEA, DECAPODA, GERYONIDAE)

RAYMOND B. MANNING

Smithsonian Institution, Washington, D.C.

L. B. HoLtuuts

Rijksmuseum van Natuurlijke Historie, Leiden

(With 6 figures)

[MS accepted 2 July 1987]

ABSTRACT

Three species of Geryon are recorded from South West Africa—Namibia and South Africa:

G. chuni Macpherson, G. macphersoni sp. nov., and G. maritae Manning & Holthuis. Geryon

ischurodous Stebbing is shown to be a species of Carcinoplax.

CONTENTS

PAGE

MMERO CME ON eperemenre Pree se Ade OS Mas, 6 Sue NRG eee a wie 4,16 48 77

The status of Geryon ischurodous Stebbing, 1923 ................. 78

The South African species of Geryon sensu stricto................. 80

INCKNOWICUSCIMEMtSar xan we A Are nk ang eae ernie eA ecadduc. asin oad 90

I CLERCTI CCS MR eee Bed Mir see aes eet Ms wee nen y ler 8 ww 91

INTRODUCTION

Until recently, three species of Geryon had been reported from South

African waters, Geryon quinquedens Smith (1879: 35), G. ischurodous Stebbing

(1923: 2), and G. maritae Manning & Holthuis (1981: 112), with most authors

identifying material from South Africa with the American G. quinquedens. Also

until relatively recently, many authors had identified material of Geryon from a

wide variety of localities with either G. affinis A. Milne Edwards & Bouvier,

1894, originally named from the north-eastern Atlantic, or G. quinquedens

Smith, described from the north-western Atlantic. In spite of the fact that

A. Milne Edwards & Bouvier (1894: 41-45) and Chace (1940: 38-40) showed

that G. affinis differed from G. quinquedens in having the dactyli of the walking

legs laterally compressed rather than dorso-ventrally depressed, some authors,

including Barnard (1950), considered the two species to be conspecific. Even

those who recognized the differences in the structure of the dactyl considered

Ann. S. Afr. Mus. 98 (3), 1988: 77-92, 6 figs.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

G. affinis to be widely distributed around the world. Thus G. affinis has been

recorded from the western Atlantic (Chace 1940), Japan (Sakai 1978), Australia

(Griffin & Brown 1976), and East Africa and the Valdivia Bank (Doflein 1904).

Manning & Holthuis (1981), in a review of the West African crabs, described

G. maritae, the first of six species of Geryon to be recognized in this decade. The

others are: G. chuni Macpherson (1983: 23), from South West Africa—Namibia;

G. fenneri Manning & Holthuis (1984: 666), from localities off Florida, U.S.A.;

G. erytheiae Macpherson (1984: 86), from the Valdivia Bank; G. gordonae Ingle

(1985: 90), from the north-eastern Atlantic; and G. inghami Manning & Holthuis

(1986: 366), from Bermuda.

As part of a long-term study of the genus Geryon, material of this genus in

the collections of the South African Museum was examined. The results of that

study are presented here.

The following abbreviations are used in the text: cb—carapace breadth;

cl—carapace length; fm—fathom; m—metre; mm—millimetre. Repositories

have been identified by the following abbreviations: BM—British Museum

(Natural History), London; ICM—Instituto de Ciencias del Mar, Barcelona;

MNHP—Muséum national d’Histoire naturelle, Paris; RMNH— Rijksmuseum

van Natuurlijke Historie, Leiden; SAM—South African Museum, Cape Town;

USNM—National Museum of Natural History, Smithsonian Institution,

Washington; ZMC— Zoological Museum, Copenhagen.

Co-ordinates in the text given in square brackets have been taken from

gazetteers of the United States Board on Geographic Names.

THE STATUS OF GERYON ISCHURODOUS STEBBING, 1923

Geryon ischurodous was described by Stebbing (1923: 2) from a single

specimen, cl 16mm, cb 26mm, taken from coral washed up on a beach at

Durban. No additional material identified with this species has been recorded in

the literature since then, and its status has remained uncertain. Barnard (1950:

293) commented on similarities between Stebbing’s species and Geryon trispino-

sus (Herbst). He concluded that ‘There seems to be little doubt that ischurodous

should become a synonym of trispinosus’.

Among the material of Geryon examined from the collections of the South

African Museum was a single small specimen taken off Natal in 700-680 m and

reported by Kensley (1977: 163) as Geryon sp. It proved to be identifiable with

Geryon ischurodous Stebbing.

In our opinion, Stebbing’s species did not belong in the genus Geryon. In

seeking to place it in another genus, we looked through a variety of accounts of

deep-sea crabs in the literature. Among these sources was the recent account by

Guinot & Richer de Forges (1981a, 1981b) of deep-sea crabs from the Indo-

Pacific. The similarity between the South African material of Geryon ischurodous

and a species taken in 600 m off the New Hebrides and described by Guinot &

Richer de Forges as Carcinoplax eurysternum was immediately recognized.

LSS

SOUTH AFRICAN SPECIES OF GERYON 719

Fig. 1. Carcinoplax ischurodous (Stebbing), Natal. Male, cl 4,1 mm.

Guinot & Richer de Forges (1981b: 251) pointed out that few species of

Carcinoplax have only two teeth on the anterolateral margin of the carapace, and

that few species have one of these teeth as an exorbital spine. Geryon ischurodous

has but two anterolateral teeth on the carapace, and one of these is a well-

developed exorbital tooth (Fig. 1). Further, the figures of C. eurysternum

provided by Guinot & Richer de Forges show a species with an inner spine on the

merus of the cheliped and an inner and outer spine on the carpus of the cheliped,

with the inner carpal spine bifurcate. The southern African specimens are

similarly spined and, although the smaller lacks the secondary spine on the inner

carpal spine, it is clearly shown on the type by Stebbing (1923, pl. 11). However,

the gonopods of the two species are slightly different— compare Stebbing’s figure

(1923, pl. 11, plp.2) with that of Guinot & Richer de Forges (1981), fig. 10H),

and the shape of the carapace is somewhat different in the two nominal species,

being broader in C. eurysternum.

Comparison of the specimen from off Natal and the type of Geryon

ischurodous with the original account of Carcinoplax eurysternum convinces us

that these three specimens are very closely related and that Carcinoplax

eurysternum Guinot & Richer de Forges, 1981, may prove to be a junior synonym

of Geryon ischurodous Stebbing, 1923. The correct name for Stebbing’s species is

Carcinoplax ischurodous (Stebbing, 1923).

We have examined the following specimens of Carcinoplax ischurodous from

South African waters:

BM 1928.12.1.104: Durban [29°51'S 31°01'E]; on a coral washed up, leg.

Mr H. W. Bell Marley; 3/20, Stebbing collection; 1 male, cl 16 mm, cb 26 mm;

holotype of Geryon ischurodous Stebbing.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-—A15289: off Natal, 27°14'S 32°54'E, 700-680 m, Agassiz trawl,

20 May 1976, R.V. Meiring Naude station SM 67; 1 male, cl 4,1 mm, cb 5,9 mm.

Synonymy of Carcinoplax ischurodous:

Geryon ischurodous Stebbing, 1923: 2, pl. 11. Barnard, 1950: 292, fig. 54k. Guinot, 1967: 276 [no

record].

Geryon sp.: Kensley, 1977: 163.

THE SOUTH AFRICAN SPECIES OF GERYON SENSU STRICTO

Geryon chuni Macpherson, 1983

Figs) 2, 3516€

Geryon quinquedens: Stebbing, 1905: 36; 1910: 313. Barnard, 1950: 291, fig. 54f-i [part.].

Geryon affinis: Takeda, 1976: 64, fig. S. Afr.—69 [colour]. [Non G. affinis A. Milne Edwards &

Bouvier, 1894.]

Geryon chuni Macpherson, 1983: 23, figs 10-15, 17a—c.

Material

South West Africa—Namibia

RMNH (no number): off South West Africa—Namibia, 23°50’'S 13°03,6’E,

761-768 m, mud, 30 May 1982, Valdivia I, station P—26; 1 male, 1 female.

South Africa

USNM 205337: Cape Columbine [32°49’S 17°51’'E] region; 1 female. SAM-—

A19513: WNW of Dasseneiland [= Dassen Island, 33°26’S 18°05’E],

160-180 fms (293-329 m), 25 February 1965, collected by S. Kannemeyer;

1 female. SAM—A12073: off Dasseneiland, 165 fms (302 m); 1 male, 1 ovigerous

female. SAM-—A12207: west of Slangkop [= Slang Kop Point, 34°09’S 18°19’E],

200 fms (366m), 24 February 1965, collected by S. Kannemeyer: 1 male,

3 females. BM 1928.12.1: Cape Point Lighthouse [Cape Point = 34°21’S

18°29’E], 470 fms (860 m), Stebbing collection; 1 male, 1 female, 3 juveniles.

SAM-A836, A837, A1626: off Cape Point (three lots with different bearings

combined), 345, 500, and 760 fms (631, 915, and 1 391 m); 6 juveniles. SAM-—

A675, A6075: off Cape Point Lighthouse (two lots with different bearings

combined), 360 and 470 fms (659 and 860m), S.S. Pieter Faure, stations

P.F. 6004, 16655; 2 males, 2 females (1 ovigerous). SAM—A15385: west of Cape

Point, 250 fms (458 m); 2 males, 1 female. USNM 228311: 34°49’S 18°17’E,

515 m, 1985, R.V. Africana, collected by R. Melville-Smith; 1 male, 1 female.

SAM-A676, A681: off Table Mountain [33°58’S 18°25’E], 250 fms (458 m) and

39°49'S 21°14’E, 560 fms (1 025 m) (two different lots combined); 5 females

(1 ovigerous). ZMC (no number): Agulhas Bank [35°30’S 21°00’E]; exchange

from South African Museum; 2 males.

Description

A large Geryon, cl to 95 mm, cb to 114 mm in adults, with five anterolateral

teeth on the carapace and laterally compressed dactyli on the walking legs.

Carapace about 1,2 times broader than long. Median pair of frontal teeth

SOUTH AFRICAN SPECIES OF GERYON 81

Fig. 2. Geryon chuni Macpherson, Agulhas Bank. A-D. Male, cl 24mm. A. Caxapace.

B. Orbit, ventral view. C. Fifth leg. D. Abdomen. E. Male, cl 53 mm. Gonopod.

Fig. 3. Juveniles of Geryon chuni Macpherson, off Cape Point. A. cl 19,5 mm, dorsal

view. B—E. Outline of carapace. B.cl13 mm. C.cl11mm. D.cl10mm. E. cl 9 mm.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

separated by deep V-shaped sinus. Distance between submedian frontal teeth less

than distance between them and lateral frontal teeth. Second and fourth

anterolateral teeth often obsolete in adults, fourth smallest of all; distance from

first to third tooth less than that from third to fifth tooth. Carapace lacking distinct

raised ridge mesial to fifth anterolateral tooth, granular posterolaterally, smooth

dorsally. Suborbital tooth well developed, extending to level of lateral frontal

teeth. Cheliped with sharp tooth subdistally on upper margin and with distal

dorsal tooth or lobe on merus; carpus roughened dorsally, with distal outer spine

or angled lobe, anterior margin smooth; propodus with at most distal angled

projection. Meri of walking legs usually with distal, dorsal tooth or projection.

Dactyli of walking legs compressed, height at midlength greater than width. Fifth

leg: merus about 4,5 (range 3,8—5,2) times longer than high, length about half

carapace width; carpus with line of denticles or low spinules dorsally; propodus

length about four times height, subequal to dactylus in length (longer in some,

shorter in others).

Size

Carapace lengths of males 24 to 92 mm, of non-ovigerous females 47 to

69 mm, of ovigerous females 54 to 65 mm, of juveniles 8,5 to 19,5 mm. Carapace

widths of males 32 to c.100 mm, of non-ovigerous females 62 to 87 mm, of

ovigerous females c.68 to 85 mm, of juveniles 12 to 26 mm. Macpherson (1983)

reported the following range of measurements in his material: males, cl 51 to

114 mm, cb 61 to 95 mm; females, cl 56 to 73 mm, cb 68 to 86 mm; ovigerous

female, cb 86 mm.

Depth range

The specimens were taken in depths ranging between 302 m and 1 391 m.

Three records were from about 300 m, two from about 400 m, one from 500 m,

two from 600 m, one from 700 m, three from about 900 m, one from about

1 000 m, and one from 1 391 m. All of the juveniles, cl 19,5 mm or less, came

from depths in excess of 600 m.

Remarks

Adults of G. chuni can be distinguished at once from both G. maritae and

G. macphersoni by the laterally compressed dactyli on the walking legs and by the

larger suborbital spine. This species further differs from G. maritae in having a

distinct dorsal spine on the meri of the walking legs, especially the posterior two

pairs, it is a smaller species than either G. maritae or G. macphersoni, and it is a

much smoother species than the latter.

At smaller sizes, differences in the shape of the dactyli of the walking legs

become less clear. The series of juveniles from SAM-—A836 all lack distal spines

on the meri of the walking legs. The three juveniles from the lot of five specimens

from the collection of the British Museum all lack distal spines on the meri of the

walking legs as well as erect spinules on the carpi of the walking legs; the meral

SOUTH AFRICAN SPECIES OF GERYON 83

spines are present, at least on the last legs, in the two larger specimens in the same

lot.

The configuration of the carapace spines in several specimens from lot

SAM-A836 are shown in Figure 3.

The specimens reported by Stebbing (1905, 1910) and Barnard (1950) were

examined. Part of the material reported by Barnard, at least the specimens from

off Cape Point, are identifiable with this species; his figure 54f-h probably is

based on the juveniles from lot SAM—A836. His material from East London is

referred below to G. macphersoni sp. nov. As Barnard did not list individual lots,

it is not possible to determine exactly which specimens that he might have studied

actually belong here, although it is likely that he saw all of the early specimens in

the collection.

The range of G. chuni overlaps with that of G. maritae off South West

Africa—Namibia and with that of G. macphersoni sp. nov. off South Africa. In

the material examined, G. chuni was found together with G. macphersoni

sp. nov. in the lots from off Dasseneiland in 293-329 m, that taken by the

R.V. Africana in 515m, that from off Slangkop in 366m, and at Cape

Columbine. According to R. Melville-Smith (in litt. August 1985), this species

and G. macphersoni were taken together off Cape Columbine at 32°25’S 16°29°E,

in 496 m.

The species reported from South Africa by Takeda (1976) probably is this

species. His colour figure shows a reddish-tan crab with blunt anterolateral teeth

and distal spines on the meri of the walking legs, and the dactyli of the walking

legs appear to be compressed.

Distribution

Off south-western coast of Africa, from 23°50’S, off South West Africa—

Namibia, to Agulhas Bank, South Africa.

Geryon macphersoni sp.-nov.

Figs 4, 5, 6A-B

Geryon quinquedens: Barnard, 1950: 291, fig. 54f-i [part.]. Kensley, 1977: 163. Paula e Silva,

1985: 8 [p.p.]. [Non Geryon quinquedens Smith, 1879.]

Geryon sp.: Kensley, 1981a: 61; 1981b: 41 [no records].

Material

South Africa

SAM-A12206: holotype, WNW of Dasseneiland [= Dassen Island, 33°26'S

18°05’E], 160-180 fms (293-329 m), 25 February 1965, collected by S. Kanne-

meyer; 1 male.

USNM 221699: paratype, ?0ff Cape Columbine [32°49’S 17°51’E], c.500 m;

1 male. SAM-—A19514: paratype, W of Slangkop [= Slang Kop Point, 34°09'S

18°19’E], 200 fms (366 m), 24 February 1965, collected by S. Kannemeyer;

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

1 male. USNM 228316: paratypes, 34°49'S 18°17’E, 515 m, 1985, R.V. Africana

(R. Melville-Smith); 1 male, 1 female. ICM (no number): paratypes, 3 males,

1 female (data as for USNM 228316). SAM-—A19515: paratype, off Table

Mountain [33°58’S 18°25'E], 250 fms (458 m) and 39°49’S 21°14’E, 560 fms

(1 025 m) (two different lots combined); 1 male. RMNH no. Crust.D.36594:

paratype, off Cape Town, R. Melville-Smith; 1 male. SAM—A680: paratype, off

Bashee River Beacon [Bashee River = 32°15’S 28°54’E], 300 fms (549 m),

S.S. Pieter Faure, station P.F. 12650; 1 female (dry). SAM—A16773: paratypes,

32°15'S 29°09’E, 580m, D.B. trawl, R.V. Meiring Naude station SM 233,

25 June 1979; 2 males, 2 females. SAM-—A15890: paratype, 30°32’'S 30°52’E,

900-625 m, beam trawl, R.V. Meiring Naude, station SM 121, 10 May 1977;

1 female. SAM-—A15282: paratypes, off Natal, 28°21’S 32°34’E, 775-825 m,

R.V. Meiring Naude station SM 38, 28 May 1975; 3 males, 1 female. ZMC (no

number): paratypes, off Natal, 25°20’S 35°17’E, 680-730 m, sand bottom,

Galathea, station 203, 21 February 1951; 1 male, 1 female.

Mozambique

RMNH no.Crust.D.36593: paratype, off Mozambique, P-—51, I. Riera;

1 female.

Description

A large Geryon, cl to 105 mm, cb to 126 mm in adults, with five antero-

lateral teeth on the carapace and dorso-ventrally depressed dactyli on the walking

legs. Carapace about 1,2 times broader than long. Median pair of frontal teeth

separated by shallow sinus. Distance between submedian frontal teeth less than

distance between submedian and lateral frontal teeth. Second and fourth

anterolateral teeth reduced, fourth often completely obsolete, distance from first

to third tooth subequal to distance from third to fifth. Carapace with distinct

raised ridge mesial to fifth anterolateral tooth, and with strong granulation on

branchial, cardiac, and protogastric regions, especially in males. Suborbital teeth

low, falling short of level of lateral frontal teeth. Cheliped rough, surface with

sharp, erect granules; upper margin of merus with sharp subdistal spine; carpus

rough, anterior margin, lateral to inner spine, lined with sharp granules;

propodus rough dorsally, unarmed distally, with an outer granulate ridge. Meri of

walking legs usually with distal, dorsal spine. Dactyli of walking legs dorso-

ventrally depressed, width near midlength greater than height. Fifth leg: merus

3,9 (range 3,6—5,0) times longer than high, with distal dorsal spine, length 0,5—0,6

carapace width; propodus with line of erect spinules dorsally, length about 3,5

times height, slightly longer than dactylus.

Size

Carapace lengths of males 23 to 105 mm, of females 30 to 83 mm; carapace

widths of males 34 to 126 mm, of females 40 to 100 mm. Paula e Silva (1985)

reported females with cb 105 mm and males with cb more than 120 mm from the

population off Mozambique.

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SOUTH AFRICAN SPECIES OF GERYON 87

Depth range

The specimens were taken in depths of 293-329 m to 900-625 m; inter-

mediate depths included 366 m, 458 m, c. 500 m, 549 m, 580 m, 680-730 m, and

775-825 m. The smallest specimens were taken in depths of 900-625 m and

775-825 m. Paula e Silva (1985) reported that this species was taken in

250-850 m off southern Mozambique.

Remarks

This species can be distinguished from both G. chuni and G. maritae by the

extent of granulation on the carapace, chelipeds, and dorsal margins of the

walking legs. The smallest specimen of G. macphersoni examined is much

rougher dorsally than the largest specimen we have seen of G. chuni. Geryon

macphersoni resembles G. maritae and differs from G. chuni in having dorso-

ventrally depressed dactyli on the walking legs. It further differs from G. maritae

in having a distal spine on the meri of the walking leg, a distinct line of erect

spinules on the carpus of the walking legs, and tubercles or denticles on the

anterior margin of the carpus of the cheliped.

In roughness of the carapace, this species resembles G. granulatus Sakai

(1978: 11) from Japan, and the Japanese species also has depressed dactyli on the

walking legs. Geryon granulatus, however, lacks the distal spines on the meri of

the walking legs and those legs are much smoother dorsally, and the carapace has

a much more rounded shape and is more inflated dorsally as well.

Barnard (1950: 291) considered G. paulensis Chun (1903: 531), taken in

2 068 m, north of St. Paul and New Amsterdam, Indian Ocean, to be the juvenile

of his G. quinquedens. That species, however, known only from the type, has

compressed dactyli on the walking legs, which are very long and slender, and has

very long first, third, and fifth anterolateral spines on the carapace. Geryon

paulensis appears to be distinct from any of the species known from southern

African waters.

The specimen recorded by Barnard (1950) from off East London [33°02'S

27°55'E] in 300 fms (549 m) is probably the specimen recorded here from off the

Bashee River. The specimens reported by Kensley (1977, 1981a) were also

examined and both are referable to this species.

This species apparently occurs together with G. chuni in the area around the

Cape but has a much greater eastward distribution than G. chuni, which has not

been taken from east of Agulhas Bank.

The specimen shown in Figure 5 has a branchial parasite on its right side.

The male from off Cape Town carries small lepadid barnacles on the mesial side

of the legs and the base of the carapace. The female from Mozambique shows two

balanid barnacles on the right cheliped.

Etymology

We are pleased to dedicate this species to our colleague Enrique Macpherson

of the Instituto de Ciencias del Mar, Barcelona, whose careful work has recently

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

revealed the existence of two previously undescribed species of Geryon, G. chuni

and G. erytheiae.

Distribution |

Southern Africa, from off Cape Columbine to 35°17’E, off Natal, and of

Mozambique.

Geryon maritae Manning & Holthuis, 1981

Fig. 6D

Geryon quinquedens Beyers & Wilke, 1980: 9 [non G. quinquedens Smith, 1879].

Geryon maritae Manning & Holthuis, 1981: 112, figs 24a, 25, 26 [part., not references to Doflein

1903, 1904=Geryon erytheiae Macpherson, 1984]. Macpherson, 1983: 22, figs 16-17d.

Melville-Smith, 1983: 123, figs 2, 3; 1985: 55, fig. 2 (left); 1986: 257, figs 1-3; 1987:

143 [p.p.].

Material

South West Africa—Namibia

MNHP (no number): off South West Africa—Namibia, 18°41,5’S 11°24,5’E,

480 m, 27 June 1971, ‘Walda’—0067—CMO3; 1 male. USNM 205342: ?off South

West Africa—Namibia, 19°30'S, 400 m; 1 ovigerous female. SAM (no number):

just north of Liideritz [26°39’S 15°09’E], 600 m; 1 male.

Description

A very large Geryon, adults with cl of up to 140 mm, cb to more than

160 mm, with five anterolateral teeth on the carapace and dorso-ventrally

depressed dactyli on the walking legs. Carapace about 1,1 times broader than

long. Median pair of frontal teeth separated by wide sinus, base of sinus at about

level of lateral frontal teeth, submedian frontal teeth distinctly over-reaching

lateral frontal teeth. Distance between submedian frontal teeth less than distance

from each to lateral frontal tooth. Second and fourth anterolateral teeth greatly

reduced, fourth often completely obsolete, distance between first and third tooth

less than distance between third and fifth tooth. Carapace with short, oblique

ridge on each side mesial to fifth anterolateral tooth. Surface of carapace variably

granular, granulations particularly well developed on branchial regions. Suborbi-

tal spine very short. Cheliped with blunt subdistal lobe on upper margin of merus,

but lacking distal projection there, carpus lacking outer spine, propodus lacking

distal dorsal spine. Meri of walking legs with no trace of distal dorsal spine.

Dactyli of walking legs dorso-ventrally depressed, width at midlength greater

than height. Fifth leg: merus about four (3,8—4,3) times as long as high, length

about two-thirds carapace width; carpus with irregularly tuberculate dorsal ridge,

lacking line of erect tubercles; propodus length about four times height, shorter

than dactylus.

Size

Carapace length of males 85 and 112 mm, of ovigerous female 82 mm.

Carapace widths of males 97 and 131 mm, of ovigerous female 97 mm. Manning

SOUTH AFRICAN SPECIES OF GERYON

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90 ANNALS OF THE SOUTH AFRICAN MUSEUM

& Holthuis (1981) recorded males as large as cl 140 mm, cb 160 mm, and females

as large as cl 89 mm, cb 95 mm. Beyers & Wilke (1980) reported males as large

as cb 172 mm. Macpherson (1983) reported males with cl 63 to 155 mm, females

with cl 54 to 86 mm.

Depth range

The specimens were taken in 400, 480, and 600 m. Manning & Holthuis

(1981) reported that the species had been recorded from depths between 100 and

936 m. Beyers & Wilke (1980) surveyed the fishery for this crab off South West

Africa—Namibia in depths between 295 and 849 m. Macpherson (1983) studied

material that had been collected between 270 and 615 m.

Remarks ie

Geryon maritae agrees with G. macphersoni and differs from G. chuni in

having the dactyli of the walking legs strongly depressed dorso-ventrally, wider

than high. In G. chuni, which is a smaller species, the dactyli of the walking legs

are distinctly compressed, higher than wide. Geryon chuni further differs from

G. maritae in having a distinct dorsal spine distally on the merus of each walking

leg. Differences between this species and G. macphersoni are discussed under the

account of that species.

In the southern extreme of its range, off southern South West Africa—

Namibia, the distribution of G. maritae overlaps that of G. chuni, which is known

to occur off southern South West Africa—Namibia and off South Africa.

In the fisheries literature, this species is known as the ‘deep-sea red crab’, the

common name now in use for G. quinquedens proper. This name probably is in

use because G. quinquedens was believed to occur in South African waters. New

common names should be coined for all three species of Geryon from South

Africa.

Distribution

Atlantic coast of western Africa, from numerous localities between South

Morocco to the north and South West Africa—Namibia to the south. Beyers &

Wilke (1980) and Melville-Smith (1983, 1985, 1986, 1987) reported on aspects of

the fisheries for this species off South West Africa-—Namibia, and Macpherson

(1983) provided additional records from off South West Africa—Namibia.

ACKNOWLEDGEMENTS

We thank our colleagues at the museums in Copenhagen, London, and Paris

for allowing us to work with their material. E. Macpherson, Instituto de Ciencias

del Mar, Barcelona, provided us with valuable comparative material.

R. Melville-Smith, Sea Fisheries Research Institute, Cape Town, gave us

information on South African Geryon and references, including some that we

might not otherwise have seen. We thank Brian Kensley, Smithsonian Institution,

SOUTH AFRICAN SPECIES OF GERYON 91

and Enrique Macpherson, for their comments on a draft of the manuscript. We

appreciate the help and patience of Mrs M. G. van der Merwe, South African

Museum, who made that collection available to us. Roy Kropp, University of

Maryland, took the photographs for us, and all of the figures were prepared by

Lilly King Manning. Our work on Geryon has been supported in part by FAO,

Rome.

REFERENCES

BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and

shrimps). Annals of the South African Museum 38: 1-837.

Beyers, C. J. DE B. & WILKE, C. G. 1980. Quantitative stock survey and some biological and

morphometric characteristics of the deep-sea red crab Geryon quinquedens off South West

Africa. Fisheries Bulletin, Division of Sea Fisheries, South Africa 13: 9-19.

Cuace, F. A. yr. 1940. The brachyuran crabs. Reports on the scientific results of the Atlantis

expeditions to the West Indies, under the joint auspices of the University of Havana and

Harvard University. Torreia 4: 1-67.

Cuun, C. 1903. Aus den Tiefen des Weltmeeres. 2nd ed. Jena: Gustav Fischer.

CrosnigR, A. 1970. Crustacés décapodes brachyoures et macroures recueillis par l’‘Undaun-

ted’ au sud de l’Angola. Description de Scyllarus subarctus sp. nov. Bulletin du Muséum

national d’ Histoire naturelle, Paris (2) 41 (5) [for 1969]: 1214-1227.

DoFLEIN, H. 1903. Die Augen der Tiefseekrabben. Biologisches Zentralblatt 23 (16, 17):

570-593.

DoF ein, H. 1904. Brachyura. Wissenschaftliche Ergebnisse des Deutschen Tiefsee-Expedition

auf dem Dampfer ‘Valdivia’ 1898-1899 6: i-xiv, 1-314.

GriFFIN, D. J. G. & Brown, D. E. 1976. Deepwater decapod Crustacea from eastern

Australia: brachyuran crabs. Records of the Australian Museum 30 (11): 248-271.

Gurinot, D. 1967. La faune carcinologique (Crustacea, Brachyura) de l’océan Indien occidental

et de la Mer Rouge. Catalogue, remarques biogéographiques et bibliographie. Mémoires de

l'Institut fondamental d’ Afrique noire 77: 235-352.

Guinot, D. & RICHER DE ForGEs, B. 198la. Crabes de profondeur, nouveaux ou rares, de

V’Indo-Pacifique (Crustacea, Decapoda, Brachyura). Bulletin du Muséum national dHis-

toire naturelle, Paris (4) 2 (A, 4) [for 1980]: 1113-1153.

Guinot, D. & RICHER DE ForGEs, B. 1981b. Crabes de profondeur, nouveaux ou rares, de

l’Indo-Pacifique (Crustacea, Decapoda, Brachyura). Bulletin du Muséum national d His-

toire naturelle, Paris (4) 3 (A, 1): 227-260.

INGLE, R. W. 1985. Geryon gordonae sp. nov. (Decapoda Brachyura, Geryonidae) from the

northeastern Atlantic Ocean. Crustaceana 48 (1): 88-98.

KENSLEY, B. 1977. The South African Museum’s Meiring Naude cruises. Part 2. Crustacea,

Decapoda, Anomura and Brachyura. Annals of the South African Museum 72 (9):

161-188.

KENSLEY, B. 1981a. The South African Museum’s Meiring Naude cruises. Part 12. Crustacea,

Decapoda of the 1977, 1978, 1979 cruises. Annals of the South African Museum 83 (4):

49-78.

KENSLEY, B. 1981b. On the zoogeography of southern African decapod Crustacea, with a

distributional checklist of the species. Smithsonian Contributions to Zoology 338: 1-64.

MacPHERSON, E. 1983. Crustéceos decapodos capturados en las costas de Namibia. Resultados

Expediciones cientificas (Supplement to Investigacion Pesquera, Barcelona) 11: 3-80.

MAcPHERSON, E. 1984. Crustdceos decApodos del Banco Valdivia (Atlantico sudoriental).

Resultados Expediciones cientificas (Supplement to Investigacion Pesquera, Barcelona) 12:

39-105.

Manninc, R. B. & Hottuuis, L. B. 1981. West African brachyuran crabs (Crustacea:

Decapoda). Smithsonian Contributions to Zoology 306: i—xii, 1-379.

MANNING, R. B. & Hottuuts, L. B. 1984. Geryon fenneri, a new deep-water crab from Florida

(Crustacea: Decapoda: Geryonidae). Proceedings of the Biological Society of

Washington 97 (3): 666-673.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

MANNING, R. B. & Ho.tuHuis, L. B. 1986. Notes on Geryon from Bermuda, with the

description of Geryon inghami, new species (Crustacea: Decapoda: Geryonidae). Proceed-

ings of the Biological Society of Washington 99 (2): 366-373.

MELVILLE-SMITH, R. 1983. Abundance of deep-sea red crab Geryon maritae in South West

African waters from photography. South African Journal of Marine Science 1: 123-131.

MELVILLE-SMITH, R. 1985. Density distribution by depth of Geryon maritae on the northern

crab grounds of South West Africa/Namibia determined by photography in 1983, with notes

on the portunid crab Bathynectes piperitus. South African Journal of Marine Science 3:

55-62.

MELVILLE-SMITH, R. 1986. Red crab (Geryon maritae) density in 1985 by the technique of

effective area fished per trap on the northern fishing grounds off South West Africa. South

African Journal of Marine Science 4: 257-263.

MELVILLE-SMITH, R. 1987. Movements of deep-sea red crab (Geryon maritae) off South West

Africa/Namibia. South African Journal of Zoology 22 (2): 143-152.

MILNE Epwarps, A. & Bouvier, E.-L. 1894. Crustacés décapodes provenant des campagnes

du yacht l’Hirondelle (1886, 1887, 1888). Premiére partie. Brachyures et anomoures.

Résultats des campagnes scientifiques accomplies sur son yacht par Albert I°’, prince souverain

de Monaco 7: 1-112.

PAULA E SILVA, Rur DE. 1985. The deep crab—Geryon quinquedens, first notes on its biology

off Mozambique. Revista de Investigacao Pesqueira, Maputo 13: 5-25.

SakAl, T. 1978. Decapod Crustacea from the Emperor Seamount Chain. Researches on

Crustacea 8 (supplement): 1-39.

SmiTH, S. I. 1879. The stalk-eyed crustaceans of the Atlantic coast of North America north of

Cape Cod. Transactions of the Connecticut Academy of Arts and Sciences 5 (1): 27-136.

STEBBING, T. R. R. 1905. South African Crustacea. Part III. Marine Investigations in South

Africa 4: 21-123.

STEBBING, T. R. R. 1910 General catalogue of South African Crustacea. Annals of the South

African Museum 6 (4): 20-584.

STEBBING, T. R. R. 1923. Crustacea of Natal. Report. Fisheries and Marine Biological Survey,

Union of South Africa (for 1922) 3 (Special Report 3): 1-16.

TAKEDA, M. 1976. Geryon affinis. Colored illustrations of bottomfishes collected by Japanese

trawlers 2: 1-188. Tokyo: Japan Deep Sea Trawlers Association.

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Family Nuculanidae

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

RAYMOND B. MANNING & L. B. HOLTHUIS

SOUTH AFRICAN SPECIES OF THE GENUS

GERYON (CRUSTACEA, DECAPODA,

GERYONIDAE)

—

‘OF THE SOUTH AFRICAN

MUSEUM

CAPE TOWN

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

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

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

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

May 1988 Mei

Part 4 Deel

, SMITHSON

JUL 28 1988

: LIBRARIES

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A NEW SPECIES OF MYSIDOPSIS (MYSIDACEA)

FROM COASTAL WATERS OF SOUTHERN

AFRICA AND A KEY TO THE KNOWN

SPECIES FROM THE SUBCONTINENT

T. H. WOOLDRIDGE

Cape Town Kaapstad

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A NEW SPECIES OF MYSIDOPSIS (MYSIDACEA) FROM COASTAL

WATERS OF SOUTHERN AFRICA AND A KEY TO THE KNOWN

SPECIES FROM THE SUBCONTINENT

T. H. WooLpRIDGE

Department of Zoology, Institute for Coastal Research,

University of Port Elizabeth

(With 5 figures)

[MS accepted 1 September 1987]

ABSTRACT

A new species of Mysidopsis was collected from warm temperate coastal waters on the east

coast of southern Africa. Morphological features distinctive to M. buffaloensis sp. nov. include

the narrowly triangular and apically acute rostrum and the armament along the inner margin of

the uropod, these spines on the endopod extending only to the midlength of the segment. The

lateral margins of the telson are also partly unarmed and the segment is posteriorly narrowed and

tapering, terminating in two long spines. A key to the known species of Mysidopsis in southern

Africa is given.

CONTENTS

PAGE

INthodactio merrier ee aio shined as sean ance aes Gal 93

DYESS OCOD Se ra eres ee or 94

INCI ARK SHI eR Ray ee EOE, cree one eens ioe Vahey 6 aes 100

Key to the southern African species of Mysidopsis ............ 100

PNCKMOWICGSEMEMES eek ey aire tase aan = ci ease ya cy teal he Fan ahha sans talat 102

I NEMESMNGES 6 a5 SS eg Gere ee ey eRe cee ae eet tee eee 102

INTRODUCTION

Bentho-pelagic mysid shrimps are an important subtidal component of sandy

beaches in southern Africa (Wooldridge 1983). Beach mysids are usually

represented by the genus Gastrosaccus, with other genera sometimes present.

Such multigeneric assemblages usually occur when beaches are interrupted by

other substrata or are influenced by estuaries.

Dredging in the shallow surf (1,5 m depth) adjacent to a concrete harbour

pier bordering Orient Beach, East London, revealed the presence of an

undescribed species of Mysidopsis. Underwater observations showed them to be

present in large numbers on the surface of the sand and up to 25 m from the pier.

Associated with these mysids were relatively low numbers of Gastrosaccus

psammodytes, a species common along the beaches on the west and south coasts

of southern Africa (Brown & Talbot 1972; Wooldridge 1983). Further distant

from the pier G. psammodytes was the principal species, with G. bispinosa

(unpublished pers. obs.) occasionally recorded.

Ann. S. Afr. Mus. 98 (4), 1988: 93-103, 5 figs.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

DESCRIPTION

Mysidopsis buffaloensis sp. nov.

Figs 1-5

Holotype

SAM-A39552, South African Museum, Cape Town. Adult female from

Orient Beach, East London (33°01’S). Collected on 9 March 1985 by

T. Wooldridge.

Paratypes

SAM-—A39553, South African Museum, Cape Town. Numerous adult males

and adult females. Collection data as for holotype.

Etymology

The name refers to the robust nature of the body of this mysid and the

proximity of the Orient Beach to the Buffalo River harbour, East London.

Description

General form robust. Total length of adult females 10,3-14,0 mm (mean of

25 specimens: 12,1 mm); adult males 9,0-11,5 mm (mean of 20 specimens:

10,5 mm).

Rostrum acutely pointed, extending to base of third segment of antennular

peduncle, or further in large specimens (Fig. 1A). Anterolateral carapace

margins rounded; posterior border emarginate leaving last two thoracic segments

exposed in dorsal view. Antennular peduncle (Figs 1A, B) extending barely

beyond eyes, first segment equal in length to second and third combined. Male

lobe large and hirsute, typical for the genus.

Antennal sympod (Fig. 1C) with a strong spine on outer distal margin.

Margins of scale setose, about four times as long as greatest width. Outer margin

of scale straight, inner margin convex. A small suture present in distal one-

twentieth.

Mandible (Fig. 2A) with incisor process and lacinia mobilis well developed.

Molar process reduced. Palp three-segmented, second segment large and

expanded, producing a distinct ‘heel’ along outer margin of segment. Maximum

width about half the segment length. Terminal segment two and one half times as

long as wide, armed at apex with a strong barbed spine. A row of about seven

smaller barbed spines on inner margin on distal half of segment. Setation of palp

as shown.

Maxilla (Fig. 2B) with long, slender exopod armed along outer margin with

about 13 graduated setae. Second endopod segment truncate, two and one-half

times as long as wide. Armament of endopod and endites as illustrated.

Maxillule (Fig. 2C) with three setae on inner lobe. Outer lobe with about

nine barbed spines at distal end. First thoracic limb (Fig. 3A) typical for the

genus, endopod robust with second and third joints fused. Dactylus small, slightly

A NEW SPECIES OF MYSIDOPSIS FROM SOUTHERN AFRICA 95

Fig. 1. Mysidopsis buffaloensis sp. nov. A. Carapace in dorsal view. B. Antennule.

C. Antenna.

wider than long, and armed with long claw three times length of segment. Exopod

with first joint expanded, outer distal angle approximately 90 degrees. Setation of

segments as illustrated (Fig. 3A).

Second thoracic limb (Fig. 3B) with endopod more slender than in first

thoracic limb. Merus three times as long as wide, carpo-propodus shorter and

more robust than merus. Dactylus slightly longer than wide, with claw equal in

length to segment.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

(

A-C 0,5mm

Fig. 2. Mysidopsis buffaloensis sp. nov. A. Mandible. B. Maxilla. C. Maxillule.

Remaining thoracic appendages slender, subequal. Carpo-propodus of three

subsegments, third shorter in length than first and second combined. Dactylus

small, nail long and slender (Fig. 4A).

Pleopod 1 and remaining pleopods in female (Fig. 3C) simple setose plates,

which become progressively longer posteriorly.

Pleopod 1 in male (Fig. 3D) with sympod sub-rectangular. Endopod one-

third length of exopod, bearing a distinct lobe at base. Lobe with five small

terminal spines. Second male pleopod with rectangular sympod (Fig. 4B). Rami

seven-segmented, endopod subequal in length to exopod. Basal lobe on endopod

expanded in distal half and bearing five small terminal setae. Fourth male

pleopod (Fig. 4C) similar in general form to pleopod 2, terminal exopod segment

bearing a long, curved modified seta (Figs 4C, 5A) barbed along distal half.

A NEW SPECIES OF MYSIDOPSIS FROM SOUTHERN AFRICA

Fig. 3. Mysidopsis buffaloensis sp. nov. A. First thoracic appendage. B. Second thoracic

appendage. C. First pleopod of female. D. First pleopod of male.

ANNALS OF THE SOUTH AFRICAN MUSEUM

C. Fourth pleopod of male.

Fig. 4. Mysidopsis buffaloensis sp. nov. A. Eighth thoracic appendage. B. Second pleopod of

male.

A NEW SPECIES OF MYSIDOPSIS FROM SOUTHERN AFRICA

Fig. 5. Mysidopsis buffaloensis sp. nov. A. Terminal exopod segment of fourth pleopod

of male. B. Uropod. C. Telson.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Subterminal barbs nearly twice as long as others. A second seta also present on

seventh exopod segment (Fig. 5A), not reaching midpoint of barbed seta. Three

minute spinules near base of barbed seta. Remaining setation as illustrated.

Uropod (Fig. 5B) with exopod having entire margin setose; outer margin

straight, the inner convex. Endopod as wide as exopod and about two-thirds as

long. Inner endopod margin armed with row of 25-30 closely-set graduated

spines, the smallest opposite statocyst. Spine row extending to mid-region of

segment, proximal spines about one-quarter the length of those more distal.

Outer endopod margin with a dense patch of setae proximally, distal two-thirds of

this margin with about 12 relatively short setae set dorso-laterally to the longer

marginal setae.

Telson (Fig. 5C) triangular, almost twice as long as basal width. Armament

along lateral margins discontinuous in anterior half, unarmed part about one-

quarter length of telson. Lateral margins slightly concave in mid-region,

becoming convex towards the pointed apex. Four lateral spines anterior to

unarmed part of telson, slightly longer than posterior spines, which number about

forty. Terminal pair of spines at least twice as long as those adjacent to them.

REMARKS

Mysidopsis buffaloensis sp. nov. represents the eighth species of the genus

currently known from coastal waters around southern Africa. Previously

described are M. major (Zimmer, 1912), M. schultzei (Zimmer, 1912) and

M. similis (Zimmer, 1912), from Liideritz Bay. Their range extends southwards

(Tattersall 1955) and along the south-east coast as far as Algoa Bay (Wooldridge

1983). Mysidopsis camelina Tattersall, 1955, is recorded from False Bay only,

while M. eremita Tattersall, 1962, is represented by one ovigerous female

collected on the west coast.

Mysidopsis bispinosa Tattersall, 1969, and M. suedafrikana Tattersall, 1969,

were each described from a single damaged specimen and had originally been

reported by him (Tattersall 1958) as Mysidopsis species A and Mysidopsis species B,

respectively. The current known range of M. bispinosa extends from Table Bay

(Tattersall 1969) eastward to Algoa Bay (Wooldridge 1983), while M. sued-

afrikana is recorded from Langebaan Bay and False Bay (Tattersall 1969).

Species of Mysidopsis presently known from southern Africa are readily

distinguishable from Mysidopsis buffaloensis sp. nov.; the key features separating

them are recorded below.

KEY TO THE SOUTHERN AFRICAN SPECIES OF MYSIDOPSIS

A key to the known species of adult Mysidopsis recorded from coastal waters

around southern Africa is presented. Characteristics included are based on

published descriptions, so that in some cases data refer to relatively few

specimens or even single animals.

Ike

A NEW SPECIES OF MYSIDOPSIS FROM SOUTHERN AFRICA 101

One or two distinct nodules along mid-dorsal line of carapace............. zy

— Carapace without nodules along mid-dorsalline ........................ 3

Two protuberances along dorsal midline of carapace. First abdominal somite

produced posteriorly into a broad plate, which covers dorsal surface of second

somite. Apex of telson truncate, the entire border along each side armed with

nine or ten spines. Apical spines are not markedly longer than remaining

SOINeSIOn telsOMa. 3 foe. ho. ne oe ee te ee. M. camelina O. Tattersall, 1955

A single protuberance on dorsal surface of carapace. First abdominal somite

not produced along mid-dorsal line. Apex of telson rounded, the apical spines

markedly longer than the evenly spaced lateral spines, which number 19 or 20

WMS ACINESIGC Meee ee a ene geen hc. sex nl tot M. eremita O. Tattersall, 1962

WaAtcKalimianrcins On Ltelsonwithout SpIneS © 4.95265. 48-.... 5045 05e soe eens. 4

Lateral margins of telson at least partially armed withspines.............. 2)

Telson with lateral margins straight, converging to a narrow apex that is

armed with two long spines .............. M. bispinosa O. Tattersall, 1969

Telson with lateral margins straight, converging towards a blunt, almost

truncate apex that is armed with three pairs of spines. Outermost pair small,

almost lateral. Inner pair longer than those adjacent tothem ...............

Pe arn ita ls, Fe wel ee a ees M. suedafrikana O. Tattersall, 1969

Spines along lateral margins of telson continuous, without a gap in the

AUTTMAVONETTE <2 5 a occaye ce opkeeh Enka Cee Ton aE ete eda 6

Spines along lateral margins of telson discontinuous. Telson distinctly

pointed, the two apical spines about three times the length of the spines

adjacent to them. Lateral margins of telson with four spines on each side near

the base, proximal to an unarmed section that is equal to one-quarter the

length of the telson. Endopod of uropod with a graduated row of spines along

inner margin, extending almost to midlength of endopod. Rostrum long and

acutely pointed, reaching base of the third segment of antennular peduncle .. .

Ne Ao eo is ot Sy canes cee: Ne Ls M. buffaloensis sp. nov.

PMC SOc |ISOMGOUNGEM! 50s Fak Se cle eae Sav heed eee oe Heres 7

Apex of telson distinctly pointed, the two apical spines markedly longer than

those immediately adjacent to them. Spines along inner margin of endopod of

uropod graduated, extending only a short distance beyond statocyst .........

3 5 8 oo ob ip SiG a6. 66: Ry One an eo M. similis (Zimmer, 1912)

Apex of telson broadly rounded, marginal spines around apex increasing

evenly in length towards the median line. Armature along inner margins of

endopod of uropod graduated, the more distal spines not markedly longer

than the others. These spines closely set and extending about three-quarters

thedencilnomhe~endopod (2.0 .0...256.....+.. M. major (Zimmer, 1912)

Telson tapering posteriorly, apex smoothly rounded with marginal spines

increasing evenly in length towards the median line. Spines along inner margin

of endopod or uropod distinctly graduated, those more distal long and

slender. Spines extend along entire length ofendopod .....................

M. schultzei (Zimmer, 1912)

owe) (8) (6) (elms! is) te) in) 16, (6) (e| 'e; (6) (se) (ope) (6; :e) (6) [e) fee: ‘e|) (©: a) (eb (0-9) (a) je) oi je) @) Je fe) \e; ‘e; (o; Ie: (e) ie

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnostic features described for Mysidopsis buffaloensis also allow for clear

separation from the remaining 36 described forms. The most diacritical is the

discontinuity in the armament along the lateral margins of the telson. Only

M. hellvillensis Nouvel, 1964, M. taironana Brattegard, 1973, and M. tortonesei

Bacescu, 1968, have a comparable arrangement of spines on the telson. In

M. tortonesei the apex of the antennal scale is sharply pointed, the spines arming

the inner margin of the uropod extending along the entire length of the segment.

In the proximal half these spines are thick and robust. The telson is apically

rounded and the marginal spines increase evenly in length towards the apex.

Apical spines are therefore not distinctly longer than those immediately adjacent

to them (Baéescu 1968; Brattegard 1969, 1973).

In M. taironana the antennal scale lacks the distal segment and the inner

margin of the uropod is armed with only 2—4 spines, just distal to the statocyst.

The telson is triangular, apically rounded and about 1,2 times as long as the basal

width. Unarmed sections along the lateral margins of the telson are not always

distinct and there are two pairs of long, slender apical spines (Brattegard 1973).

Mysidopsis hellvillensis was described by Nouvel (1964) from Madagascar

and is clearly distinguishable from M. buffaloensis. In the former species the

spines along the inner margin of the endopod of the uropod extend along the

entire length of the segment. The rostrum is obtund and there are only two spines

on each side of the telson near the base. Posterior to these spines, the unarmed

portion is followed by a single spine and a further discontinuity in the spinal

arrangement along each side. In M. hellvillensis the apex of the telson is obtund

with two pairs of terminal spines subequal in length.

An indistinct discontinuation of marginal spines on the telson is sometimes

apparent in M. coralicola (Bacescu, 1975). This species was described by Bacescu

from Tanzanian coastal waters (Bacescu 1975), but it is more closely allied to

M. similis from southern Africa than to M. buffaloensis.

ACKNOWLEDGEMENTS

Financial assistance from the Department of Environment Affairs is

acknowledged.

REFERENCES

Baéescu, M. 1968. Etude des quelques Leptomysini (Crustacea Mysidacea) des eaux du Brésil

et de Cuba; description d’un genre et de cing autres taxons nouveaux. Annale Museo civico

de Storia naturale ‘Giacome Doria’, Genova 77: 231-249.

Bacescu, M. 1975. Contributions to the knowledge of the mysids (Crustacea) from the

Tanzanian waters. University of Dar es Salaam Science Journal 1 (2): 39-61.

BRATIEGARD, T. 1969. Marine biological investigations in the Bahamas. 10. Mysidacea from

shallow water in the Bahamas and southern Florida. Part 1. Sarsia 39 (1): 17-106.

BRATTEGARD, T. 1973. Mysidacea from shallow water on the Caribbean coast of Colombia.

Sarsia 54 (1): 1-65.

Brown, A. C. & TatBot, M.S. 1972. The biology of the sandy beaches of the Cape Peninsula,

South Africa. Part 3. A study of Gastrosaccus psammodytes Tattersall (Crustacea:

Mysidacea). Transactions of the Royal Society of South Africa 40 (4): 309-333.

A NEW SPECIES OF MYSIDOPSIS FROM SOUTHERN AFRICA 103

NovuveL, H. 1964. Mysidacés récoltés par S. Frontier a Nosy-Bé. 1. Description de deux

Leptomysini. Bulletin de la Société d Histoire naturelle de Toulouse 99 (1/2): 107-123.

TATTERSALL, O. S. 1955. Mysidacea. Discovery Reports 28: 1-190.

TATTERSALL, O. S. 1958. Further notes on Mysidacea from South African waters. Transactions

of the Royal Society of South Africa 35 (4): 373-383.

TATTERSALL, O. S. 1962. Report on a collection of Mysidacea from South African offshore and

coastal waters (1957-59) and from Zanzibar (1961). Proceedings of the Zoological Society

of London 139 (2): 221-247.

TATTERSALL, O. S. 1969. A synopsis of the genus Mysidopsis (Mysidacea, Crustacea) with a

key for the identification of its known species and descriptions of two new species from

South African waters. Journal of Zoology, London 158 (1): 63-79.

WOOLDRIDGE, T. 1983. Ecology of beach and surf-zone mysid shrimps in the eastern Cape,

South Africa. In: McLACHLAN, A. & Erasmus, T. eds. Sandy beaches as ecosystems:

449-460. The Hague: Junk.

ZIMMER, C. 1912. Siidwestafrikanishe Schizopoden. Jenaische Denkschriften 17: 1-11.

ha Sa a

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

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

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Punctuation should be loose, omitting all not strictly necessary

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

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

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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.

T. H. WOOLDRIDGE

A NEW SPECIES OF MYSIDOPSIS (MYSIDACEA)

FROM COASTAL WATERS OF SOUTHERN

AFRICA AND A KEY TO THE KNOWN

SPECIES FROM THE SUBCONTINENT

VOLUME 98 PART 5 JULY 1988 ISSN 0303-2515

7 AY ai

i a

ANNALS

“OF THE SOUTH AFRIC

CAPE TOWN

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Examples (note capitalization and punctuation)

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

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

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

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

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

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

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

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

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

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

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

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band CNITHSC

July 1988 Julie :

Part- 5» DeeK. ~™) 66-00

A REDESCRIPTION OF AFROCHILTONIA

CAPENSIS (K. H. BARNARD, 1916) WITH A

REVIEW OF THE GENERA OF THE FAMILY

CEINIDAE (CRUSTACEA, AMPHIPODA)

WOLFGANG ZEIDLER

Cape Town Kaapstad

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Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A REDESCRIPTION OF AFROCHILTONIA CAPENSIS

(K. H. BARNARD, 1916)

WITH A REVIEW OF THE GENERA OF THE FAMILY CEINIDAE

(CRUSTACEA, AMPHIPODA)

WOLFGANG ZEIDLER

South Australian Museum, North Terrace, Adelaide, Australia

(With 4 figures)

[MS accepted 2 February 1988]

ABSTRACT

Afrochiltonia capensis (Barnard, 1916) is redescribed and figured, including the first

description of the male, and a lectotype is selected. In the light of this redescription it is

concluded that A. capensis is sufficiently different from currently recognized Australian

congeners to warrant generic recognition. The genera of Ceinidae are reviewed briefly and

Austrochiltonia Hurley, 1959, is resurrected for the Australian species with Afrochiltonia K. H.

Barnard, 1955, being restricted to A. capensis from South Africa. A key to the genera of

Ceinidae is provided.

CONTENTS

PAGE

MINGRO GUC H OI rarest ee A as oi Ok ces ee A ea eae ve 105

Miatental Sram deme th Ods yar ot er ih ab ieelages he eae on cucg Be oieteees 106

SWSUCIIUAUICS megane mite etreane i oon A ae ats oR RK Nee 107

Reviewrolmche tantly; Ceimidacenne sae see a een oer 114

Key tothe ceneraorthe family @emidaeny..).-e.+- 08-4 oe 118

PNCknOWIEMCeMeMUS) himts ape ri ens Apne cue hua Oe ei mie atte. e 118

INGER CIN CE Sie es yeti an nee tek a, ene ear ea A GPA 118

PND DRE VIAN OMS My meni Ae el Se ee Ne mie oe sei Sak 119

INTRODUCTION

Afrochiltonia capensis (Barnard, 1916) has never been adequately figured.

The male was not correctly recognized until Griffiths (1976b) and has never been

described, although Rtihe (1914), referring to six young females (as Chiltonia

subtenuis), illustrated the second gnathopod of what is clearly a male.

Earlier confusion of non-ovigerous females with males led to a misleading

generic diagnosis, which has never been resolved satisfactorily. Barnard (1916)

suggested widening the diagnosis of Chiltonia to accommodate what he thought

was a species with gnathopod one and two alike in both sexes. Later Hurley

(1954) noted an unusual modification of the first pleopod in the male of New

Zealand species of Chiltonia, where the inner ramus forms 1-3 dorsally directed

whip-like lashes. This feature was thought to be of generic significance and led to

105

Ann. S. Afr. Mus. 98 (5), 1988: 105-119, 4 figs.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

the erection of Afrochiltonia Barnard, 1955, for the South African species and

Austrochiltonia Hurley, 1959, for the Australian species, with the New Zealand

species remaining in Chiltonia Stebbing, 1899. Thus Afrochiltonia was dis-

tinguished from Austrochiltonia by the condition of the male second gnathopod.

There the matter remained until Griffiths (1976b) discovered some males with the

characteristically enlarged second gnathopod. He consequently synonymized

Afrochiltonia with Austrochiltonia and furthermore Afrochiltonia capensis with

Austrochiltonia subtenuis. However, Afrochiltonia has priority over Austrochil-

tonia, as noted and corrected by Barnard & Karaman (1982). Despite all of these

systematic changes the South African species remained poorly known and the

male was still not described or figured.

Barnard’s (1916) description is considered inadequate by today’s standards

and is insufficient for comparisons with similar fauna in New Zealand and

Australia. Because ‘Chiltonias’ are very common and widespread in the

freshwaters of southern Australia, with several undescribed species, it is

important to establish clearly the systematic status of the South African species.

It is the purpose of this paper to redescribe Afrochiltonia capensis (Barnard,

1916), including the first description of the male, and to clarify the status of the

genera of Ceinidae, particularly Afrochiltonia and Austrochiltonia.

MATERIALS AND METHODS

Barnard’s (1916) type specimens of Afrochiltonia capensis were borrowed

from the South African Museum and examined in detail. As no other material of

A. capensis was available from the South African Museum, additional specimens

were collected at my request, from Milnerton Lagoon (see ‘Material examined’)

by Dr C. L. Griffiths. These specimens are deposited in the South Australian

Museum, except for the male described herein, which has been transferred to the

South African Museum. Specimens of Austrochiltonia in the collections of the

South Australian Museum were also examined for comparison.

Specimen length was measured along a lateral parabolic line drawn from the

anterior extremity of the head through the middle of the body to the posterior

limit of the telson. Barnard’s type material was not used for size comparisons

between males and females as his sample may have been biased towards larger

specimens.

The thoracic limbs are referred to as gnathopod 1 and 2 followed by

pereopods 3-7 to avoid confusion. Size comparisons of gnathopods exclude the

coxa and dactyl and of the pereopods the coxa, with articles being measured down

the middle.

Unless indicated otherwise dissected appendages were taken from the left

hand side of the animal. The mouthparts of the lectotype and the mouthparts and

appendages from the male, described herein, are mounted in poly-vinyl

lactophenol on microscope slides. All other appendages, remains of dissected

specimens, and other specimens are preserved in 75 per cent alcohol.

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 107

SYSTEMATICS

Family Ceinidae J. L. Barnard, 1972

Afrochiltonia capensis (Barnard, 1916)

Figs 1-4

Chiltonia subtenuis Rithe, 1914: 35, figs 13, 14a—c [non Sayce, 1902].

Chiltonia capensis Barnard, 1916: 224, pl. 27 (figs 38-40).

Afrochiltonia capensis Barnard, 1955: 93. Griffiths, 1974a: 253; 1974b: 327; 1975: 168; 1976a: 75,

fig. 47.

Austrochiltonia subtenuis Griffiths, 1976b: 30.

Type locality

Salt River, Cape Town, South Africa, by present designation of lectotype.

Material examined

Type material. K. H. Barnard’s syntypes consist of two lots.

SAM-—A28835 labelled ‘Type Specms’ from Salt River, Cape Town, collected

by Dr W. F. Purcell, October 1898, consisting of 8 females in alcohol, one of

which has been selected as lectotype; the remainder have been designated

paralectotypes and have been transferred to SAM—A39685.

SAM-—A2886 from Milnerton near Cape Town, collected by K. H. Barnard,

25 October 1913, consisting of 20 females (some damaged) in alcohol and a

microscope slide of the appendages of at least three specimens. All of this

material has been designated paralectotypes. The slide material, although

labelled ‘Type’, could not be used as the lectotype as it did not consist of a single

specimen and the mountant had become crazed with age, thus obscuring the finer

detail of the mounted appendages.

Other material. South Australian Museum No. C4165: 30 females and

6 males from Milnerton Lagoon, Cape Town, collected by C. L. Griffiths,

20 January 1987. SAM—A39686: male (the specimen described and illustrated

herein), collection data as for C4165 but transferred to the South African

Museum. South Australian Museum No. C4166: 40 females and 69 males from

mouth of Milnerton Lagoon, near Cape Town, collected by C. L. Griffiths, June

1987.

Description

Female

Lectotype, 3,7 mm, non-ovigerous, SAM—A2885. Coxal gills present

from G2 to P6. Oostegites, dorsally folded, present from G2 to P5.

Head as long as deep, length equivalent to first 1,5 pereonites; eyes black (in

alcohol), ovato-circulate with some ommatidia diffuse dorsally.

Antenna 1 short, about twice head length or equivalent to 0,2 times body

length; article 1 of peduncle almost twice as long as wide and 1,5 times length of

article 2; article 3 slightly shorter than 2; flagellum slightly longer than peduncle,

of six articles with a ventral aesthetasc at the base of each of the last two articles.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Afrochiltonia capensis (Barnard). A. Lectotype female, 3,7 mm (SAM-—A2885) (gills,

oostegites and pleopods not shown). Scale bar = 0,5 mm. B. Oostegites from paralectotype

specimen (SAM-—A39685), shown in order from G2—P5. Scale bar = 0,2 mm. Other appen-

dages from lectotype. Scale bar = 0,2 mm.

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 109

Fig. 2. Afrochiltonia capensis (Barnard). Lectotype female, 3,7 mm, gnathopods and

mouthparts. Scale bars = 0,2 mm and 0,1 mm, respectively.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Antenna 2 little more than 0,75 length Al, gland cone pressed firmly against

head—not visible laterally; article 1 of peduncle twice as wide as long; article 2 as

wide as long, twice length article 1 and slightly less than half length of article 3;

flagellum only slightly longer than peduncle, of six articles.

Upper lip as wide as long, apically rounded, bearing numerous short setae

distally.

Lower lip without inner lobes; outer lobes subovate with setose distal and

inner margins.

Mandibles without palp: left with incisor of seven teeth, lacina mobilis of five

teeth, spine row of three feathered spines and triturative molar with one long

feathered seta; right with incisor of five teeth, lacina mobilis of three teeth, spine

row of three feathered spines and molar like left.

Maxilla 1: outer plate without palp but notched at palp’s normal position,

with eight comb-like spines apically; inner plate very narrow with only two

feathered spines apically.

Maxilla 2: outer plate slightly longer than inner, about 0,75 times as wide;

both apically setose; inner plate with one large seta on inner margin at end of setal

TOW.

Maxilliped: inner plate reaching extremity of article 1 of palp, rectangular,

about three times as long as broad, with three stout spines apically, the inner one

very small; outer plate reaching 0,75 along article 2 of palp, ovate, about as wide

as inner plate bearing several setae apically and along inner margin; palp article 1

with oblique distal margin, length outer margin about 2,5 times inner; palp

article 2 slightly broader than long and slightly shorter than outer margin of

article 1, bearing a few setae on inner distal corner and distal half of inner margin;

palp article 3 about as long as broad and as long as article 2, with sparse long

setae on distal and inner margins; palp article 4 small, conical, slightly longer than

wide, about half length of article 3; dactylus sharp, as long as article 4.

Gnathopod 1: coxa length about 1,5 times width, longer than article 2 with

antero-dorsal corner slightly produced; article 5 length twice maximum width,

postero-distal lobe not produced, with row of six stout pectinate spines; article 6

as long as article 5, about twice as long as wide, postero-distal corner rounded,

distal face with spine on either side of dactyl and two long medial setae, antero-

distal corner with four long setae, posterior margin with small spine near

postero-distal corner; dactyl as long as width of palm and fitting neatly against

palm.

Gnathopod 2: similar to G1 but 1,25 times as long; coxal gill sac-like, more

than twice as long as wide, as long as article 2; coxa slightly longer than wide, as

long as article 2; article 5 with slightly produced postero-distal lobe with row of

five stout pectinate spines; article 6 without small spine on posterior margin,

otherwise as in Gl.

Pereopod 3: length 1,35 times G2; coxal gill like that of G2; coxa like that of

G2 but slightly larger, slightly longer than article 2; article 4 broad, about

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 111

0,7 times length; article 5 like 4 but not as broad and only 0,75 times as long;

article 6 length 1,6 times article 5, about three times as long as wide; dactyl

length 0,4 times article 6; all articles sparsely setose as illustrated.

Pereopod 4 identical to P3 except for coxa. Coxa with shallow antero-dorsal

excavation, maximum width 1,5 times length, slightly longer than article 2,

posterior margin oblique so that width at distal margin is only about half

maximum width.

Pereopod 5: smallest pereopod, length about 0,9 times P4; coxal gill similar

to P4 but a little wider; coxa width almost twice width article 2, length of anterior

lobe about half maximum width coxa, length of posterior lobe about 0,7

maximum width coxa or as long as article 2; article 2 slightly longer than wide

with typical expanded posterior margin and postero-distal lobe overlapping and

almost reaching to distal margin of article 3; article 4 length 1,2 times width, with

postero-distal corner produced; article 5 length about 1,2 times article 4 and of

similar shape except postero-distal corner is not as produced; article 6 length

1,6 times article 5, about three times as long as wide; dactyl length 0,4 times

article 6; all articles sparsely spinose as illustrated.

Pereopod 6 longest pereopod, length 1,25 times P5; like PS except articles

3-6 somewhat longer in proportion to their width; coxa as wide as article 2,

anterior lobe small, length about half width coxa, posterior lobe as long as coxa

width.

Pereopod 7 a little shorter than P6; coxa semi-circular, slightly wider than

long; article 2 as wide as long, postero-distal lobe extending beyond article 3,

posterior margin slightly serrate and minutely spined with acute proximal corner;

otherwise similar to P6.

Pleonal epimera with very small postero-ventral tooth.

Uropod 1 longer than U2; rami subequal, about 0,8 times as long as

peduncle, outer ramus with two large and two small spines at tip, inner ramus

with two large and three small spines at tip; peduncle with large spine on inner

and outer-distal corner and two more on dorsal outer margin.

Uropod 2: rami subequal, a little more than 0,8 times as long as peduncle;

peduncle and rami with spines as in U1 except outer ramus also has a spine on

middle of inner margin (only on right in lectotype).

Uropod 3 one-articulate, half length of telson, conical in shape with one long

outer and one short inner seta at tip.

Telson entire, hemispherical, slightly wider than long.

Oostegites from paralectotype (SAM-—A39685), ovigerous, most eggs

released. All with curled margins and numerous small hooks forming a tight

marsupium. First is pentagonal with long distal and posterior margins, almost as

wide as deep; second is rectangular, length 1,8 times width; third is also

rectangular, length a little more than 1,8 times width and is longest oostegite,

about 0,6 times length of P4; fourth is sub-rectangular with oblique distal margin

and excavate postero-distal corner, maximum length 1,3 times maximum width.

LZ, ANNALS OF THE SOUTH AFRICAN MUSEUM

Male

Hypotype 2,5 mm (SAM-—A39686), generally like female but differs as

follows.

Coxal gills relatively smaller.

Antenna 1 flagellum a little shorter than peduncle, consisting of only five

articles.

Antenna 2 only slightly shorter than Al.

Mandibles: only right molar with long feathered seta.

Gnathopods and pereopods with articles not so stout.

Gnathopod 1: coxa narrower distally, about 0,7 times dorsal width, without

antero-dorsal corner produced; article 6 slightly longer than article 5 with two

spines on distal face in addition to those on either side of the dactyl.

Gnathopod 2 with enlarged article 6, unlike G1; length about 1,4 times G1;

coxa length 1,2 times width, only 0,9 times length article 2; article 4 with right-

angled bend; article 5 small, without pectinate spines; article 6 a little longer than

article 2, maximum length 1,6 times maximum width, postero-proximal corner

forming distinct lobe for almost 0,4 length article, palm oblique with several small

spines on either side of ‘cutting edge’ followed proximally by small groove for tip

of dactyl; dactyl claw-like, length 0,8 times maximum length article 6.

Pereopod 3: length 1,1 times G2; coxa like that of G2 only slightly longer.

Pereopod 4: coxa maximum width a little more than length, only slightly

more narrow distally—posterior corner of excavation not produced as in female.

Pereopod 5 as long as P4; coxa width 1,5 times width article 2, length

antericr lobe slightly less than half maximum width coxa; length posterior lobe

less than 0,6 times maximum width coxa or only 0,7 times length article 2.

Pereopod 6: coxa, length anterior lobe 0,3 times width coxa, length posterior

lobe a little less than coxa width.

P7, U1—U3 and telson like female.

Pleopods all of normal structure (not modified as in Chiltonia).

Variations

Females ranged in size from 1,0 to 4,1 mm with a mean of 3,2 + 0,2 mm

(+ 95 % c.1.; n = 70). Males ranged in size from 1,5 to 2,6 mm with a mean of

2,2 + 0,06 mm (+ 95 % c.1.; n = 76). Males are thus significantly smaller than

females (P<<0,01). No specimens were as large as 4,5 mm as recorded by

Barnard (1916) and one must assume this measurement included the antennae or

to be an error.

All of the specimens examined varied little from the above descriptions. In

some specimens the ommatidia of the eyes were more diffuse at the edges and

almost confluent dorsally. Antenna 1 and 2 usually had a flagellum of six articles,

increasing to seven in some females, or decreasing to five in some males but rarely

in females. The antennae were almost equal in length in some specimens and

varied from 0,2 to 0,25 times the body length. The condition of the mandible

with a feathered seta on the molar usually occurred only on the right, rarely on

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 113

Fig. 3. Afrochiltonia capensis (Barnard). Lectotype female, 3,7 mm, pereopods 1-7.

Scale bar = 0,2 mm.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

the left as in the lectotype. The rami of Ul were sometimes as short as 0,6 times

peduncle, particularly in males. The inner ramus of U2 without a medial spine

was rare. Oostegites of females varied considerably in size but were expanded and

as illustrated in ovigerous specimens.

Remarks

Griffiths (19766) synonymized Afrochiltonia capensis with Austrochiltonia

subtenuis on the basis that the male G2 was enlarged and U3 was one-articulate.

However, apart from a number of minor differences, Afrochiltonia capensis is

clearly distinguished from Austrochiltonia subtenuis by: (1) the shape of coxa 4;

(2) the relative lengths of the antennae; (3) the stout nature of the pereopods,

particularly in the female; (4) the lack of marginal spines on the rami of U1

and U2; (5) the gland cone on A2 is not visible laterally; and (6) P7 is shorter

than P6. The synonymy proposed by Griffiths (1976b) is thus considered invalid.

The earlier confusion on non-ovigerous females with males might be

explained by the fact that the males are considerably smaller than the females and

may have been overlooked when sorting samples or confused with juveniles of

other common species (e.g. Melita zeylanica). There is also a likely seasonal

variation in the number of males present in the population, as is evidenced by the

two random samples collected in January and June 1987, which contained 6/36

and 69/109 males, respectively. It is therefore possible that, when Barnard’s

samples were collected in October, few males were present in the natural

population and were thus not represented in his samples. Further evidence of a

seasonal breeding cycle is provided by the fact that in January 15/30 females were

Ovigerous as compared to only 9/40 in June. However, a more detailed study of

the life cycle of this species is required to determine breeding seasons and

seasonal variations in the male/female ratio.

Now that the male has been described it is possible to re-evaluate the

systematic position of this species with that of similar taxa in New Zealand and

Australia. In particular, I recommend retaining the genus Austrochiltonia Hurley,

1959, for the Australian species and reserving Afrochiltonia Barnard, 1955, for

the single South African species.

Distribution

Kosi Bay, Zululand, to Olifants River, western Cape, in brackish—estuarine

environments (endemic).

REVIEW OF THE FAMILY CEINIDAE J. L. BARNARD, 1972

The family Ceinidae is divided into two subfamilies, the Ceininae—consist-

ing of marine forms with cleft telsons—and the Chiltoniinae—consisting of

freshwater—brackish forms with uncleft telsons (Barnard 1972b). It is generally

distinguished from the Hyalidae (= Talitridae) by the form of U3 (e.g. Barnard

1972b), which is diagnosed by Barnard (1972a) as ‘composed only of peduncle

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 1S)

Fig. 4. Afrochiltonia capensis (Barnard). Male, 2,5 mm (SAM-—A39686), pereopods 1-7.

Scale bar = 0,2 mm.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

.... This diagnosis needs to be amended as the Australian species Austrochil-

tonia australis (Sayce, 1901) has a third uropod consisting of two articles, a feature

used to distinguish it from the only other described Australian species

A. subtenuis (Sayce, 1902) (Williams 1962). In addition, I have recently

discovered some undescribed Australian species that also have a third uropod

consisting of a peduncle and a small ramus. The family diagnosis therefore needs

amending as follows: ‘third uropod usually composed of peduncle only, some

freshwater species (Australian) with additional small article’.

The problem now remains to determine a more satisfactory way to

distinguish this family from the Hyalidae.

The subfamily Ceininae J. L. Barnard, 1972, consists of entirely marine

species placed in the three genera, Ceina Della Valle, 1893, Taihape J. L.

Barnard, 1972, and Waitomo J. L. Barnard, 1972. These genera have been

adequately diagnosed by Barnard (1972b) and they will not be discussed further

here.

The subfamily Chiltoniinae J. L. Barnard, 1972, consists of brackish—

freshwater species currently placed in only two genera, Chiltonia Stebbing, 1899,

and Afrochiltonia K. H. Barnard, 1955.

Chiltonia, endemic to New Zealand, is readily distinguished from the other

Chiltoniinae by the unusual character of the first male pleopod, in which the inner

ramus is modified to form 1-3 dorsally directed whip-like lashes. The three

species concerned are adequately dealt with by Hurley (1954), who also provided

a Satisfactory generic diagnosis.

Afrochiltonia was erected by Barnard (1955) for the South African species

A. capensis, based on the mistaken belief that the gnathopods were alike in both

sexes, a feature used to distinguish it from Austrochiltonia, erected by Hurley

(1959) for the Australian species. When males of Afrochiltonia capensis were

eventually recognized correctly and found to have an enlarged G2 (Griffiths

1976b), the distinction between the two genera was considered invalid and

Griffiths (1976b) promptly synonymized Afrochiltonia with Austrochiltonia.

However, Afrochiltonia has priority over Austrochiltonia as was noted and

corrected by Barnard & Karaman (1982). This synonymy came about without an

examination of the Australian species and in the absence of an adequate

description Afrochiltonia capensis. Having examined both the South African and

Australian species I find this synonymy unsatisfactory, as A. capensis is

considered sufficiently different from Australian species (including known

undescribed ones) to warrant the resurrection of Austrochiltonia. Afrochil-

tonia is thus restricted to A. capensis from South Africa and is diagnosed as

follows. :

Afrochiltonia K. H. Barnard, 1955

Males only about two-thirds size of females. Eyes ovato-circulate, occa-

sionally with diffuse ommatidia at edges. Antenna 1 and 2 subequal in length,

only about 0,2 times body length. Antenna 1 with ventral aesthetasc at base of

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 117

last two flagella articles. Gland cone of A2 not visible laterally. Maxilla 1 without

palp, notched at palp’s normal position. Coxae 1—4 deep. Coxa 4 with large,

shallow posterior excavation, maximum width more than length (up to 1,5 times

in female). Gnathopods 1 and 2 subchelate in both sexes; article 6 of G2 enlarged

in male. Pereopods stout, particularly in female; P7 shorter than P6. Pleopod 1

not modified in male. Uropod 3 one-articulate. Telson entire, hemispherical,

with evenly rounded distal margin.

Austrochiltonia is here resurrected for the Australian species A. subtenuis

(Sayce, 1901), A. australis (Sayce, 1902) and several undescribed species, and is

diagnosed as follows.

Austrochiltonia Hurley, 1959

Males of similar size or only marginally smaller than females. Eyes ovato-

circulate without diffuse edges. Antenna 1 longer than antenna 2, more than

0,3 times body length. Antenna 1 with ventral aesthetasc at base of distal four or

more flagella articles. Gland cone of A2 large, visible laterally. Maxilla 1 without

palp, notched at palp’s normal position. Coxae 1—4 deep. Coxa 4 with shallow or

deep posterior excavation, maximum width equal to, or less than, length.

Gnathopods 1 and 2 subchelate in both sexes; article 6 of G2 enlarged in male.

Pereopods slender, P7 longer than P6. Pleopod 1 not modified in male. Uropod 3

with single ramus or ramus absent. Telson entire, subrectangular to slightly

concave.

Although Afrochiltonia superficially resembles Austrochiltonia there are

several differences that collectively are considered of generic significance. In

particular the large coxa 4 of Afrochiltonia capensis is most unusual and is unlike

any species of Austrochiltonia or Chiltonia. The small size of the male relative to

the female is also unusual. In his revision of Austrochiltonia, Williams (1962)

noted that the largest specimens were males but, in a more detailed study of

A. australis, Smith & Williams (1983) found that the largest male was about

0,8 times as long as the largest female. In the present study of Afrochiltonia

capensis, in which 76 males and 70 females were measured, the largest male was

2,6 mm and the largest female was 4,1 mm, a ratio of only slightly more than 0,6!

Other Gistinguishing characters of Afrochiltonia, apart from those given in

the diagnosis and other minor differences, are the lack of marginal spines on the

rami of uropod 1 and uropod 2 (except for one on the inner ramus) and the

relatively few pectinate spines on article 5 of gnathopod 1 and gnathopod 2

(female).

Also of possible significance is that Afrochiltonia is only found in estuarine

habitats, while Austrochiltonia— although found in brackish—freshwater habitats

throughtout the southern half of Australia—has not been recorded from an

estuarine environment.

In view of the above, the genera of Ceinidae may now be distinguished

according to the following key.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

KEY TO THE GENERA OF THE FAMILY CEINIDAE

1. Telson’shightly cleft, marine (Ceiminae)) 9 55.4524 50s) oa eee 2

— Telson entire, freshwater—brackish (Chiltoniinae) ...................... 4

2. Pereopod 3 with numerous long setae on anterior margins of articles 2-4 ....

rae eer ria NC WINN. UR cana Bion 5,8 6 0 '0,0 0 0 2 Taihape

= Pereopod 3 lacking longisetaeonvanticles)2=49— 5 5 a ee 3

3. Head attached to body in underslung fashion, mandibular molar forming long

smooth thomy v4 sea a Gaeta s wine ae ee nee ee A eee Ceina

— Head attached normally to body, mandibular molar large and heavily

{ritwralive: on. oe 5 Ae eek ee Ot ae Waitomo

4. Pleopod 1 of male with inner ramus modified to form 1-3 dorsally directed

whip-like lashes (NewZealand) eee eee ee Chiltonia

— Pleopod lot male of normalstructure 4-2-2545. 4542 eee 5

5. Coxa 4 maximum width 1,5 times length in female, only marginally wider

than long in male; estuarine (South Africa) ................. Afrochiltonia

— Coxa 4 maximum width equal to or marginally less than length in both sexes;

brackish—freshwater, non-estuarine (Australia) ........... Austrochiltonia

ACKNOWLEDGEMENTS

I thank Mrs M. G. van der Merwe, South African Museum, for the loan of

Barnard’s syntypes and for checking Museum records for more material. I am

particularly grateful to Dr C. L. Griffiths, Zoology Department, University of

Cape Town, for collecting and sending me more specimens on two separate

occasions, thus making this study possible. Mrs K. L. Gowlett-Holmes is thanked

for typing the manuscript.

REFERENCES

BARNARD, J. L. 1972a. Gammaridean Amphipoda of Australia. Part I. Smithsonian

Contributions to Zoology 103: 1-333.

BARNARD, J. L. 19726. The marine fauna of New Zealand: Algae-living littoral Gammaridea

(Crustacea Amphipoda). Memoirs. New Zealand Oceanographic Institute 62: 1-216.

BARNARD, J. L. & KARAMAN, G. S. 1982. Classificatory revisions in Gammaridean Amphipoda

(Crustacea). Part 2. Proceedings of the Biological Society of Washington 95 (1): 167-187.

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. 1955. Additions to the fauna-list of South African Crustacea and

Pycnogonida. Annals of the South African Museum 43 (1): 1-107.

GrirFitus, C. L. 1974a. The Amphipoda of southern Africa. Part 3. The Gammaridea and

Caprellidea of Natal. Annals of the South African Museum 62 (7): 209-264.

GrirFitHs, C. L. 1974b. The Amphipoda of southern Africa. Part 4. The Gammaridea and

Caprellidea of the Cape Province east of Cape Agulhas. Annals of the South African

Museum 65 (9): 251-336.

GrirFitHs, C. L. 1975. The Amphipoda of southern Africa. Part 5. The Gammaridea and

Caprellidea of the Cape Province west of Cape Agulhas. Annals of the South African

Museum 67 (5): 91-181.

GrirFfitHs, C. L. 1976a. Guide to the benthic marine amphipods of southern Africa. Cape

Town: South African Museum.

REDESCRIPTION OF AFROCHILTONIA CAPENSIS (BARNARD) 119

GriFFITHS, C. L. 1976b. Some new and notable Amphipoda from southern Africa. Annals of

the South African Museum 72 (2): 11-35.

Hurey, D. E. 1954. Studies on the New Zealand amphipodan fauna. No. 2. The family

Talitridae: the freshwater genus Chiltonia Stebbing. Transactions of the Royal Society of

New Zealand 81 (4): 563-577.

Hurtey, D. E. 1959. Austrochiltonia, a new generic name for some Australian freshwater

amphipods. Annals and Magazine of Natural History (13) 1 (12): 765-768.

RuvuweE, F. E. 1914. Die Stisswassercrustaceen der Deutschen Stdpolar-Expedition 1901-1903

mit ausschluss der Ostracoden. Deutsche Stidpolar-Expedition 6 (8): 5—66.

SaycE, O. A. 1902. Description of some new Victorian freshwater Amphipoda, No. 2.

Proceedings of the Royal Society of Victoria 15 (1): 47-58.

SmiTH, M. J. & Wituiams, W. D. 1983. Reproduction cycles in some freshwater amphipods in

southern Australia. Memoirs of the Australian Museum 18: 183-194.

WituiaMs, W. D. 1962. The Australian freshwater amphipods. I. The genus Austrochiltonia

(Crustacea: Amphipoda: Hyalellidae). Australian Journal of Marine and Freshwater

Research 13 (2): 198-216.

ABBREVIATIONS

Al antenna 1 Gl gnathopod 1

A2 antenna 2 G2 gnathopod 2

Mxp maxilliped P3—P7 pereopod 3 to pereopod 7

Mx1 maxilla 1 U1-U3 uropod 1 to uropod 3

Mx2 maxilla 2 aly telson

Md mandible Epl-Ep3 first to third pleonal epimeron

UL upper lip R right

LL lower lip L left

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

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

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Punctuation should be loose, omitting all not strictly necessary

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

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

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

WOLFGANG ZEIDLER

A REDESCRIPTION OF AFROCHILTONIA CAPENSIS

(K. H. BARNARD, 1916) WITH A REVIEW

OF THE GENERA OF THE FAMILY CEINIDAE

(CRUSTACEA, AMPHIPODA)

VOLUME 98 PART 6 = DECEMBER 1988 : ISSN 0303-2515

‘OF THE SOUTH AF RIGAN-

MUSEUM

CAPE TOWN

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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

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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-

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 98 Band

December 1988 Desember

Part 6 Deel

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES

PART 17. PYCNOGONIDA

FRANCOISE ARNAUD

C. ALLAN CHILD

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

PART 17. PYCNOGONIDA

FRANCOISE ARNAUD

Station marine d’Endoume et Centre d’Oceanologie, F-13007 Marseille, France

C. ALLAN CHILD

Smithsonian Institution, Washington, D.C. 20560, USA

(With 14 figures)

[MS accepted 27 May 1987|

ABSTRACT

Thirty-nine species of Pycnogonida collected off the east coast of South Africa are

recorded. One new genus, Safropallene (family Callipallenidae) and the following 13 new

species are described: Pantopipetta armata, Pantopipetta bilobata, Callipallene africana, Para-

pallene invertichelata, Safropallene longimana, Hedgpethia magnirostris, Nymphon barnardi,

Nymphon bicornum, Nymphon granulatum, Nymphon obesum, Nymphon_paralobatum,

Nymphon pedunculatum, and Nymphon serratidentatum.

A brief discussion of the distribution and zoogeography of the pycnogonids collected by the

R.V. Meiring Naude indicates the presence of a considerable endemic pycnogonid fauna on the

continental shelf and slope of the east coast of South Africa.

CONTENTS

PAGE

MACKO AUCH OMe yaa. eee A as cee ate lnns biel yearn Ae il eta se 121

SIE CICSMIS Cremer retest: Mercure cite otra cee ceeeta as lato care eh ac Ww

SVSlematiciacCOUME oc nati en Gai Lae oles Sick aPeeaniconian lo Nia 124

RatmilyeAmmmopneidac sy cae Gan kook: a) oe, ees 124

RamilyBhoxichilididac msec. een ete seco oe ee 130

RamulysAustrodecidac’ cs acu. 2 ses aga oles ace: 31

Ramuly«@allipallemidac. 4 ce. cscs aes sea hehe: 7)

Ramulyshycnosomidacyean cs tye tern aan ok ae Gya cee 150

Famuily/@olossendeidac i. ee as sles hee ee hate Ye 1153)

FamulysNymphomidacne ac ono 40 cae: ieee wee 156

Distributiontand:zoopeorgraphyense aca ae oe 180

PXCKNOWIECECIIEMUS A: Bungee ry retentte. 0 Sate hh yale els iets ccc 184

| RUE LEVEN CES oe cheno Gesetes one dane, Sib etapa ee eee 185

INTRODUCTION

The paper deals with the pycnogonids taken during four cruises by the

South African Museum on R.V. Meiring Naude in 1976, 1977, 1978 and 1979.

No pycnogonids have come to light from the 1975 cruise. The specimens

Ann. S. Afr. Mus. 98 (6), 1988: 121-187, 14 figs.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

originate from continental shelf and slope localities from just south of the

Mozambique-—South African border to the East London area and from depths of

80 to 1 300 m. For full station data as well as background information on the

cruises the reader is referred to Louw (1977, 1980).

All specimens are deposited in the South African Museum, Cape Town,

under SAM-A catalogue numbers, except for a few surplus or duplicate lots of

material deposited in the U.S. National Museum of Natural History (USNM),

Washington, and the Museum National d’Histoire Naturelle (MNHN), Paris.

Material unidentified at species level is either damaged or immature.

SM Station

Family Ammotheidae

Ammothella setacea (Helfer)

Ascorhynchus ornatum (Helfer)

Boehmia tuberosa Mobius

Cilunculus sewelli Calman

Tanystylum thermophilum (Barnard)

Family Phoxichilidiidae

Anoplodactylus typhlops Sars

Family Austrodecidae

Pantopipetta armata sp. nov.

Pantopipetta bilobata sp. nov.

Family Callipallenidae

Callipallene phantoma (Dohrn)

Callipallene africana sp. nov.

Callipallene sp.

Pallenopsis capensis Barnard

Pallenopsis longirostris Wilson

Pallenoides sp.

Parapallene algoae Barnard

Parapallene invertichelata sp. nov.

Pseudopallene gilchristi Flynn

Safropallene longimana gen. et sp. nov.

SPECIES LIST

no.

6S wena 22 Qe

4 — 6

3 — il

(3 OO, 22, INS)

—_ a 1

3 — 1

CUO Gc 3 LS, uns.)

14 — 3

4 — it

— — 1

1 a =

1 = or

4 pul 1

— _ 1

1 _ 1

= — 4

— 1

— 5

3 oat we

aes as 1

— — 1

1 2s 1

1 — il

— — 1

_ — 2:

CLG, 22)

12 iS)

juv.

|»

2 |

fesexl Tt fos

[pce r=

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 123

SM Station

HO, Cne OG Ce WeRyiel QQ ii.

Family Pycnogonidae

Pycnogonum (Nulloviger) africanum

Calman 162 — 1 — — —

232 — — — 1 —

Pycnogonum crassirostrum Sars Veay — 1 — — —

Pycnogonum forte Flynn 185 — — — 1

226 — — — 1 1

Pycnogonum nodulosum Dohrn 86 — — — Zz —

250 — — — 3 —

Family Colossendeidae

Hedgpethia magnirostris sp. nov. 103 — 1 if — —

Hedgpethia sp. 226 — — — — 1

Rhopalorhynchus gracillimus Carpenter 164 — — — 1 —

Family Nymphonidae

Nymphon barnardi sp. nov. 250 — — — 2 —

Nymphon bicornum sp. nov. 60 — 1 — — —

129 — — 2 ae

Nymphon comes Flynn 179 — — —— 1 —

233 — 1 — — —

Nymphon crenatiunguis Barnard 179 — 1 oa —- —

250 — — — 1 —

Nymphon distensum Mobius 86 — 2 — 2 —

179 1 — — 1 —

185 1 — il 1 3

232 — 1 — —- —

Nymphon granulatum sp. nov. 129 — — — 1 —

162 — 3 1 —

226 — 1 — 3 —

232 — — — 1 3

236 — — — 1 il

Nymphon microctenatus Barnard M33) — — 1 — —

Nymphon modestum Stock 86 1 1 — -- —

239 — — 1 — —

250 — 3 — 1 7

Nymphon obesum sp. nov. 179 — — — 1 _

Nymphon paralobatum sp. nov. 226 — — — 1 —

Nymphon pedunculatum sp. nov. 163 2 1 — — 1

164 — — — 3 —

Nymphon phasmatodes B6hm 180 — — — 1 —

185 — — — 1 1

Nymphon pilosum Mobius 86 6 3) — 4 5

103 2 1 4 — 3

129 — 1 — - —

226 — — — 1 —

250 — 1 — 2 —

Nymphon serratidentatum sp. nov. 226 — 1 — 1 1

Nymphon sp. 185 — — — — 1

250 — — —— — 1

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATIC ACCOUNT

Family Ammotheidae Dohrn, 1881

Ammothella Verrill, 1900

Ammothella setacea (Helfer, 1938)

Rice

Kyphomia setacea Helfer, 1938: 179-181, fig. 9A—C. Barnard, 1954: 144, fig. 28A.

Ammothella setacea Stock, 1953a: 41; 1954: 118 (text), 120 (key).

Material

Transkei area. Neotype, SAM—A10221, SM 226, 32°28,6’S 28°58,8’E,

710-775 m, 1 ovigerous ¢. Paraneotype, SAM-—A10220, SM 226, 1 gravid 9°.

SAM-A10240, SM 226, 104 dd, 22. USNM 216758, SM 226, 2 dd, 2 98.

MNHN-Py 593, SM 226, 1 6, 1 2. SAM-A10241, SM 232, 620-560 m, 14 dd,

3 22, 1 juv. SAM-—A10222, SM 236, 670-660 m, 4 dd, 1 29. SAM-A10242,

SM 250, 150-200 m, 1 &.

Zululand area. SAM-—A10236, SM 86, 550m, 4646, 6292, 4 juv.

SAM-—A10219, SM 103, 680 m, 3 dd, 1 &.

Durban area. SAM-A10237, SM 129, 850 m, 93 dd, 22. SAM-A10238,

SM 151, 900 m, 1 @.

East London area. SAM-A10239, SM 163, 90m, 246d, 198, 1 juv.

F. Arnaud’s collection, 1 6, SM 163.

Redescription

Male. Trunk fairly slender, first anterior segmentation line present, second

usually absent, third lacking. Lateral processes long, separated by their diam-

eters or slightly less proximally, armed with three long dorsodistal slender

tubercles, except for the posterior pair which have two, and two laterodistal long

spines. The longest tubercle measures over twice diameter of lateral process.

Trunk armed with single anterolateral slender tubercle as long as chelifore basal

diameter. Ocular tubercle very long, slender, eyes at distal swollen tip, slightly

pigmented; tubercle armed with one, two, or three posterior pointing slender

tubercles of varying lengths, but some specimens without posterior tubercles.

Proboscis moderately inflated, without constrictions, lips almost flat. Abdomen

typical of genus, long, with ventral bend in posterior half, armed with set of

extremely variable dorsal tubercles consisting of from two to six single or double

slender tubercles equal to and in varying lengths as the lateral process tubercles,

and a pair of distal setae.

Chelifore slender, flaring distally, not as long as proboscis, the first segment

armed with slender distal tubercle, the longer second segment with variable

number (4—7) of similar dorsal tubercles and several long and short distal setae.

Chela vestigial, without fingers, sometimes everted, sometimes extended beyond

chelifore distal cup. Chela armed with single seta.

Palp typical of genus, distal five segments subequal, armed with many

ventral and lateral setae slightly longer than each segment diameter.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES IDS

Fig. 1. Ammothella setacea (Helfer), male. A. Trunk, dorsal view. B. Trunk, lateral view.

C. Chelifore tip. D. Trunk anterior, oblique lateral view. E. Third leg, with enlargement of

cement-gland tube. F. Distal segments of third leg. G. Oviger with egg attached. H. Strigilis

segments of oviger.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oviger also typical, with second segment longest. Distal four segments each

armed with one denticulate spine, except terminal segment with two. Spines

with five or six serrations per side. Egg larger than for others of genus, equal in

diameter to length of fourth segment.

Leg very slender, armed with long dorsal and lateral spines equal to or

longer than segment diameters and dorsal blunt spines equal to or shorter than

segments. Femoral cement-gland tube very long, dorsodistal, carried at low

angle. Tarsus extremely short, propodus long, slender, armed with few long

dorsal setae, three slender heel spines, and from seven to ten short sole spines.

Claw slender, less than half propodal length, auxiliaries very slender, almost as

long as main claw.

Proboscis, chelifores and palps originating in an anterior trunk concavity,

slightly similar to that found in the genus Cilunculus.

Female. Slightly larger in all measurements except for oviger, which is

reduced in size with a fourth segment of reduced length. Posterior trunk seg-

mentation lines present or absent. Tubercles on chelifore and on lateral trunk

shorter, as are those of lateral processes and abdomen.

Previous record

Agulhas Bank, 126 m.

Remarks

This species was described from a single female specimen and the above

records apparently constitute only the second time this species has been taken.

The type specimen was evidently lost during wartime (Stock 1953a: 41), so we

have designated a neotype and a paraneotype from this large collection. This

heretofore enigmatic species appears to have some characters of the closely

related genus Cilunculus. The accompanying figures reveal several characters

not illustrated by Helfer (1938). The proboscis, chelifore and palpi originate in a

cephalic segment ‘hood’ typical of Cilunculus species, but this species has the

longer chelifores of two segments, long tubular spines and general habitus of the

genus Ammothella. The differences cited in the literature for Cilunculus are this

cephalic hood, shorter chelifores of one or two segments, usually long pointed

spines on the appendages, and a femoral gland showing as a long conical out-

growth of the femur. The genus Ammothella usually lacks the cephalic hood, has

chelifores usually as long as the proboscis, long hollow clubbed or tubular

spines, and an articulated distal tube as the emergent part of the femoral gland.

These characters are mixed in A. setacea. Among the known species of both

genera there are exceptions to every character used to diagnose the two.

Cilunculus has usually been characterized by short chelifores but one species,

C. acanthus, has two-segmented chelifores almost as long as the proboscis. On

the other hand, several species of Ammothella (A. tubicen, A. thetidis, and

others) display an ocular segment anterior extension suggestive of a hood, and

two other genera, Boehmia and Oorhynchus, have projecting cephalic collars or

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES OF

hoods over the chelifore insertion. The number of scape segments is not diag-

nostic for Cilunculus, but all known Ammothella species have long two-

segmented scapes. Cilunculus has eight known species with two-segmented

scapes and seven with a single segment. The cement glands of Ammothella

species are characterized as tubes that are sometimes articulated, but at least

two species (again A. tubicen and A. thetidis) have what appear to be long

conical gland tubes, which have no articulation with the femur. The cement

glands of Cilunculus are not known to be articulated but appear similar to those

Ammothella species mentioned. This leaves the long tubular spines of Ammo-

thella as a character not known to be shared by Cilunculus. If all Ammothella

species had tubular blunt spines, the character might serve as diagnostic for the

genus, but less than half of the known species have these spines. The remainder

have long (sometimes only short) pointed spines, to all appearances the same as

those on species of Cilunculus. Therefore, spine type cannot be a valid diag-

nostic character of Ammothella. At least one other genus, Eurycyde, has species

with similar hollow blunt spines. |

Cilunculus is known for its deep-water species and indeed, most are found

in waters deeper than 100 m. There are exceptions to this aspect also. Ciluncu-

lus australiensis has a known depth range of 40-70 m and C. cactoides has been

taken in as little as 38 m (unpublished National Museum of Natural History

specimens). Cilunculus species have never been found in the littoral, whereas

Ammothella is usually a littoral genus, but A. profunda was taken in over

1000 m. This species was found to have a cephalic hood and was recently

moved to Cilunculus (Nakamura & Child 1983: 32).

We believe, in spite of the evidence presented here, that the two genera

should be kept separate for now. The differences remain, even with exceptions,

and those of the tubular blunt spines, shallow- versus deeper-water habitats, pre-

dominantly short chelifores, and hooded cephalic segment continue to be sufficient

evidence, in our opinion, to arrange the species in two separate genera.

Distribution

Extended considerably from Agulhas Bank to Transkei coast, in depths

from 90 to 900 m.

Genus Ascorhynchus Sars, 1877

Ascorhynchus ornatum (Helfer, 1938)

Ainigma ornatum Helfer, 1938: 181, fig. 10a—f. Barnard, 1954: 144, fig. 28B.

Ascorhynchus ornatum Stock, 1953a: 41, 43-44, fig. 6; 1953b: 304 [key].

Material

East London area. SAM—A19597, SM 163, 90 m, 1 &.

Remarks

This is apparently only the fourth specimen and second capture recorded

for this species. The types consist of a male and two females from the Agulhas

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bank, and the sexual characters of the male have not been described. From an

unreported male collected south-west of the Cape of Good Hope in 708 m in the

National Museum of Natural History collections, the following sexual characters

are noted. The cement-gland pores appear to be a series of 6—8 tiny cups in a

single row along the dorsal surface of the femur. The cement gland itself is not

discernible in this young male Cape specimen. The ovigers are very similar in

both sexes with the major segments of the male not much longer than those of

the female. The strigilis terminal claw is slender, with no serrations, and is

longer than the terminal segment. Sexual pores are very indistinct on the fourth

legs, second coxae of the male. Stock (1953a: 41) is correct in stating that this is

a highly characteristic, easily recognizable species. It should be noted that, in

the two specimens examined, neither has eyes on the tall blunt ocular tubercle.

Distribution

This is an apparently endemic South African species and the Cape of Good

Hope specimen is from the deepest known locality.

Genus Boehmia Hoek, 1881

Boehmia tuberosa Mobius, 1902

Fig. 3H

Béhmia tuberosa Mobius, 1902: 189-190, pl. 28 (figs 13, 14). Schimkewitsch, 1909: 3-4,

figs 1B, D. Barnard, 1954: 136-137, fig. 23.

Bohmia spinosa Mobius, 1902: 190 [/apsus calami].

Boehmia tuberosa: Stock, 1957: 96 [text]; 1962: 278.

Material

Transkei area. USNM 216759, SM 233, 540-580 m, 1 od.

Remarks

This species had been described as smaller than Boehmia chelata, but both

species should be on hand for this comparison to be relevant. It is easier to dif-

ferentiate between the males of the two species than to identify isolated females.

The male cement-gland area of B. tuberosa consists of a smooth or slick patch in

an otherwise papillose and setose femur dorsum, and contains two rows of tiny

transverse slits (Barnard 1954, fig. 23). The gland area covers the proximal half

or two-thirds of the femur. The gland area of B. chelata is shorter, broader

and has an uneven row of tiny circular pores in place of the slits present in

B. tuberosa.

The palps of the three known species of this endemic genus have seven or

eight segments. Boehmia longirostris (known only from a female) has seven palp

segments, while B. chelata and B. tuberosa have eight—the short basal segment

of these having been unnoticed by previous authors. A figure of the palp of

B. tuberosa is provided (Fig. 3H). The terminal segment is shorter than the pen-

ultimate segment, and in the single specimen on hand, the two terminal

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 129

segments measure only half or a little less than the length of the fifth segment.

Barnard (1954: 133) lists two other differences in his key. In B. chelata the

femur is shorter, measuring only three or four times longer than its width, and

the cephalic segment is parallel-sided or widens anteriorly. In B. tuberosa (the

Meiring Naude specimen), the femur is slightly longer, being 4,2 times the

width, and the cephalic segment narrows anteriorly. The cephalic segment

anterior in B. longirostris is more or less parallel-sided, but the leg is not

figured, nor are measurements given. The proboscis of the latter species is a

longer tube than in the other two species and its lateral processes are spaced

further apart.

Distribution

Of the few collecting records for B. tuberosa, the depth ranges from 11 m

to 550 m, and it has been taken from off Cape Town and the Agulhas Bank to

the East London area and now the Transkei area.

Through the courtesy of Dr G. Rack (Zoologisches. Institut und Zoo-

logisches Museum, Hamburg), we have been permitted to examine the holotype

of B. longirostris. This is a smaller species than the two others of the genus. It is

similar in habitus to B. chelata, but the following differences are evident: the

legs decrease in length from anterior to posterior on the trunk; the three coxae

are subequal in length with the second swollen with genital pores; the femur is

about 1,5 times its width and bears many acute conical tubercles, each bearing a

seta; the tibiae are shorter with tibia 2 slightly longer than tibia 1; the proboscis

reaches 0,6 as long as the ventral trunk, while that of B. chelata is only half as

long as the trunk; and the abdomen is articulated, carried ventrally and is square

tipped. The following key is presented to help separate the three known species.

There is controversy as to whether or not B. dubia Hedgpeth, known from a

larva, is actually a member of this genus, and it is omitted from the key.

1. Legs about four times as long as body; cephalic segment narrow anteriorly;

femuUn(o )4—5 times longer than’ broad .)........2.......5.. B. tuberosa

— Legs not more than twice as long as body; cephalic segment parallel-sided

or widened anteriorly; femur little more than twice as long asbroad ...... 2

2. Palp 7-segmented; proboscis length equals 0,6 of ventral trunklength ......

ey eee Ce ee Git oe a ne eas eo B. longirostris

— Palp 8-segmented; proboscis length equals half of ventraltrunklength .....

sisiahl 21S oak SRLS SP etd ck oR oe ee B. chelata

Genus Cilunculus Loman, 1908

Cilunculus sewelli Calman, 1938

Cilunculus sewelli Calman, 1938: 161-163, figs 8-9. Clark, 1963: 75. Stock, 1968: 13.

Ammothella gigas Fage, 1956: 173-175, figs 9-14.

Material

East London area. SAM—A19607, SM 185, 90 m, 3 juvs.

Transkei area. SAM—A19608, SM 226, 710-775 m, 1 juv.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks and distribution

There is nothing remarkable about the two sets of juveniles except for the

great difference in their depths of capture. The 90 m at SM 185 marks the shal-

lowest this species has ever been taken, but its wide depth range of 183-1 789 m

easily permits the addition of shallower or deeper depths to the record. This

species has been taken off the Natal coast, Zanzibar, and as far east as New

Zealand.

Genus Tanystylum Miers, 1879

Tanystylum thermophilum (Barnard, 1946)

Austroraptus thermophilus Barnard, 1946: 62; 1954: 146-148, figs 29-30.

Tanystylum thermophilum Stock, 1956: 92-93; 1957: 94-95, fig. 12; 1962: 277. Clark,

19773336:

Material

Transkei area. SAM—A19596, SM 250, 150-200 m, 2 do.

Remarks

These are typical males as figured by Barnard (1954, figs 29-30). The

species has a six-segmented palp and a curved bottle-shaped proboscis, which

makes it easy to distinguish.

Distribution

The maximum depth at this station off Transkei is the deepest at which this

species has been taken (previously recorded from 11 m to a little over 100 m).

Family Phoxichilidiidae Sars, 1891

Genus Anoplodactylus Wilson, 1878

Anoplodactylus typhlops Sars, 1888

Anoplodactylus typhlops Sars, 1888: 341-342; 1891: 29, pl. 2 (fig. 3a—e). Hedgpeth, 1948: 228,

229, fig. 29a—c [literature]. Child, 1982a: 21.

Anoplodactylus neglectus Hoek, 1898: 293-295, figs 7-10.

Anoplodactylus pelagicus Flynn, 1928: 25-27, fig. 14a—b. Barnard, 1954: 128, fig. 19a—g.

non Anoplodactylus pelagicus Stock, 1981: 463-464, fig. 6.

Material

Durban area. SAM—A10223, SM 123, 690 m, 1 o.

Remarks

This male specimen closely resembles both A. pelagicus Flynn and A. typh-

lops Sars. The blind flattened ocular tubercle, the large gaping chelifores, the

legs on which the cement-gland tube rises distally at 0,7 of the femur length, and

the general habitus all conform to both descriptions. We believe they are insep-

arable and have little hesitation in synonymizing Flynn’s species with Sars’.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 131

We do not believe that Stock’s nine specimens reported (as A. pelagicus)

from campaign ‘Walvis 1’, station 75, 32°18,2’S 13°15,9’E, in 3 550 m, are this

species. Stock (1981: 464, fig. 6a) illustrated three different acutely pointed

ocular tubercles for Flynn’s species, while the original description and figure

stated ‘the ocular tubercle is vey low and flattened on top’ (Flynn 1928: 25,

fig. 14b). The oviger and legs of Stock’s specimens are very close to this species,

but the tall ocular tubercle is so unlike that of Flynn’s and Sars’ species descrip-

tions that we believe the Walvis specimens may be another or new species.

There is a curious anomaly present in the Meiring Naude male. The cement-

gland tubes are found only on the anterior three pairs of legs and not on the

posterior pair. This specimen must be an adult as the sex pores appear ventrally

on the second coxae of the posterior pair of legs. Perhaps the posterior legs were

lost and then regenerated. The cement-gland duct joins the femur at a more

obtuse angle (135°) than those shown by Flynn (1928, fig. 14a).

Distribution

This species, under its various names, is a deep-water blind species of wide

distribution. It is known from the North and South Atlantic and its range

extends into the Indian Ocean in the sub-Antarctic zone of Prince Edward and

Crozet islands, and it has been taken in the Gulf of Alaska. Its depth range

extends to over 3 600 m.

Family Austrodecidae Stock, 1954

Genus Pantopipetta Stock, 1963

Key to the genus Pantopipetta

Pe NUalitamy claws PLeSEMeseVES PNESEME (258.2 4.8 ay. oe. s se ec Se ne es 2

— Auxiliary claws lacking; eyes present, lacking, or notevident............ 3

2. With very tall spurs on lateral processes; auxiliary claws from 0,3 to 0,4 of

Aida Claw alle MNP UIN are A kee eer Ay iy Neon Laan nic eo hone de s God hus P. auxiliata

— Without spurs or tubercles on lateral processes; auxiliary claws less than

OS arama lava le mp Gli dere: eee Fal eset il lard a aeus's gee os P. oculata

3. Lateral processes without distal spurs or tubercles; palps with 4 distal

segments; first coxae without spur; eyes lacking or not evident ........... 4

— Lateral processes with tall or short spurs or low tubercles; palps with 3 or

4 distal segments; first coxae with or without spurs; eyes present, lacking, or

MNOGCVACCIGhos rennet een rend Cec ea hc e a tic 8a Tal ete «8 6

4. Tarsus very short; tarsus—propodus length ratio 1:7; propodal claw

NUN OM GSE Ta AVIONS Pee. ircke dR Gaee eae ade Goduit o& aes Gala wikiyels Gland rack P. australis

— Tarsus longer, tarsus—propodus length ratio from 1: 1,5 to 1: 2,5; pro-

podaltclawawithionwilioOuluserralions| meee lee est) eon ))

5. Tarsus—propodus length ratio 1: 2,5; palp segment 5 equal to terminal

segment; propodal claw smooth (including P. brevicauda)................

956.160. oo. CERES csisics oie Stes Ries CRA EO ee npn a eee P. longituberculata

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

— Tarsus—propodus length ratio 1 : 1,5; palp segment 5 distinctly longer than

terminal segment; propodal claw endally serrate .............. P. weberi

6. Palp with 3 short distal segments; eyes present or lacking ............... 7

— Palp with 4 short distal segments; eyes lacking or notevident ........... 10

7. Lateral processes with low rounded tubercles, not taller than wide; first

coxae with 2 low slender tubercles; eyes lacking............... P. aconae

— Lateral processes with tall spurs or tubercles; first coxae with or without

Spurs or tubercles: eyes present omlacking 45 4..0 ee ee 8

8. Lateral processes with low tubercle on first pair, tall spur on posterior

3 pairs; first coxae and femorae with broad rounded dorsal tubercles; eyes

PFESEME Se ye ne he eee net Arn eet P. bilobata sp. nov.

— All lateral processes with tall spurs; first coxae with or without tubercles or

spurs; eyes lacking or nOt evident yy. 8 aia ae eile ae a ayes ea 3)

9. First coxae with tall dorsal spur and shorter ventral spur, matching those of

lateral PROCESSES. 02 ie Nene aie ee nee eel ns rcON enue P. armata sp. nov.

-—— First’coxae withoutspurso1 tubercles: 7405-224 P. armoricana

10. Lateral processes with tall spurs on posterior 3 pairs only; propodal claws

Slightly’ Se trates (ie se yale tes Ee a WOR ita te i are ae AE mene te P. capensis

— Lateral processes with low inconspicuous tubercles; propodal claw smooth

EAN mim cA nm MM LalEts MMI usta g's" 5.¢0! 0 0 « I.

11. Femur with large dorsodistal tubercle almost as long as segment diameter;

tarsus—propodus length ratio 1 : 2,6; propodal claw shorter than tarsus

wns la SHOE Bild ve nlc ve ak RS ace area IMO a MR Pa Ve P. angusta

— Femur with very low dorsodistal tubercle less than half segment diameter;

tarsus—propodus length ratio 1: 3,1; propodal claw longer than tarsus

diet 6 Sle aa ccc Ca Genta al eo] ADA ah DUATUID RN) ae ann Ue Ne Ue ee P. lata

Pantopipetta armata sp. nov.

ieee

Material

Holotype, SAM-—A19574, SM 103, 28°31,7’S 32°34,0’E, 680 m, 1 6. Para-

types, SAM-A19575, 16, 1. Paratype, USNM 228131, SM 103, 1d.

Paratype, MNHN-Py 591, SM 103, 1 6. Paratype, SAM-—A19576, SM 109,

28°41,0’S 32°36,8’E, 1 300 m, 1 &.

Description

Male. Trunk completely segmented, including segmentation line at base of

abdomen. Lateral processes long, widely separated, armed with long slender

spur on first pair, slender spurs twice length of first pair on posterior three pairs,

and shorter slender spurs on ventrodistal lateral processes. Abdomen reaching

to midlength of coxa 2 of fourth pair of legs, slender, cylindrical, glabrous.

Proboscis slightly swollen proximally, tubular and annulated in distal two-thirds,

almost as long as trunk, curved slightly ventrally. Palps seven-segmented,

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 133

second segment longest, fourth segment 0,6 of second segment length. Terminal

three segments armed with ventral short setae, sixth segment twice length of

fifth, seventh only a tiny knob.

Ocular tubercle a tall erect cone tapering to small distal bulb, without eyes.

Oviger 10-segmented, typical of genus. Strigilis segments increasingly smal-

ler, armed with short broad denticulate spines in formula 3: 2:2: 3, with

terminal claw only slightly larger than spines.

Legs long, slender, armed with few distal setae and long dorsodistal seta

longer than segment diameter on major segments. First coxae with tall dorso-

distal spur and short ventrodistal spur, third coxae with matching dorsodistal

Fig. 2. Pantopipetta armata sp. nov. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Third leg, with enlargement of sole spine. D. Distal seg-

ments of palp. E. Oviger. F. Distal segments of oviger.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

spur. Cement-gland tube tapered, longer than femur diameter, placed distally

on anterolateral or posterolateral surface and extending beyond flexed first tibia.

Anterior or posterior gland placement random, without apparent pattern.

Tarsus short, about 0,25 propodal length, both segments armed with row of

short sole spines, few ectal setae. Claw 0,4 propodal length, auxiliary claws

lacking. Femur and tibia 2 equal in length, tibia 1 slightly longer.

Female. Only slightly larger than male, except for oviger which is smaller,

with fewer denticulate spines.

Measurements (holotype, in mm)

Trunk length, 1,66; trunk width (across 2nd lateral processes), 1,3; pro-

boscis length, 1,54; abdomen, 0,84; third leg, coxa 1, 0,36; coxa 2, 0,35;

coxa 3, 0,24; femur, 0,94; tibia 1, 0,98; tibia 2, 0,94; tarsus, 0,12; pro-

podus, 0,49; claw, 0,2.

Remarks

Of the 12 known species of Pantopipetta, none has the slender spurs on

both the dorsal and ventral surfaces of the lateral processes and first coxae. At

least one other species (P. capensis) has reduced spurs on the first pair of lateral

processes; others have no spurs but instead low tubercles (as in P. bilobata), but

the presence of ventral spurs is sufficient to make this a new species.

Distribution

The stations at which these specimens were taken are on the Natal coast,

between the St Lucia Estuary and Richards Bay, and this blind deep-water

species was taken in 680 m and 1 300 m.

Etymology

The specific name ‘armata’ refers to the unusual lateral process and first

coxa armament of dorsal and ventral spurs.

Pantopipetta bilobata sp. nov.

Fig. 3A-—G

Material

Transkei area. Holotype, SAM—A10256, SM 250, 31°59,3'S 29°22,5’E,

150-200 m, 16. Paratypes, SAM-A10260, SM 250, 4 9. Paratype,

MNHN-Py 592, SM 250, 1 &.

Zululand area. Paratypes, SAM-—A10259, SM 86, 27°59,5’S 32°40,8’E,

Sine IS. It 2 |

East London area. Paratypes, SAM—A10257, SM 163, 33°04,6’S 28°06,6’E,

90m, 3 992 PRaradtypes, WWSNM 228132) SMolosi 1G al as barabyper

SAM-—A10258, SM 164, 33°04,6’S 28°06,6’E, 90 m, 1 &.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 135

Description

Male. Trunk completely segmented including abdomen base. Lateral

processes moderately long, about twice their maximum diameter, separated by

approximately their own diameters, armed with tiny low single or double

tubercles on first pair, and very tall slender spurs on others. Without ventral

armament. Tall slender spurs may be present only on posterior two pairs of

lateral processes or only on posterior pair. Ocular tubercle tall, erect, tapering

Fig. 3. A-—G. Pantopipetta bilobata sp. nov. A—F. Holotype, male. A. Trunk, dorsal

view. B. Trunk anterior, lateral view. C. Third leg, with enlargement of cement-gland

tube. D. Oviger. E. Oviger distal segments. F. Distal palp segments. G. Paratype,

female. Femur. H. Boehmia tuberosa Mobius. Palp.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

to tiny eyes at tip. Proboscis typical of genus, swollen proximally, tubular and

annulated distally, 0,25 longer than trunk length, down-curved distally.

Abdomen moderately robust, cylindrical, extending to distal end of second

coxae of fourth legs, armed with pair of short distal setae.

Palp seven-segmented, second longest, fourth 0,6 as long as second.

Terminal three segments heavily setose ventrally, setae longer than segment

diameters. Sixth segment slightly longer than fifth, seventh only a knob.

Oviger of typical recurved shape; second, fourth and sixth segments sub-

equal. Strigilis segments of increasingly reduced size distally, armed with an

ectal seta and endal denticulate spines in the formula 3 : 3 : 3 : 2, with terminal

claw the same size as spines. Spines slender, margins closely serrate.

Legs long, slender, armed with few short setae and a very long spine dorso-

distally on a femur tubercle as long as femur diameter and on single dorsodistal

swellings on both tibiae. Femur with conical cement-gland tube ventrally along

0,6 of its length and large broad-based tubercle dorsally just opposite cement-

gland tube. First coxae with large broad-based dorsodistal tubercle and tiny hint

of a low tubercle on ventral side. Third coxae with tall slender dorsal tubercle or

spur. Tarsus short, slightly over 0,2 propodal length, armed with few ventro-

distal setae. Propodus slender, well curved, with few ectal setae and row of short

sole spines. Claw robust, short, only slightly longer than tarsus, without

serrations.

Female. Entire animal slightly larger than male except for oviger which is

typically smaller. Femur distinctive from that of male. Dorsal surface with two

broad low tubercles, the distal tubercle larger than the proximal one and dorso-

distal slender tubercle bearing spine longer than in male, measuring 1,5 the

segment diameter. Dorsal broad tubercles only on posterior four legs and absent

from anterior two pairs.

Measurements (holotype, in mm)

Trunk length, 1,5; trunk width (2nd lateral processes), 1,25; proboscis

length, 2,1; abdomen, 0,71; third leg, coxa 1, 0,41; coxa 2, 0,36; coxa 3, 0,21;

femur, 0,96; tibia 1, 1,11; tibia 2, 1,06; tarsus, 0,12; propodus, 0,53; claw, 0,15.

Remarks

This species is characterized by the presence of eyes, lack of auxiliary claws,

and by the curious variability of the tall lateral-process spurs occurring on the

posterior six, or four, or only on the last pair of lateral processes. This latter

character is not exactly unique among species of Pantopipetta, where at least two

species lack tall spurs on the anterior pair of lateral processes, but no other

species appears to have this ‘diminishing’ of spurs toward the anterior in some,

but not all, specimens examined. Perhaps this is a sexual character related to

maturity of the gonads, but many more specimens would have to be examined,

including ovigerous specimens, in order to confirm this conjecture. This charac-

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 37

ter of ‘gain’ or ‘loss’ of tall spurs appears sufficient in our opinion to propose this

as another new species of Pantopipetta.

In the twelve species of Pantopipetta, including the two new species

described herein, there are only three with eyes (a few others are described in

which the eyes are not evident and supposedly are lacking), while the majority

(10) lack auxiliary claws. Besides this species, P. auxiliata and P. oculata have

eyes, but the latter two have auxiliary claws whereas the new species lacks them.

The majority of species have four short distal palp segments, except for four

species—including P. bilobata—that have three segments. The other species

with three short segments are P. aconae, P. armoricana, and P. armata (des-

cribed herein, see pp. 132-134). Pantopipetta bilobata differs from P. aconae,

which has low pointed lateral-process tubercles, paired tubercles on the first

coxae, and other morphological differences. The other two species bear long

slender tubercles of varying lengths on all eight lateral processes, have other

tubercles where P. bilobata has none, and also lack eyes.

Distribution

This new species appears to be a typical element of the eastern warmer-

water pycnogonid fauna of South Africa and has a rather wide geographic and

bathymetric distribution from Zululand to East London in 90-550 m.

Etymology

This species name refers to the bilobed appearance of both the male femur,

with a dorsal tubercle and opposed ventral cement-gland tube, and the female

femur, with its twin dorsal tubercles.

Family Callipallenidae Hilton, 1942

Genus Callipallene Flynn, 1929

Callipallene phantoma (Dohrn, 1881)

Pallene phantoma Dohrn, 1881: 196-197, pl. 14 (figs 1-9).

Callipallene phantoma Child, 1982a: 26-27 [literature]. Arnaud, 1987: 49.

Material

Durban area. SAM—A10225, SM 129, 850 m, 1 6 with larvae.

Remarks

This specimen possesses the typical habitus of the species, the long neck,

long legs with few spines, and straight propodus without a heel, but with long

auxiliary claws measuring over half the main claw. The auxiliaries of this

specimen have basal denticulation like those figured for C. panamensis Child,

1979 (fig. 14e), and the proboscis is less angular than that of Mediterranean

specimens.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (in mm) of the third right leg of this specimen are:

coxa 1, 0,116; coxa 2, 1,23; coxa 3, 026; femur, 15955 tibial 7 tibiaae eee

tarsus, 0,05; propodus, 0,51; claw, 0,34; auxiliaries, 0,25.

Distribution

This species is distributed from the Mediterranean to New England,

Florida, the Bahamas and Japan. It is a new record for the Indian Ocean off

South Africa and was taken at a surprising depth of 850 m. Other known collect-

ing depths are much shallower (3 m from Sargassum linifolium— Krapp-Schickel

& Krapp 1975, and from 10 to 35 m in Marseille Gulf, from Posidonia oceanica

blades— Arnaud 1987: 49).

Callipallene africana sp. nov.

Fig. 4

Material

Transkei area. Holotype, SAM—A10226, SM 226, 32°28,6’S 28°58,8’E,

710-775 m, 1 3 (incomplete). Paratypes, SAM—A10217, SM 226, 710-775 m,

2463 (damaged, without legs). Paratype, SAM—A10227, SM 250, 31°59,3’S

29°22,5'E, 150-200 m, 1 (damaged, 1 complete leg only).

Description

Male. Size moderately small, leg span 24,3 mm. Trunk completely seg-

mented, lateral processes short, 1,5 times longer than their diameters or less,

separated by slightly less than their diameters, glabrous. Neck moderately long,

posterior parallel-sided part as long as anterior expanded part, or ‘crop’. Oviger

implantation well in front of first lateral processes, against posterior extremity of

neck. Ocular tubercle low, blunt, as tall as diameter, with slender tubercle at

apex not as tall as ocular tubercle itself. Eyes present, slightly pigmented.

Abdomen short, blunt, semi-erect, glabrous. Proboscis short, not as long as

neck, blunt, broad, without ventrodistal bulges, lips flat.

Chelifores massive, curved against proboscis anterior. Scape not as long as

proboscis, slightly curved inward, armed with few lateral and distal setae. Chela

broad, semiglobular, armed with fringe of setae before fingers. Fingers straight

except at curved tips, armed with row of low crenulations.

Ovigers typical, third segment slightly elongate, fourth about 0,74 length of

fifth, which has distal apophysis and row of six or seven ectal setae. Strigilis seg-

ments increasingly shorter distally, armed with monomorphic denticulate spines

in the formula 12 : 11: 11: 12. Spines bear one or two broad proximolateral

teeth and finely serrate distolateral edges. Distal spines longer than proximal

spines. Legs missing.

Female. Trunk and appendages slightly larger than male, oviger fifth

segment shorter, without distal apophysis.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 139

Fig. 4. Callipallene africana sp. nov. A-D. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Oviger, with enlargements of proximal and distal denticulate

spines. D. Chela. E. Paratype, female. Third leg, distal segments.

Legs elongate, armed with very few setae not as long as segment diameters.

Femur and first tibia equal in length, second tibia about 0,2 longer. Propodus

very slightly curved, without heel, armed with five or six heel spines arranged in

two rows, many shorter sole spines flanked by row of short lateral setae. Claw

moderately slender, slightly curved, auxiliaries about 0,6 main claw length.

Measurements (in mm)

Holotype, male: trunk length 2,22; trunk width, 0,92; proboscis length, 0,7;

abdomen length, 0,23.

Paratype females sles——coxa) lm 0)32-scoxa 2, 1-37; coxa 3, 0,460; femur,

2,76; tibia 1, 2,75; tibia 2, 3,37; tarsus, 0,12; propodus, 0,54; claw, broken.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

There are no long-necked species of the genus Callipallene known in south-

ern African waters. We hesitate to describe this as a new species for lack of

suitable undamaged material, but it is unlike anything known from the region.

There are two species known from South Africa, Callipallene vexator Stock,

1956, and Callipallene sp. Barnard, 1954, each with sufficient differences as to

prevent confusion with this new species.

The taxonomy of this genus is in a state of confusion and there also seems

to be some evidence that neck length is a function of age (Stock 1952: 10,

figs 25-27). Callipallene phantoma (Dohrn) and its controversial subspecies

appear to be the nearest relatives of the new species, but they have longer

necks, many more setae, different leg length ratios, and other characters that

differ from this new species. They are also separated from the new species by

vast geographical distances and, although this is never sufficient reason to name

a new species, this is a contributing factor, in addition to the morphological

differences, in naming this new species.

The combination of moderately long neck, five or six heel spines in two

rows, lack of significant setae and spines on the legs, presence of chela finger

serrations, and the compact trunk segments are sufficient, we believe, to

describe this as a new species.

Distribution

Known only from the type locality, the Transkei area, in 710-775 m, and

slightly to the north-east in 150-200 m.

Etymology

The specific name for this species is derived from the African continent, its

place of origin.

Callipallene sp.

Material

East London area. SAM-—A10228, SM 163, 90m, 16 with eggs.

SAM-—A10247, SM 185, 90 m, 1 9.

Remarks

These damaged specimens may represent a new species, but they share

many characters with known species and are sufficiently damaged that we refrain

from designating them either as known or new.

The trunk is either completely segmented or lacks the posterior suture

between third and fourth segments. The lateral processes are compact, separ-

ated by less than their diameters and only slightly longer than their maximum

diameters. The neck is short and the chelifores are robust and short. The palm

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 141

has many ectal setae and the fingers, about as long as the palm, have serrate

teeth. The abdomen and proboscis are quite short and the proboscis has ventro-

distal bulges.

The oviger is typical, with a large fifth segment apophysis in the male. The

strigilis spines are alike and have the formula 10: 8: 8: 10. The legs are robust,

with few setae, the femur is subequal to tibia 1, and tibia 2 is longest. The pro-

podus is very setose lateral to two distinct rows of sole and heel spines. There

are four to six heel spines. The claw is short and the auxiliaries are over 0,7 as

long as the claw. Both auxiliaries have endal rugosities or patches of tiny blunt

spines numbering from five to eight.

These specimens share characters of Callipallene belizae Child, C. pectinata

(Calman), C. californiensis Hall, and C. emaciata (Dohrn). The pectinate or

rugose auxiliary claws do not appear on all specimens of the species bearing

them (C. pectinata, C. panamensis, and possibly others). Therefore, it is not

always a reliable taxonomic character in the genus.

This unnamed species is closest to C. pectinata, also its closest geographical

associate. Although C. pectinata has much larger rugosities (even teeth) endally

on the auxiliary claws, the leg and propodus are similar except for having fewer

lateral propodal setae and a longer main claw. The neck and general trunk

habitus are the same in both species, as are the ovigers and chelifores. It is poss-

ible that when more specimens of this species are taken from South African

waters, it will be found that they conform more closely with C. pectinata and can

be assigned to that species with more confidence.

Of the other species near these Meiring Naude specimens, C. emaciata and

C. belizae have shorter propodi with less curvature and fewer lateral setae.

Also, C. belizae lacks chelifore teeth and ventrodistal proboscis bulges. Callipal-

lene panamensis has a much longer propodus with fewer setae and spines, while

the propodus of C. californiensis has fewer lateral setae and auxiliary claw rugos-

ities that appear more like tiny setae than the blunt spines of Callipallene sp.

These specimens are, of course, quite different from the two known South

African species, C. vexator Stock, 1956, and Callipallene sp. Barnard, 1954.

Genus Pallenopsis Wilson, 1881

Pallenopsis capensis Barnard, 1946

Pallenopsis capensis Barnard, 1946: 62; 1954: 116 [key], 118-119. Stock, 1956: 83-84, fig. 7a;

19622 23519832 34-

Material

East London area. SAM—A10216, SM 185, 90 m, 1 6d, 1 &.

Remarks

These two specimens agree well with P. capensis as described. They have

the long auxiliaries, a few long spines, and a club-shaped sixth oviger segment

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

densely covered with minute spines or setae. The male of this species has a very

short cement-gland tube opening ventrally, slightly proximal to the middle of

each femur. The second left leg of the male measures 73 mm, whereas that of

the female is smaller.

Distribution

This is a typical South African species, known from the Cape to East

London area in 51-342 m.

Pallenopsis longirostris Wilson, 1881

Pallenopsis longirostris Wilson, 1881: 252-253, pl. 4 (figs 19-22), pl. 5 (figs 24-25). Hedgpeth,

1948: 210-211, fig. 21a—b [literature]. Arnaud, 1973: 150. Stock, 1981: 463, fig. 5a-f.

Pallenopsis oscitans: Barnard, 1954: 122, fig. 17.

Material

Zululand area. SAM—A10215, SM 107, 1 200-1 000 m, 1 d, 1 &.

Remarks

This is a large handsome species with well-pigmented eyes.

Distribution

It has a wide distribution, predominantly in the North Atlantic, but this

appears to be the first record of it in the Indian Ocean. Barnard (1954: 122)

recorded it from off Cape Point, so it is not surprising to find it off the coast of

Zululand. It has a depth range of 135-3 065 m.

Genus Pallenoides Stock, 1951

Pallenoides sp. nr amazonica Stock

Compare:

Pallenoides amazonica Stock, 1975: 1012-1015, figs 23-24.

Material

East London area. SAM—A10218, SM 185, 90 m, 1 larva.

Remarks

Despite its young stage of development (its fourth legs are only pointed

knobs), this specimen shares the main characters of an adult of P. amazonica as

originally described. It has three trunk suture lines, a short proboscis with

ventrodistal bulges, stout scapes and chelae with smooth movable fingers and

very small crenulations on the immovable finger, a low conical ocular tubercle, a

short angular abdomen, and robust legs. The anterior four legs are completely

developed, whereas the posterior four are less developed to undeveloped. The

eyes are not visible and the curved propodus has a heel with only two basal

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 143

spines and five shorter sole spines. The auxiliary claws are as long on the

anterior legs as in the type of P. amazonica.

The larva measures 1,44 mm in trunk length and 0,63 mm across the second

lateral processes.

It is possible that this specimen is either P. proboscideum Barnard, or

P. magnicollis Stock, both known from the South African area (Algoa Bay and

Luderitz Bay respectively). However, it lacks the median trunk, lateral process

and chelifore tubercles of P. proboscideum, and lacks the distal trunk setae and

has a different shaped proboscis than P. magnicollis. It agrees more in the

trunk, proboscis and leg characters with P. amazonica than with the two South

African species. Stock’s type was taken in northern Brazil, and although this

Meiring Naude record is certainly questionable, it would not be the first time a

pycnogonid species has such disparate distribution.

Genus Parapallene Carpenter, 1892

Parapallene algoae Barnard, 1946

Parapallene algoae Barnard, 1946: 61; 1954: 111-112, fig. 12a—c. Stock, 1956: 81-82, fig. Sa—e;

1962: 284.

Material

Zululand area. SAM—A10252, SM 86, 550 m, 1 &.

Remarks

This is a large, handsome, long-necked species and this specimen bears

several orange or white ova in its distended femorae. The trunk length, from the

base of the chelifores to the abdomen base, is 10,3 mm.

Distribution

This is apparently an endemic South African species, known from the south

and east coasts in 55-159 m. This record extends the known range well into the

Indian Ocean and, at 550 m, is from much deeper water.

Parapallene invertichelata sp. nov.

Fig. 5

Material

Zululand area. Holotype, SAM-—A19572, SM 86, 27°59,5'S 32°40,8’E,

550m, 1 larvigerous ¢. Paratypes, SAM-—A19573, SM 86, 550m, 2 2?

(1 without legs), 1 juv., 2 larvae.

Description

Male. Size fairly small, leg span slightly greater than 21 mm. Slender

species with long appendages. Trunk completely segmented, neck moderately

144

ANNALS OF THE SOUTH AFRICAN MUSEUM

WSs ie x

F ) y, WF

Fig. 5. Parapallene invertichelata sp. nov. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Third leg. D. Distal segments of leg. E. Oviger, with

enlargements of denticulate spine and terminal claw. F. Chela, with enlargement of

teeth.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 145

short, lateral processes separated by their own diameter or little more, as long as

1,5 times their maximum diameter, armed with short posterodistal seta on

second and third pairs. Ocular tubercle short, not as tall as its diameter; eyes

large, anterior pair larger than posterior pair, slightly pigmented. Abdomen

short, erect, armed with pair of short distal setae. Proboscis slender, constricted

in midregion, tapering distally to very small mouth bearing tightly packed fringe

of oral setae.

Chelifores large, carried in inverted or inferior position, arching ventral to

proboscis. Scape large, equal in length to proboscis, armed with several distal

setae. Chela long, fingers shorter than palm. Fingers swollen for most of length,

tapering to slender tips that cross when fingers close. Armed with many short

setae on immovable finger only, and row of coalesced teeth similar to propodal

lamina on both fingers.

Oviger moderately long, slender, third segment slightly longer than com-

bined length of first two, fifth longest, 0,2 longer than fourth, armed with

several lateral setae and low distal apophysis bearing three setae. Sixth through

tenth segments progressively shorter, armed with few lateral and distal setae.

Strigilis armed with denticulate spines in formula 16: 15: 14: 13, spines

slender, with pair of large basal serrations and many tiny serrations distally. Ter-

minal claw short, about half length of terminal segment, well curved, armed

with eleven short endal teeth.

Legs long, very slender, armed with short setae increasing in numbers dis-

tally. Second coxa slightly longer than first and third combined. Second tibia

longest with first tibia and femur progressively shorter. Tarsus very short, armed

with several ventral setae and single spine equal to ventrodistal spine on second

tibia. Propodus very slightly curved, without heel, armed with three heel spines

of equal length and row of eight or nine sole spines, which are flanked by rows

of short lateral setae. Claw robust, moderately curved, half length of propodus.

Auxiliary claws absent. Cement glands not evident.

Female. Sexual pores dilated on all second coxae, femorae with few ova.

Trunk size almost matching male, equally or slightly less setose. Oviger seg-

ments four and five equal, fifth without setose distal apophysis.

Measurements (holotype, in mm)

Trunk length, 1,97; trunk width (2nd lateral processes), 0,93; proboscis

length, 0,76; abdomen, 0,2; third leg—coxa 1, 0,29; coxa 2, 0,78; coxa 3, 0,36;

femur, 2,0; tibia 1, 2,23; tibia 2, 3,37; tarsus, 0,16; propodus, 0,6; claw, 0,3.

Remarks

This new species shares characters of both Parapallene and Pseudopallene

and, indeed, there is little to differentiate the two genera (Stock 1953b:

296-297; 1954: 50). Without wishing to undertake an analysis of both genera,

we propose this new species as belonging to Parapallene. Its proboscis is very

much like that of Pseudopallene hospitalis (Loman) (formerly Parapallene

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

hospitalis). One of the few differences between the two genera has been the

configuration of the lips and whether or not they have a corona of setae. This

means of differentiation has apparently been abandoned, as there are species in

both genera with differing lip configurations and with or without lip setae. The

presence or absence of chela finger teeth has been used as a differentiating

character, but there are known species in both genera with and without chelae

teeth. We have placed this new species under Parapallene because, in our

opinion, it shares more characters with that genus than it does with Pseudo-

pallene, according to Stock’s (1954: 50, 61) definitions of these genera.

The inverted flexure of the chelifores in this new species is very rare among pycno-

gonids and is unique among the species of Parapallene (and Pseudopallene).

The adults of this new species have an apparent wrinkle artifact anterior to

the oviger implantation on the neck.

Distribution

Known only from the type locality, off Zululand in 550 m.

Etymology

The specific name ‘invertichelata’ refers to the inverted flexion of the cheli-

fores present in all specimens examined.

Genus Pseudopallene Wilson, 1878

Pseudopallene gilchristi Flynn, 1928

Pseudopallene gilchristi Flynn, 1928: 23-25, fig. 13a—c. Barnard, 1954: 107. Stock, 1968: 39,

fig. 14e—g.

Material

Transkei area. SAM-—A10254, SM 226, 710-775 m, 1 larvigerous <4.

Remarks

This male specimen agrees well with the original description and figure 13b

of Flynn’s male, in regard to the proboscis shape and lip fringe, the chelae, legs

and ovigers. The few small differences involve the expected variation in oviger

spine counts and differences in measurement of various long segments. The claw

measures 0,7 of the propodus length and there are no auxiliaries present.

Despite the depth at which this specimen was dredged, the low, rounded ocular

tubercle bears well-pigmented eyes.

Distribution

This record apparently marks only the third capture of this species and its

range, from so few records, is restricted to the Indian Ocean from Transkei and

Port Natal to Lourengo Marques, from ‘tow net’ depth to the present record of

710-775 m.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 147

Safropallene gen. nov.

Diagnosis

Trunk elongate, completely segmented. Neck present, with oviger implan-

tation at midlength. Lateral processes well separated, without adornment.

Chelifores extremely elongate, scape one-segmented, chelae with many teeth.

Palps lacking entirely. Proboscis straight, cylindrical, without protruding lips.

Oviger long, slender, 10-segmented, strigilis with denticulate spines and serrate

terminal claw. Legs very slender, elongate, with few very long spines. Cement-

gland pore(s) not evident. Propodus without heel or long sole spines, claw long,

without auxiliaries.

Type species: Safropallene longimana sp. nov.

Etymology

The name is an anagram derived from the area of origin, southern Africa,

and ‘pallene’, the suffix used for most genera of this family. Gender feminine.

Remarks

This new genus joins 22 others in this broad-based but fairly homogeneous

family. In the latest key to the family (Child 1982a: 24-25), Safropallene can be

traced to couplet 14, where it joins Seguapallene in the list of characters. This

latter genus differs from Safropallene in its much shorter, more normal appear-

ing trunk and appendage lengths, and in the presence of auxiliary claws on the

two known species.

Safropallene longimana sp. nov.

Fig. 6

Material

Transkei area. Holotype, SAM-—A10243, SM 226, 32°28,6'S 28°58,8’E,

710-775 m, 1 3d. Paratypes, SAM—A10244, 5 299; SAM-—A10245, 4 dd, SAM

—A10229, 15 specimens, all SM 226, 710-775 m. Paratypes, USNM 216760,

SIVME206,, (102/75 mina, iG, 2 2.

Durban area. Paratypes, SAM—A10246, SM 129, 30°53,4’S 30°31,7’E,

850 m, 18 dd, 2Y. Paratypes, MNHN-Py 599, SM 129, 850 m, 2 dd, 2 22.

Description

Male. Moderately small, extremely tenuous, leg span 12,3 mm. Trunk fully

segmented, slender, with moderately long neck bearing oviger implantation

bulges at midlength, armed with two small anterolateral knobs at chelifore inser-

tion. Lateral processes separated by more than twice their diameters, about

twice their diameters in length, without adornment. Ocular tubercle low, broad-

based, with small unpigmented eyes, situated at anterior of cephalic segment.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Safropallene longimana gen. et sp. nov. A-F. Holotype, male. A. Trunk,

dorsal view. B. Trunk, lateral view. C. Trunk anterior, ventral view. D. Third leg.

E. Chela, with enlargement of teeth. F. Oviger. G. Paratype, female. Oviger with

enlargements of denticulate spine and terminal claw.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 149

Proboscis fairly short, cylindrical, slightly swollen distally, lips rounded.

Abdomen moderately long, sausage-shaped, glabrous, extending to distal rim of

first coxae of fourth legs.

Chelifores scapes almost twice proboscis length, armed with few distal setae

not longer than scape diameter. Chelae as long as scapes, carried folded inward

above proboscis, cylindrical, slightly curved. Fingers 0,3 of chela length, cross-

ing at tips, armed with 15 large teeth on immovable finger and 12-13 on

movable finger, teeth separated by gaps holding one or two very small teeth in

each. Chela armed at insertion of movable finger with small slender tubercle and

few setae mostly distally and ectally on immovable finger.

Palps entirely lacking.

Oviger very long, slender, fifth segment over 14 times longer than maxi-

mum diameter, fourth segment measuring 0,6 length of fifth. Sixth segment short,

curved; it and fifth armed with few setae. Strigilis segments subequal, each

armed with ectal seta and endal denticulate spines in the formula 8:6: 5: 6.

Spines with smooth proximal edges, distal edges closely serrate. Terminal claw

shorter than tenth segment, well curved, armed with three large endal teeth.

Legs very slender, long, major segments armed with few very long spines,

some over five times segment diameter. Femoral cement gland a long dark mass

0,6 as long as segment length, pores or tubes not evident. Tarsus slender, 0,6

propodal length, armed with few dorsal and ventral setae. Propodus slightly

curved, without heel, armed with few dorsal setae, a very long dorsodistal spine,

seven or eight short sole spines. Claw almost 0,5 propodal length, well curved,

auxiliaries lacking.

Female. Oviger fourth and fifth segments much shorter, almost equal in

length. Strigilis segments with denticulate spines in the formula 8:6: 4: 7,

with three teeth on the claw. Femur dilated only proximally in gravid specimens.

Measurements (holotype, in mm)

Trunk length, 1,57; trunk width (2nd lateral processes), 0,59; proboscis

length, 0,6; abdomen, 0,36; third leg—coxa 1, 0,18; coxa 2, 0,5; coxa 3, 0,24;

femur, 0,91; tibia 1, 1,25; tibia 2, 1,7; tarsus, 0,28; propodus, 0,54; claw, 0,24.

Remarks

This new species has several attributes found in the genus Nymphon, such

as the generally slender habitus, the oviger implantation, the heavily armed

chelae, and the long slender legs without auxiliary claws, but the total lack of

palps removes it from consideration in that genus. There is, to our knowledge,

no other known pycnogonid with such long chelifores that, when extended,

measure over four times the length of the proboscis.

Distribution

Known from Transkei and Durban areas with a bathymetric range of

710—850 m.

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The specific name lJongimana alludes to the very long characteristic

chelifores.

Family Pycnogonidae Wilson, 1878

Genus Pycnogonum Brinnich, 1764

Pycnogonum (Nulloviger) africanum Calman, 1938

Pycnogonum africanum Calman, 1938: 163-165, fig. 1OA—C.

Pycnogonum (Nulloviger) africanum Stock, 1968: 61, fig. 22A—B.

Material

East London area. SAM—A10250, SM 162, 630 m, 1 o.

Transkei area. SAM—A10251, SM 232, 620-560 m, 1 9°.

Remarks

Stock (1968: 61) created the subgenus Nulloviger to accomodate P. africa-

num and several other species without ovigers in the male, and this is confirmed

by the Meiring Naude male. This specimen (SAM-—A10250), while lacking

ovigers, is a male due to the placement of the sex pores, but fits Calman’s (1938)

female paratype better than his female holotype. It has lateral processes devoid

of the three tubercles, having only a single large conical one, and the ocular

tubercle is produced into a tall acute spine with four distinct but unpigmented

eyes. The three median pointed tubercles have some sparse short spines. First

coxae of each leg have two low posterior tubercles, second coxae with only an

integument of large granules, and third coxae devoid of ornamentation. The

male femur is distally produced into a shorter anterior and a longer posterior

tubercle, the first tibia has two subequal prominences, and the second tibia has a

brush of equal and very short spines over its ventrodistal third, which is con-

tinued over the sole of the tarsus and propodus. The reticulation is usually

inconspicuous, as stated by Calman, but the integument is granular with one

short spine on some of the granules.

Some measurements (in mm) of the Meiring Naude male are: trunk from

anterior of cephalic segment to base of abdomen, 5,22; proboscis, 3,44; and

abdomen, 1,46.

The female from station SM 232 is much larger than the male of SM 162.

Some of the female’s measurements (in mm) are: trunk, 4,59; proboscis, 6,31;

and abdomen, 1,15. The proboscis in the female is long and styliform and is

0,33 longer than the trunk. The female specimen is also more tuberculate with

three stout anterior-projecting median trunk tubercles which, like most of the

integument, are granular, of an uneven size, and bear a few short setae or

spines.

The male sexual pore is quite visible on the posteroventral face of the

second coxae on the fourth legs. The female legs are slightly larger than those of

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Sl

the male, but are otherwise similar. The differences between characters of the

male and female, as noticed by Stock (1968) are confirmed by these two

specimens.

Distribution

This species was known from off Zanzibar and Lourenco Marques in

183—1 228 m, and the above records extend its distribution further south in the

Indian Ocean.

Pycnogonum crassirostrum Sars, 1888

Pycnogonum crassirostre Sars, 1888: 340; 1891: 12, pl. 1 (fig. LA-H). Hedgpeth, 1948: 279,

fig. 52B [literature]. Stock, 1966: 402 [key].

Pycnogonum crassirostrum: Stock, 1975: 1086-1087, fig. 59e.

Material

Transkei area. SAM—A10255, SM 232, 620-560 m, 1 6 juv.

Remarks

This small subadult specimen agrees more closely with Pycnogonumi crassi-

rostrum than with the similar P. guyanae Stock. It has the small rounded median

dorsal tubercle on the fourth trunk segment, the median dorsal trunk tubercles

are smooth (but granular), tibia 2 is slender, and the tarsus and propodus have a

dense ventral cover of spinules, bifurcate at their tips. The oviger has eight seg-

ments although it is not fully formed (the ultimate segment is a knob without the

claw) in this juvenile, the ocular tubercle is short, broad, and has four well-

pigmented eyes, and it is shorter than the three anterior median trunk tubercles.

Stock (1975: 1086-1087) remarked on the resemblances or similarities between

these P. crassirostrum and P. guyanae, but the differences appear to be suf-

ficient to keep the two separate.

Distribution

This species has been taken from water ranging from 100-400 m deep off

Iceland, Greenland, Norway, and eastern North America, and is thus a moder-

ately deep, cold-water form. It is surprising to find it in South African waters,

but less so if the cold capture depth is considered.

Pycnogonum forte Flynn, 1928

Pycnogonum forte Flynn, 1928: 31-33, figs 18, 19. Barnard, 1954: 152-153, fig. 31C—D. Stock,

1966: 402 [key].

Material

East London area. SAM-—A10249, SM 185, 90 m, 1 &.

Transkei area. SAM—A10248, SM 226, 710-775 m, 1 2, 1 juv.

1152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Neither the females nor the juvenile specimen shed any light on the ques-

tion of relationship between this species and Pycnogonum cataphractum Mobius

(1902: 194, pl. 30 (fig. 11)). The specimen figured by Mobius is a male, for

which he indicated the absence of ovigers despite its bearing eggs. Flynn (1928)

figured a female of P. forte, as did Barnard with a second specimen. The above

three specimens appear from the scant literature to be the third and fourth cap-

tures of this species, but still the male is either unknown or is the more

tuberculate P. cataphractum as figured by Mobius. The male in many genera of

pycnogonids is the more spinose, tubercular or setose of the two sexes, so the

more tubercular male specimen figured by Mobius may possibly be the male of

this species. However, some Pycnogonum species—such as P. africanum,

P. anovigerum, P. cataphractum, and possibly others known from females only

—lack ovigers in the male, so we cannot be sure until a male with eggs is taken.

The female status of these two adult specimens was confirmed by finding a

barely distinguishable genital pore on the posterior surface of the second coxae

of the fourth legs in both specimens (none on the second coxae of the other

legs), and groups of tiny eggs in a dissected leg of the specimen from SM 226.

Therefore, the presence or absence of an oviger in the male of this species

remains unknown.

The mid-dorsal trunk tubercles are apparently variable. Both Flynn (1928)

and Barnard (1954) illustrated a rather pronounced tubercle between the ocular

tubercle and the posterior tubercle of the cephalic segment. The corresponding

tubercle on the females of this collection is a low mound composed of many

papillae arranged on bumps all over the mound. The distal ends of all tubercles

on the trunk and legs are very papillose. The propodal sole has only a few short

pointed spines and none of the blunt bifurcate spines found on some Pycno-

gonum species.

Distribution

This species has been collected previously off the Gneka River near Port

Elizabeth and off Great Fish Point. Only one depth, 90 m (49 fathoms), has

been recorded in the literature, so the above records greatly extend the known

depth range to 775 m, and confirm that it is an endemic South Africa species.

Pycnogonum nodulosum Dohrn, 1881

Pycnogonum nodulosum Dohrn, 1881: 203-207, pl. 16 (figs 1-3). Loman, 1928: 62, 64. Fage,

1953: 381 [list]. Barnard, 1955: 106. Stock, 1958a: 5; 1966: 401 [key]. Arnaud, 1987: 50,

table 1.

Material

Zululand area. SAM—A10253, SM 86, 550 m, 2 °°.

Transkei area. SAM—A10224, SM 250, 150-200 m, 3 ¢°.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 153

Remarks

These five females are fully adult with sexual pores opening on the postero-

lateral side of fourth leg, second coxae. Their length ranges from 3,5 mm to

3,97 mm and width across the second lateral processes from 1,03 mm to

1,15 mm, sizes comparable with the Mediterranean specimens (Arnaud

1987: 50). The largest female (SM 86) is 3,97 mm long and bears three strong

almost columnar dorsal tubercles, larger than in the specimens from SM 250.

Distribution

This species was previously recorded from South Africa by Barnard (1955:

106), who listed a specimen from Algoa Bay in 9 m. It has also been found oc-

casionally in places between its type locality in the Mediterranean and Morocco

(Loman 1928), occidental Africa (Fage 1953) and South Africa. Its above occur-

rence off the Zululand and Transkei coasts is in deeper waters than previous

records, which are as shallow as 9m. Another female specimen was taken by

the University of Cape Town Ecological Survey in Saldanha Bay, at station

SB306T (33°03,7'S 17°58,5’) in 20 m, on 30 April 1963, on fine sand.

Family Colossendeidae Hoek, 1881

Genus Hedgpethia Turpaeva, 1973

Hedgpethia magnirostris sp. nov.

Fig. 7

Material

Zululand area. Holotype, SAM-19579, SM 103, 28°31,7’S 32°34,0’E,

680 m, 1 6. Paratype, SAM-—A19580, SM 103, 680 m, 1 &.

Description

Male. Size small for this genus, leg span about 34 mm. Trunk completely

segmented, with suture or separation lines crossing each segment. Each segment

posteriorly flaring, decreasing in size towards posterior. Lateral processes

extremely short, not as long as their diameters, separated by slightly less than

their diameters, armed with one dorsodistal seta or glabrous. Cephalic segment

narrow anteriorly, with small dorsolateral knob above palp insertion. Ocular

tubercle low, conical, as tall as basal diameter, eyes indistinct, unpigmented.

Abdomen extremely short, reaching to just beyond posterior lateral processes,

carried obliquely ventrally, glabrous.

Proboscis extremely large, 1,7 times length of trunk, proximally slender, in-

flating gradually to very swollen midpoint, tapering less gradually to slender

distal portion and small lips. Curved slightly downward in lateral view.

Palp third segment longest, slender, curved, fourth segment short, fifth

almost 0,6 length of third, sixth shortest, distal four segments almost subequal,

all armed with short setae, more numerous ventrodistally.

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Hedgpethia magnirostris sp. nov. Holotype, male. A. Trunk, dorsal

view. B. Trunk, lateral view. C. Palp. D. Oviger. E. Oviger terminal segment

and claw. F. Third leg, terminal segments.

Oviger fourth and sixth segments subequal, sixth armed with row of short

ectal setae. Strigilis four segments, increasingly short, armed with single ecto-

distal seta and row of spatulate endal spines. Terminal segment armed with

comb-like terminal spine forming subchela with terminal claw. Claw broad,

bottle-shaped with denticulate comb distally forming acute angle with tip.

Legs long, slender, armed with very short setae and few equally short spines

growing more numerous distally. Femur curved proximally, straight distally.

Tibia 2 only 0,64 length of tibia 1, femur 0,68 length of tibia 1. Tarsus slightly

shorter than propodus, both armed with very short sole setae, fewer short ectal

setae. Claw robust, well curved, equal in length to tarsus.

Female. Slightly larger in all measurements except for main oviger seg-

ments. Femorae slightly inflated proximally with ova.

Measurements (holotype, in mm)

Trunk length, 2,0; trunk width, 0,73; proboscis length, 3,36; third leg—

coxa 1; 0;4; coxa,2;055-1coxal3,, 0:53) femur, 34> tibia le s.0:etibiayZ or

tarsus, 1,2; propodus, 1,3; claw, 1,2.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 155

Remarks

For a genus hitherto containing only eight known species, Hedgpethia pre-

sents some problems to the taxonomist because the species are closely similar.

This new species differs from the eight known species by having a proboscis

much larger in proportion to the trunk than in any other species, and a shorter

ocular tubercle with indistinct eyes.

This species is most closely related to H. californica (Hedgpeth, 1939), and

H. bicornis (Turpaeva, 1958) (previously under the ‘articulatus-group’ (Loman,

1908)—lacking a proboscis tooth—of the genus Rhopalorhynchus). It differs

from these two in having a longer first tibia, thus making the leg segment ratios

different. It has a femur measuring only 0,68 the length of tibia 1, whereas the

femur is 0,82 of the first tibia in H. californica and is 0,8 in this ratio for

H. bicornis. The tarsus is shorter than the propodus (T = 0,92 of P) in

H. magnirostris, whereas it is equal to or longer than the propodus in H. califor-

nica and H. bicornis.

The main difference, besides greater proboscis size in the new species, is in

the ocular tubercle, which is a low truncate cone without a pointed apex,

whereas H. californica and H. bicornis both have a tall pointed ocular tubercle.

The eyes of both species are large and well pigmented, whereas those of the new

species are very indistinct and lack pigmentation. Both Meiring Naude

specimens agree in this character.

The new species shares with H. bicornis two small anterolateral cephalic

segment tubercles, which are not present in H. californica. But, H. bicornis has

middorsal trunk tubercles, whereas the other two have cowls or ridges that lack

these tubercles. The palps of H. californica have ‘a few scattered setae on the

terminal segments’ (Hedgpeth 1939: 460), whereas the palps of the other two

species are heavily setose on the terminal segments.

Distribution

Since the new species has been taken at a single station and depth, nothing

can be deduced about its distribution.

Etymology

The specific name is from the Latin magnus and rostrum, meaning a large

snout or beak, pertaining to the larger than usual proboscis of these specimens.

Hedgpethia sp. indet.

Material

Transkei area. SAM—A19585, SM 226, 710-775 m, 1 juv.

Remarks

This specimen has similarities with the species described above, but is too

juvenile for positive identification.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Rhopalorhynchus Wood-Mason, 1873

Rhopalorhynchus gracillimus Carpenter, 1907

Rhopalorhynchus gracillimus Carpenter, 1907: 99-100, pl. 13 (figs 25-32) [all except Saya de

Malha record].

Rhopalorhynchus kréyeri: Loman, 1908: 26-27 [only Siboga sta. 310 specimen]. Calman, 1923:

268-270 [part]. Barnard, 1954: 88-89, fig. 2.

Rhopalorhynchus gracillimum: Stock, 1958b: 128-132, figs 39-56.

Material

East London area. SAM—A19598, SM 164, 90 m, 1 2.

Remarks

This longitarsal specimen agrees in most characters with Carpenter’s (1907)

description and particularly with Stock’s (19585) figures 52-56. The ratio of tar-

sus—propodus-—claw to tibia 2 length is 0,72 to 1, well within the lower end of

Stock’s ratio figures of 0,7: 1 to 1: 1. The claw is slightly shorter than half the

propodal length.

The one character of the present material not in keeping with the previous

descriptions of this species is the form of the distal palp. The palp segments of

this female are notably shorter than in any figures of the palp given for the

species, and much shorter than those of the type specimen. The distal four seg-

ments are only slightly longer than their diameters, whereas the sixth segment is

about twice as long as its diameter. The shortness of the distal segments on both

palps may be due to regeneration or some other factor, and we feel that other

specimens from this locality should be examined before deciding whether or not

this character is variable in the species or if the East London material is attribu-

table to another species.

Distribution

This specimen adds nothing new to the known distribution of the species,

which has been recorded from the East Indies and Maldive Islands to the east

coast of South Africa, in depths from 0 m to 156 m.

Family Nymphonidae Wilson, 1878

Genus Nymphon J. C. Fabricius, 1794

Nymphon barnardi sp. nov.

Fig. 8

Material

Transkei area. Holotype, SAM-—A10261, SM 250, 31°59,3'S 29°22,5’E,

150-200 m, 1 @. Paratype, SAM—A10262, SM 250, 150-200 m, 1 2.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 157,

Description

Female. Moderately small size, leg span 26,3 mm. Trunk completely seg-

mented, glabrous, without tubercles. Lateral processes short, hardly longer than

their diameter, separated by half or slightly more of their diameters, glabrous.

Neck short, oviger implantation bulges touching first lateral processes. Ocular

tubercle not taller than wide, sides bulging, with tiny apical papilla, eyes large,

slightly pigmented. Proboscis moderately short, cylindrical, inflated slightly at

midlength, lips rounded. Abdomen cylindrical, tapering distally, reaching

slightly beyond posterior lateral processes, glabrous.

Fig. 8. Nymphon barnardi sp. nov. Holotype, female. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Oviger, with enlargement of denticulate spine.

D. Strigilis, with enlargement of terminal claw. E. Chela, with enlargement of

teeth. F. Third leg, terminal segments.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Chelifores typical, scapes as long as proboscis, armed with few lateral and

distal setae. Chelae moderately short, palm slightly longer than fingers, armed

with few short distal setae. Fingers slightly curved, overlapping at tips, armed

with 15 equal-sized teeth on immovable finger and 16 similar or slightly smaller

teeth on movable finger.

Palp segment 2 longest, third through fifth subequal, fairly short, armed

with many short ventral and distal setae.

Oviger fairly short, third segment slightly shorter than fourth, both with

few setae. Sixth segment half length of fifth, armed with few setae. Strigilis

segments progressively smaller distally, armed with few ectal setae, and a row

of denticulate spines in the formula 10: 7:5: 6; spines with four or five

serrations per side. Terminal claw shorter than terminal segment, moderately

curved, with row of eleven serrations extending around and forming part of the

tip of claw.

Legs long, moderately slender, armed with short setae increasing in

numbers distally. Tibia 2 the longest segment, followed by tibia 1, then the

femur. Tarsus slightly longer than propodus on first pair of legs, slightly shorter

than propodus on posterior three pairs of legs. Tarsus and propodus armed with

few short dorsal setae, a row of lateral setae and several sole spines with short

setae between each. Claw robust, moderately curved, about 0,4 length of

propodus, auxiliaries about 0,85 length of main claw.

Measurements (holotype, in mm)

Trunk length, 2,56; trunk width (across 2nd lateral processes), 1,39;

proboscis length, 1,15; abdomen, 0,44; third leg—coxa 1, 0,5; coxa 2, 0,72;

coxa 3, 0,4; femur, 2,38; tibia 1, 2,78; tibia 2, 3,95; tarsus (first leg), 0,62; pro-

podus (first leg), 0,58; tarsus (third leg), 0,7; propodus (third leg), 0,74;

claw, 0,29.

Remarks

This new species is closest to another small South African endemic species,

N. crenatiunguis. The general habitus of the two is very similar, but the differ-

ences are more than superficial. The chelae fingers are shorter and have more

teeth in N. crenatiunguis, and the third palp segment is much longer than the

short fourth and fifth segments. The ovigers of the two species are quite similar,

including the claw serrations that continue around the tip in a fan shape, but the

denticulate spines of N. crenatiunguis have three lateral lobes per side, whereas

the new species has spines with four or five narrower lobes. The femur is slightly

longer than the first tibia in N. crenatiunguis (femur shorter than tibia 1 in

N. barnardi), and in N. crenatiunguis the tarsus and propodus have many more

sole spines and shorter auxiliaries than in the new species. The tarsus is shorter

than the propodus in all legs we examined in N. crenatiunguis, but is slightly

longer than the propodus on the first pair of legs in the new species. We believe

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 159

these differences are sufficient to warrant creation of a new species for these two

females.

Distribution

This species is only known from one station, at a depth of 150-200 m.

Etymology

This species is named for Keppel H. Barnard (1887-1964) of the South

African Museum, whose contributions to the literature have greatly enriched

our knowledge of South African pycnogonids.

Nymphon bicornum sp. nov.

Fig. 9

Material

Zululand area. Holotype, SAM—A19577, SM 60, 27°09,6’S 32°58,2'E,

800-810 m, 1 o.

Durban area. Paratypes, SAM—A19578, SM 129, 30°53,4’S 30°31,7’E,

850 m, 2 29, 2 larvae.

Description

Of a small slender size, leg span 13,9 mm. Trunk long, slender, fully seg-

mented, including base of abdomen. Lateral processes separated by 1,5 to twice

their diameters, moderately short, slightly longer to 1,5 times as long as their

diameters, glabrous. Neck long, semicylindrical, slightly inflated at midpoint

with oviger bulges and ocular tubercle both well anterior to first lateral pro-

cesses. Ocular tubercle a low anterior-pointing cone with two moderately large

lateral sensory papillae, without trace of eyes. Proboscis of moderate size, cylin-

drical, constricted just behind anterior tip, lips rounded. Abdomen short,

cylindrical, constricted distally, with two distal setae.

Chelifores robust, scape slightly longer than proboscis, with a dorsodistal

seta. Chela longer than scape, slender, palm shorter than fingers, glabrous.

Fingers straight to gently curving distally and overlapping, armed with 40 small

curved teeth on immovable finger and 41 similar teeth on movable finger. The

immovable finger also bears several short setae.

Palp second segment longest, third 0,75 length of second, fourth half length

of third, fifth slightly longer than fourth, terminal three segments armed with

distal and ventral setae, some slightly longer than segment diameter.

Oviger fourth segment with ectal bulge proximally, segment 0,75 as long as

fifth, which is armed with ectal and endal rows of short setae. Sixth segment

slightly curved, half as long as fourth, armed with ectal and endal setae same as

fifth segment. Strigilis segments subequal in length, armed with two or three

ectal setae, endal denticulate spines in the formula 6:4: 3:5, spines with

spatulate centre lobe flanked by two serrations per side. Terminal claw shorter

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Nymphon bicornum sp. nov. Holotype, male. A. Trunk, dorsal view. B. Part of

anterior trunk segment, lateral view. C. Palp. D. Third leg. E. Distal segments of third

leg. F. Oviger. G. Strigilis, with enlargements of denticulate spine and terminal claw.

H. Chela, with enlargement of teeth.

than ultimate segment, slightly curved, armed with seven endal serrations not

including pointed tip.

Legs slender, tibia 2 the longest segment, major segments each armed with

few short dorsal and ventral setae and single long dorsodistal seta. Tarsus

slightly over half propodus length, both armed with row of short sole spines and

propodus with few lateral and dorsal setae. Claw robust, slightly curved, 0,6 as

long as propodus, auxiliaries lacking.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 161

Measurements (holotype, in mm)

Trunk length, 1,82; trunk width (across 2nd lateral processes), 0,78; proboscis

length, 0,6; third leg—coxa 1, 0,26; coxa 2, 0,48; coxa 3, 0,24; femur, 1,14;

tibia 1, 1,3; tibia 2, 1,58; tarsus, 0,4; propodus, 0,73; claw, 0,43.

Remarks

This new species appears to be most closely related to a deeper-water

species found in both the North and South Atlantic and Pacific oceans, N. pro-

cerum Hoek, 1881. Both have a long neck with the ocular tubercle and oviger

bases implanted well forward of the first lateral processes, and have a similar

long tenuous appearance. Both lack auxiliary claws and have long chelae fingers

with 40 or more teeth in each. The differences are in the longer neck, much

longer palp segments, longer chelae fingers with longer teeth, longer proboscis

and abdomen, and longer setae on the legs of N. procerum. The legs and ovigers

of the two species are very much alike, except that the oviger of N. procerum

has a terminal claw with about 20 fine teeth, instead of the seven or eight found

in the new species.

Distribution

The new species is known only from Natal and Zululand in 800-810 m and

850 m depths, whereas the depths known for N. procerum are much deeper,

including specimens from a trawl in over 6 000 m.

Etymology

The species name bicornum means two-lobed or two-horned and refers to

the two fairly large sensory papillae found on the ocular tubercle.

Nymphon comes Flynn, 1928

Nymphon comes Flynn, 1928: 14-16, figs 4-6. Barnard, 1954: 92-93, figs 3, 5C. Stock, 1956:

78-79, fig. 3; 1962: 282; 1965: 21-23 [list, text].

Nymphon affinis Stock, 1951: 5-7, figs 1-6. Barnard, 1954: 94.

Nymphon affine: Stock, 1965: 21-23 [list, text].

Material

East London area. SAM-—A19612, SM 179, 80 m, 1 &.

Transkei area. SAM—A19611, SM 233, 540-580 m, 1 do.

Remarks

Both of these fairly large specimens agree in most characters with descrip-

tions of both Nymphon comes and N. affine. Their differences lie in appendage

segment lengths, numbers of chela finger teeth, and oviger spine counts, all of

which can be attributed to variation within the species. The character that separ-

ated the two species, prior to our decision to unite them under N. comes, was

the presence or absence of alternating small and large teeth on the chelae

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

fingers. Both of these specimens have alternating large and small teeth, some-

times with two short teeth instead of one between two successive long teeth, in

agreement with Barnard’s (1954) figure 5c of specimens he listed as N. comes.

Since these have the alternating teeth and N. affine shares this character (Stock

1951: 6, fig. 3), we have united the two species under the senior synonym.

Flynn’s type specimen is not located at the South African Museum and is

unavailable at the moment for examination.

Distribution

This handsome species is another that has been found only along the South

African coasts and, although it has been taken in as shallow as 15 m (Stock’s

type specimen), the specimen from SM 233 was taken in 540-580 m, giving it a

rather wide depth range on the shelf and slope.

Nymphon crenatiunguis Barnard, 1946

Nymphon crenatiunguis Barnard, 1946: 60; 1954: 101-102, fig. 7.

Material

East London area. Syntypes SAM-—A10137, Pieter Faure, P.F. 907, 85 fm

(155 m), 1 6, 1 9. SAM-—A19609, Meiring Naude, SM 179, 80 m, 1 ¢.

Transkei area. SAM—A19610, SM 250, 150-200 m, 1 9.

Remarks

This apparently rare species is recorded in new material here only for the

second time. Barnard (1946, 1954) treated the same specimens—the types and

one other doubtful specimen. Through the courtesy of the South African

Museum, we have been permitted to examine the type series deposited in that

Museum.

There are several slight differences between the Meiring Naude material

and the type specimens. There are several more finger teeth on the SM 179 male

chelae (20 and 26), whereas the syntype male has 17 and 22, and both rows of

teeth appear more like the pointed ovals figured in Barnard’s (1954, fig. 7a)

enlargement of the movable finger teeth. The strigilis spine count is 10: 8: 6:7 in

the SM 179 male and there is a total of about five more spines on the female

strigilis. The tarsus is slightly shorter than that of the syntypes and the sole

spines on the propodus are fairly robust but of a single size. There are five

cement-gland openings protruding as low knobs ventrally on the femorae,

whereas the syntype male has either four or five per femur, the presence of

which add to the ease of recognition in this species.

Distribution

Barnard’s types were taken in 155 m, whereas the two Meiring Naude speci-

mens are from 80m and 150-200 m, respectively. The distribution of this

species is extended north-east to Port St Johns in the Transkei area.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 163

Nymphon distensum Mobius, 1902

Nymphon distensum Mobius, 1902: 179-180, pl. 25 (figs 1-6). Loman, 1923: 14 [key]. Flynn,

1928: 10-11. Gordon, 1932a: 105-106, fig. 6; 1932b: 28-29, [table], 35 [key]; 1944: 20

[key]. Barnard, 1954: 98-100, fig. 5b.

Material

East London area. SAM-—A10026, Pieter Faure, P.F. 12840, 17 mi SSW of

Buffalo River, trawled, 357 m, 2 66. SAM-A19600, Meiring Naude, SM 179,

80m, 1 ovigerous 6. SAM-—A19601, SM185, 90m, 2 22, 3 juvs.

MNHN-Py 594, SM 185, 1 ovig. ¢.

Zululand area. SAM—A19602, SM 86, 550 m, 1 6, 2 29°. USNM 228135,

SM 86, 550 m, 1 od.

Transkei area. SAM—A19599, SM 232, 620-560 m, 1 d.

Remarks

The male from SM 232 has 38 teeth on the immovable chela finger and

about 50 teeth on the movable finger, whereas most specimens in the literature

have from 60 to 70 and 30 to 33 teeth, respectively. The third leg tarsus is 0,85

as long as the propodus and the claw is 0,4 of that segment, differing slightly

from published figures. The strigilis has somewhat fewer denticulate spines than

in those described by Barnard (1954: 98). The formula is 12: 8:8: 8, with

9 spines on the claw. Other than for these slight differences, the specimen from

SM 232 and the others, including the Pieter Faure specimens, differ very little

from the type.

Distribution

Widely distributed from south of Cape Point to Zululand region, between

80 m and 560 m.

Nymphon granulatum sp. nov.

Fig. 10

Material

Transkei area. Holotype, SAM—A10230, SM 226, 32°28,6'S 28°58,8’E,

710-775 m, 1 d. Paratypes, SAM—A10232, SM 226, 710-775 m, 2 ¢¢. Para-

type, MNHN-Py 597, SM 226, 710-775 m, 1 6. Paratypes, SAM-—A10231,

SM 232, 32°14,9’S 29°10,4’E, 620-560 m, 1 2, 3 larv. Paratypes, SAM—A10235,

SM 236, 32°14,3’S 29°11,6’E, 670-660 m, 1 Q, 1 larv.

Durban area. Paratype, SAM—A10233, SM 129, 30°53,4’S 30°31,7’E,

od) ian, Ih

East London area. Paratypes, SAM—A10234, SM 162, 32°55,0’S 28°31,0’E,

630 m, 2 dd. Paratypes, USNM 228133, SM 162, 630 m, 1 d, 1 2.

Description

Male. Moderately small, legspan27,7 mm. Entire surface moderately papillose

except proboscis, imparting granular appearance. Trunk anterior fully segmented,

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Nymphon granulatum sp. nov. A-G. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Third leg. D. Palp. E.Chela. F. Oviger. G. Strigilis.

H. Paratype, female. Oviger, with enlargement of denticulate spine.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 165

posterior third and fourth segmentation line incomplete. Lateral processes robust,

separated by about half their diameters, as long as twice their diameters, armed with

dorsodistal and slightly laterodistal slender tubercles not as long as segment

diameter, capped with slender spine longer than segment diameter. Neck short,

expanded anteriorly to oblique angular chelifore insertions. Oviger implantation

well anterior to first lateral processes, ocular tubercle implanted directly dorsal to

ovigers, very tall, columnar, with two tiny papillae at apex, without eyes. Proboscis

cylindrical, slightly constricted proximally, lips almost flat. Abdomen long, extend-

ing to distal rim of second coxae, fourth legs, glabrous.

Chelifores large, scapes almost as long as proboscis, distal diameter almost

equal to proboscis base, armed with one or two lateral and distal setae. Chelae

longer than scapes, palm ovoid, shorter than fingers. Immovable finger moder-

ately curved, armed with slightly curved teeth of a similar size. Movable finger

straight except for curved tip overlapping immovable finger, armed with

33 smaller curved teeth only half size of those on immovable finger, of slightly

different sizes.

Palp moderately slender, second segment longest, third 0,6 as long as

second, both armed with few short lateral setae. Fifth segment slightly longer

than fourth, combined length only equal to second segment, both distal seg-

ments armed with few ventral and distal setae increasing in numbers distally,

none longer than segment diameter.

Oviger fourth segment with large ectal mound proximally, fifth longest,

straight, armed with ectal and endal rows of few setae, without distal apophysis

or setae. Sixth segment well curved, armed with few lateral setae. Strigilis seg-

ments slender, progressively shorter, armed with ectal seta distally and small

denticulate spines in the formula 5 : 3 : 2: 3; spines broad with single lateral

lobes. Claw almost as long as terminal segment, armed with three endal teeth.

Legs slender, long, armed with very few setae or spines. Second coxae with

ventrodistal tuft of setae as long as segment diameter. Femoral cement gland a

small sac at midpoint of segment, pore(s) not evident. Tibia 2 the longest

segment with tibia 1 and femur each slightly shorter than the last. Tibia 2 armed

with stout ventrodistal spine. Tarsus 0,6 as long as propodus, both armed with

very few tiny sole spines. Claw slightly less than half propodal length, almost

straight, auxiliaries lacking.

Female. Slightly larger overall than male. Oviger segments four and five

subequal, glabrous. Strigilis armed with fewer denticulate spines in the formula

3:2:2:2, with three endal teeth on the terminal claw of both female and

male. Sexual pores on coxa 2 ventrally on only posterior four legs.

Measurements (holotype in mm)

Trunk length, 3,24; trunk width (across 2nd lateral processes), 2,08; pro-

boscis length, 1,21; abdomen, 1,14; third leg—coxa 1, 0,58; coxa 2, 1,18;

coxa 3, 0,54; femur, 2,09; tibia 1, 2,28; tibia 2, 2,86; tarsus, 0,98; propo-

dus, 1,57; claw, 0,74.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This new species appears to be most closely related to the West African

species N. mauritanicum and N. prolatum, each having tall columnar ocular

tubercles, long abdomens and no auxiliary claws. This species differs from

N. mauritanicum in its slender lateral-process tubercles bearing long spines

instead of broad tubercles bearing many short setae, its longer palps and probos-

cis, its glabrous abdomen, and the legs that have far fewer setae. The chelifores

and ovigers are quite different in the two species.

It differs from N. prolatum in its much more robust and crowded general

habitus, shorter lateral processes, abdomen and neck, in having tubercles with

spines on the lateral processes, and in the chelae finger teeth that are more or

less the same size on any one finger instead of being at least two different sizes.

The legs, palps, proboscis, and ovigers are very similar in the two species except

that the fifth segment of the oviger has a distal apophysis with setae in N. pro-

latum. Eyes are lacking in the new species, whereas both of the West African

species have clearly defined eyes.

Distribution

Well established along the east coast of southern Africa, between 620 m

and 850 m.

Etymology

The new species name, granulatum, refers to the overall granular appearance

of the integument.

Nymphon microctenatus Barnard, 1946

Nymphon microctenatus Barnard, 1946: 60; 1954: 95-96, fig. 4. Stock, 1965: 22-23 [list, key].

Material

East London area. Holotype, SAM-—A10100, Pieter Faure, P.F. 12840,

17 mi SSW of Buffalo River, trawled in 357 m, 23 April 1901, 1 2.

Transkei area. SAM-—A19594, Meiring Naude, SM 237, 600-650 m,

1 gravid 9°.

Remarks

This is apparently only the second specimen of this large and very distinc-

tive species to be recorded. The specimen from SM 237 is from about 100 miles

north-east of where the type was taken and is from deeper water.

The chelae teeth are quite distinctive and make this an easily recognized

species. They number over 300 on each chela and appear similar to a fine-

toothed comb or much like the large propodal lamina found on many species of

the genus Anoplodactylus. The finger tips cross in the new specimen and may

have been broken off in Barnard’s type. The terminal palp segment appears

even more slender but no more curved than that of Barnard’s (1954: 95, fig. 4c)

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 167

type, and both the fourth and fifth segments are slightly longer in this specimen.

The terminal leg segments appear tiny in comparison to the very long major leg

segments. The sole spines of the tarsus and propodus are all of a single size and

in a single row. The oviger strigilis has the denticulate spine formula

15:11:10: 11, with 16 small teeth on the slender terminal claw. The spines

have 5—6 fine serrations on each side.

Nymphon modestum Stock, 1959

Nymphon modestum Stock, 1959: 556-558, fig. 3.

Material

Zululand area. SAM-—A19595, SM 86, 550m, 1d with eggs. USNM

228136, SM 86, 550 m, 1 d.

Transkei area. SAM—A19603, SM 239, 90 m, 1 gravid 2. SAM—A19604, SM

250, 150-200 m, 266, 12, 7juv. and larv. MNHN-Py 596, SM 250,

150-200 m, 1 o.

Remarks

We believe these specimens are N. modestum, although the variation dis-

played among these seven adults does not quite agree with the description of the

female type. Some differences seem superficial but others assume greater

importance among closely related species in the genus Nymphon.

The palp third segment is described as being about as long as segment 5 in

the type, whereas that of these specimens is variously 0,3 to 0,6 times longer

than segment 5. The abdomen varies from shorter than the fourth lateral processes

to as long as the distal end of coxa 1. The curved chelae of these specimens are dis-

tinctly shorter than the scapes that, in turn, are equal to the proboscis, whereas in

the type the chelae are longer than the scapes. The finger teeth vary in number

from 17 to 21 on the immovable finger and from 21 to 24 on the movable finger,

whereas those of the type number 23 and 27, respectively. Similarly, the oviger

strigilis spines number from as many as eight to as few as five.

The tarsus and propodus of these specimens show the greatest differences

from those of the type. The tarsus is fully 0,6 as long as the propodus and both

are more slender than those of the type. There are usually more sole spines, the

proximal heel spine is sometimes absent, and in the males from SM 250, the

auxiliary claws are only about 0,4 times as long as the main claw. In most other

characters, the variations are sufficiently small to suggest that these specimens

are well within the limits described for the type.

The male oviger has not been described before as the type, and only previous

record, is a female. The fifth segment is typically elongate, curved, laterally setose,

and bears a small distal apophysis with six or seven setae as long as the segment

diameter. The strigilis is also more setose and bears spines in the formula

8: 6:5: 7, with five teeth on the terminal claw. The distal palp segments are more

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

setose ventrally. In summary, the principal differences are the greater length of the

third palp segment and tarsus, and the shorter chelae in relation to the scape in

these specimens.

Distribution

The range of this species is extended eastwards from Lambert’s Bay in the

South Atlantic to the Indian Ocean, from off Lake St Lucia to the Transkei

coast, and the known depths are greatly extended from 23 m for the type to

90 m and 550 m by the present specimens.

Nymphon obesum sp. nov.

Fig 11

Material

East London area. Holotype, SAM—A10263, SM 179, 33°30,3’S 27°22,1'E,

80 m;, I 9.

Description

Moderately small, leg span 26,7 mm. General habitus robust, stout, append-

ages crowded. Trunk completely segmented, barrel-shaped. Lateral processes

short, thick, only as long as their diameters, separated by very slight intervals to

almost touching, glabrous. Neck short, oviger implantation against and below

first lateral processes. Ocular tubercle directly above ovigers, short, slightly

taller than wide, apex conical with small lateral papillae, eyes large, tear-shaped,

slightly pigmented. Proboscis broad, shorter than twice its diameter, cylindrical,

lips flat. Abdomen short, carried almost erect, tapering distally, glabrous.

Chelifores large, scape longer than proboscis, downcurved, armed with

several dorsal and distal setae. Chela palm longer than fingers, subcylindrical,

with few distal setae. Fingers well curved, overlapping at tips, armed with three

widely separated teeth on the immovable finger and four like teeth on the

movable finger.

Palps small, slender, second segment longest, third slightly shorter, both

armed with several setae, mostly distal. Fourth segment slightly shorter than

fifth, both with short ventral and distal setae not longer than segment diameter.

Oviger fourth and fifth segments subequal, glabrous, sixth shorter than

seventh, with several distal setae. Strigilis segments progressively shorter, armed

with several short ectal setae and denticulate spines in the formula 11 : 6: 5: 5;

with eight small blunt teeth on the short almost straight terminal claw. Spines

with three lobes per side.

Leg robust proximally, more slender distally. Main segments with few

setae, tibia 2 the longest segment, with tibia 1 and femur progressively shorter.

Tarsus very short, 0,33 the length of propodus, armed with ventrodistal spine

and few sole setae. Propodus slightly curved, armed with several short ectal

setae and five or six sole spines interspersed with short setae. Claw robust,

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 169

Fig. 11. Nymphon obesum sp. nov. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Palp. D. Chela. E. Oviger. F. Strigilis, with enlargement

of denticulate spine. G. Third leg, terminal segments.

short, about 0,3 the length of propodus, auxiliaries slender, about 0,6 as long as

main claw.

Male unknown.

Measurements (holotype, in mm)

Trunk length, 2,94; trunk width (across 2nd lateral processes), 2,04; pro-

boscis length, 1,32; abdomen, 0,66; third leg—coxa 1, 0,78; coxa 2, 0,88;

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

coxa 3, 0,76; femur, 2,4; tibia 1, 2,64; tibia 2, 2,92; tarsus, 0,4; propodus, 1,18;

claw, 0,37.

Remarks

This new species shares the character of a remarkably ‘fat’ trunk and pro-

boscis with a number of species previously named under the old junior synonym

Chaetonymphon, erected by G. O. Sars (1888) to include ‘thickset’ setose

species. Although we have found no closely related South African species at all

like N. obesum, there are at least two similar North Atlantic species, N. hirtum

Fabricius, 1794, and N. hirtipes Bell, 1853. The new species is probably closest

to N. hirtum from the high Arctic, except that it is far less setose and has a bald

appearance in comparison to N. hirtum. The chelae, legs and propodus are quite

different in the new species.

Nymphon hirtipes is also a very setose species and, in spite of its similarities

of short neck, thickset trunk and lateral processes, and large chelifores, its

ovigers, legs, chelae, and palps are very different from those of N. obesum.

Distribution

Only known from the type locality, off East London, at 80 m.

Etymology

The species name obesum refers to the robust and thickset habitus of this

unique specimen.

Nymphon paralobatum sp. nov.

ice 2

Material

Transkei area. Holotype, SAM-A19571, SM 226, 32°28,6'S 28°58,8’E,

T0=775 m, 12:

Description

Moderately small, leg span 24,4 mm. Trunk completely segmented, rather

slender, lateral processes short, little longer than their diameters, separated by

about their diameters, glabrous. Neck short, oviger bases slightly anterior to first

lateral processes; ocular tubercle between oviger bases and first lateral pro-

cesses, short, not as tall as wide, with tiny apical papilla; eyes large, slightly

pigmented; lateral papillae present, tiny. Proboscis moderately long, cylindrical,

with constriction just proximal to flat lips. Abdomen extending half length of

first coxae of fourth legs, tapering distally, armed with few dorsolateral setae.

Chelifores large, scape cylindrical, armed with few lateral and distal setae.

Chelae only as long as scapes, palm semicylindrical, armed with scattered short

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 171

Fig. 12. Nymphon paralobatum sp. nov. Holotype, female. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Palp. D.Chela, with enlargement of teeth. E. Oviger.

F. Strigilis, with enlargements of proximal and distal denticulate spines. G. Third leg,

distal segments.

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

setae. Fingers slender, slightly longer than palm. Movable finger moderately

curved, armed with 38 slender straight teeth; immovable finger almost straight,

overlapping movable finger at tip, armed with 33 slightly longer straight teeth.

Palps fairly small, second segment only slightly longer than third, both

armed with few distal setae. Terminal two segments subequal, together almost

as long as third segment, armed with many ventral and distal setae, some longer

than segment diameters.

Oviger fourth segment with small proximal bump, length about equal to

fifth segment. Sixth segment straight, 0,6 as long as fifth, both armed with row

of ectal setae and sixth with short lateral and ventral setae. Strigilis segments

armed with few short ectal setae and denticulate spines endally in the formula

8 :6:5: 6, with eight short teeth on terminal claw as long as tenth segment.

Denticulate spines shorter proximally than longer distal spines.

Legs moderately setose, major segments increasingly longer from femur to

tibia 2. Tarsus 0,56 as long as propodus, armed with six or seven sole spines,

several lateral and dorsal setae not as long as segment diameter. Propodus

armed with 10 or 11 sole spines, few lateral setae, and several dorsal setae,

some of which are longer than segment diameter. Claw about 0,45 as long as

propodus, well curved, auxiliaries about 0,7 as long as main claw.

Male unknown.

Measurements (holotype, in mm)

Trunk length, 2,58; trunk width (across 2nd lateral processes), 1,28; probos-

cis length, 1,1: abdomen, 0,6; third leg—coxa 1, 0,43; coxa 2, 0,76; coxa 3, 0,38;

femur, 1,75; tibia 1, 2,74; tibia 2, 3,84; tarsus, 0,46; propodus, 0,82; claw, 0,36.

Remarks

We hesitate to attribute this single female to a new species, particularly as it

has no outstanding or unique characters, but its combination of characters, we

feel, goes beyond the extremes of variation that might be found in its near-

namesake, N. lobatum Stock, 1962. It is also sufficiently different from any

other species in the genus to warrant being placed in a new species. This new

species is closest to N. Jobatum in its general trunk habitus, most chelifore

characters, the palp, some oviger characters, and the legs in particular. It differs

from N. lobatum in having a shorter neck, oviger bulges placed in advance of

the first lateral processes, longer chelifores in relation to the scapes (about

equal), a much shorter second coxa, and a slightly longer tarsus. Nymphon

lobatum was described from a single male specimen and this new species is a

single female, but the differences are mostly non-sexual and sufficiently numer-

ous, we believe, to warrant designating a new species. Another major difference,

although of little taxonomic importance, is the size of the two species. In most pyc-

nogonids the female is usually a bit larger than the male in most measurements

except the oviger, but this female specimen is smaller in the trunk measurements

and has about half the leg span of N. lobatum (latter about 51 mm).

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES M73

This new species also has characters in common with N. signatum Mobius,

1902, another presumably endemic South African species. In both the neck is

moderately short, the lateral processes are separated by about half their

diameters, the chela palm is approximately equal in length to the fingers that

overlap at the tips, the palps are similar with the fourth segment being about

0,85 the length of the third, and the tarsus, propodus and claw length ratios are

similar. The differences are that, in N. signatum, the oviger bases touch the first

lateral processes, the ocular tubercle is taller than wide, there are almost twice

as many teeth on the immovable finger as there are on the movable finger, and

there is a maximum of 43 denticulate spines of the same size on the strigilis seg-

ments, whereas in the new species there are only 23 spines, of which the distal

spines are taller.

Distribution

Only found at one station, off Transkei coast.

Etymology

The specific name is derived from the prefix para meaning a species beyond

the designated nomina Nymphon lobatum, its closest relation.

Nymphon pedunculatum sp. nov.

Fig. 13

Material

East London area. Holotype, SAM—A19581, SM 163, 33°04,6’S 28°06,6’E,

90 m, 1 d with eggs. Paratype, USNM 228134, SM 163, 90 m, 1 6d. Paratypes,

SAM-A19582, SM 163, 90 m, 2 2°. Paratypes, MNHN-Py 598, SM 164, 1 &.

Description

Male. Size small, leg span about 21 mm. Trunk fully segmented, lateral pro-

cesses 1,5 as long as their diameters, separated by half to less than half their

diameters, glabrous. Neck very short, only as long as oviger bases. Ocular

tubercle situated directly dorsal to oviger bases, slightly over twice as tall as

diameter, arising from constricted base giving pedunculate appearance, eyes

large, well pigmented, situated just below biconical apex. Proboscis slightly

longer than cephalic segment, cylindrical, with slight constriction distal to mid-

point, lips moderately inflated. Abdomen short, erect, extending just beyond

distal rims of fourth lateral processes, glabrous.

Chelifores robust, scape almost as long as proboscis, armed with few lateral

and distal setae. Chelae approximately as long as scapes, palms little shorter

than fingers. Immovable finger only curved distally, overlapping movable finger,

armed with fringe of proximal setae, 53 closely set teeth of equal size. Movable

finger moderately curved except distally where sharply curved, armed with

56 closely set teeth of equal size, slightly smaller than teeth of immovable finger.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

WM,

Sy YY)

LUN

ALA LS

Fig. 13. Nymphon pedunculatum sp. nov. A-G. Holotype, male. A. Trunk, dorsal view.

B. Trunk, lateral view. C. Palp. D. Chela, with enlargement of teeth. E. Third leg, distal

segments. F. Oviger. G. Strigilis. H. Paratype, female. Strigilis, with enlargements of

denticulate spine and terminal claw.

Palp segment three subequal to segment two, third armed with few distal

setae. Segment five only 0,8 as long as segment four, combined length of both

equal to second segment, terminal segments armed with many short lateral and

ventral setae, some longer than segment diameter.

Oviger segments four and five subequal, both with row of lateral setae,

short on fourth, longer on fifth. Sixth segment slightly curved, 0,3 as long as

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES leis)

fifth, armed with lateral setae and tuft of long distal setae, longer than segment

diameter. Strigilis segments progressively shorter in length, armed with ectal tuft

of setae longer than segment diameter, and endal denticulate spines in the

formula 12 : 8 : 7 : 7; spines slender, with four or five lateral serrations per side.

Terminal claw slightly shorter than terminal segment, straight, armed with

17 lobe-like serrations extending to tip.

Major leg segments with few setae, tibia 2 longest with tibia 1 and femur

each respectively shorter. Cement-gland pores not evident. Tarsus about 0,85 as

long as propodus, both armed with few dorsal setae, numerous short lateral

setae and many short sole spines of a single size. Claw 0,3 length of propodus,

moderately curved, auxiliaries about 0,6 as long as main claw.

Female. Slightly larger than male except for oviger. Bifurcate ocular

tubercle apex also pedunculate, as is base of tubercle. Oviger segments four and

five subequal, strigilis segments with fewer setae and all shorter than those of

male. Strigilis compound spines in the formula 13 : 10 : 8 : 9, with 18 blunt ser-

rations on terminal claw.

Measurements (holotype, in mm)

Trunk length, 1,99; trunk width (across 2nd lateral processes), 1,16; pro-

boscis length, 1,18; abdomen, 0,4; third leg—coxa 1, 0,41; coxa 2, 0,72; coxa 3,

0,58; femur, 1,91; tibia 1, 2,39; tibia 2, 2,64; tarsus, 0,49; propodus, 0,59;

claw, 0,19.

Remarks

This new species is closest to Nymphon pleodon Stock, 1962, a South

African species not encountered during this survey. It differs from N. pleodon in

having a taller and pedunculate ocular tubercle, chelae with fingers showing less

curvature and having more closely crowded teeth, the palp terminal segment

shorter than the fourth, oviger segment six slightly longer than the seventh, ter-

minal claw and denticulate spines of the oviger with a greater number of

serrations, femur shorter than tibia 1, and auxiliaries longer than half the main

claw length. The new species lacks the strong broad spines on oviger segments

five and six of N. pleodon.

As stated by Stock (1962: 281), N. pleodon resembles N. gruveli Bouvier,

1910, as does the new species. The principal differences between the new species

and N. gruveli are that the latter lacks the pedunculate ocular tubercle, has

dorsodistal tubercles on tibia 2 (not found in the new species), and has much

shorter auxiliary claws. Also, in Bouvier’s species the terminal palp segment is

longer than the fourth segment, whereas the opposite is true for the new species.

Distribution

Only found at one station, off East London.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The specific name pedunculatum is derived from the Latin pedis, meaning a

foot and refers to the ocular tubercle growing on a narrowed basal part that sup-

ports the larger part of the tubercle.

Nymphon phasmatodes Bohm, 1879

Nymphon phasmatodes Bohm, 1879: 173-174, pl. 1 (fig. 2). Flynn, 1928: 4. Loman, 1928: 67,

figs 2, 6, 7. Stock, 1956: 76-78, fig. 2; 1959: 558; 1962: 283-284; 1965: 22-23 [key].

Nymphon capense Hodgson, 1908: 169-170, pl. 1 (figs 2, 2A). Gordon, 1932a: 117-120,

figs 11-12; 1932b: 28, 32, 34, 62. Barnard, 1954: 91-92.

Material

East London area. SAM-—A19605, SM 180, 80 m, 1 2. SAM—-A19606,

SM 185, 90 m, 1 9, 1 juv.

Remarks

There are few differences between these three specimens and the previous

decriptions and illustrations of N. phasmatodes. We regard these differences to

be only variation within the species, because most characters in the Meiring

Naude specimens agree very well with the published figures. The differences are

a slightly shorter tarsus for these specimens, the strigilis spine and claw count

which vary by one or two spines per segment, and a similar variation in claw

teeth, a slightly lower ocular tubercle having rather large lateral papillae, and a

shorter tibia 2 which is nevertheless much longer than tibia 1 or the femur.

Apparently, the chelifores of this species are carried usually in a distinctive erect

position with the chelae extended vertically down to the sides of the mouth. In

most Nymphon species the chelae are carried in a more horizontal or oblique

position.

Distribution

This is another endemic South African species known from relatively

shallow depths. Station 185, at 90 m, marks the deepest depth from which this

species has been taken.

Nymphon pilosum Mobius, 1902

Nymphon pilosum Mobius, 1902: 179, pl. 24 (fig 8-12). Barnard, 1954: 100-101, fig. 6. Stock,

1956: 76; 1962: 278; 1968: 29.

Nymphon bipunctatum Flynn, 1928: 8-10, figs 1-2.

Material

Zululand area. SAM—A19591, SM 86, 550 m, 6 66 with eggs, 2 dd, 3 29,

5 juvs. MNHN-Py 595, SM 86, 550m, 1 6, 1 9. SAM—A19593, SM 103,

680 m, 2 dd with eggs, 1 d, 4 gravid 29, 3 juvs. SAM—A19592, SM 129,

850 m, 1 6.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES a

Transkei area. SAM—A19590, SM 226, 710-775 m, 1 2. SAM-A19589,

SM 250, 150-200 m, 1 6, 2 9°.

Remarks

This fairly common species is easily recognized among the many South

African Nymphon species by its extremely long leg setae, some of which are up

to five times longer than the segment diameter. As is true for all described

Nymphon species examined for this report, the appendage measurements of this

species vary and assume slightly different ratios among the segments, the oviger

strigilis spines vary in number—some with as little as 18 spines, and the number

of chelae finger teeth show a reduction in number in some of the above speci-

mens, possibly due to age, but the specimens agree very well in most characters

with the specimens previously figured. The cement-gland tube cones on the

ventral surface of the femur vary in number from one male with two per leg to

several specimens with four or five per femur. This, again, may be a function of

age.

Distribution

As far as its distribution is known, this species is confined to South African

waters, and was previously recorded over a rather wide depth range from 20 m

to 366 m. The depths at which it was taken by the Meiring Naude increase the

maximum known depth to 850 m.

Nymphon serratidentatum sp. nov.

Fig. 14

Material

Transkei area. Holotype, SAM-—A19583, SM 226, 32°28,6’S 28°58,8'E,

710-775 m, 1 subadult 2. Paratypes, SAM—A19584, SM 226, 710-775 m, 1 6

(damaged), 1 juv.

Description

Female. Size moderately small, leg span 27,8 mm. General habitus slender,

tenuous. Trunk long, completely segmented, with lateral processes separated by

twice their diameters, less than twice as long as their diameters, glabrous. Neck

moderately long, triangular in dorsal profile anterior to ocular tubercle and

Oviger implantation, which are well anterior to first lateral processes in an

inflation of the neck. Ocular tubercle low, rounded, wider than tall, eyes

present, with little pigmentation. Proboscis slender, moderately short, with

slight median inflation, lips rounded. Abdomen very short, not extending

beyond distal fourth lateral processes, with few dorsodistal setae.

Chelifores large, scape 1,2 times longer than proboscis, glabrous. Chelae

slightly longer than scapes, curved inward, palm less than half length of fingers,

with few short distal setae. Fingers very slender, slightly curved, overlapping at

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. Nymphon serratidentatum sp. nov. Holotype, female. A. Trunk, dorsal

view. B. Trunk, lateral view. C. Chela, with enlargements of teeth. D. Third

leg, terminal segments. E. Oviger. F. Strigilis, with enlargements of denticulate

spines.

tips, armed with very many closely crowded narrow teeth, numbering 95 on

immovable finger, 118 on movable finger. Some proximal teeth on movable

finger with finely serrate edges with two to four tiny serrations per margin. No

serrate teeth on immovable finger, plain spine-like teeth on both fingers distally.

Finger tips slightly curled laterally.

Palps long, tenuous, third segment longest, 1,3 times longer than second,

fourth almost as long as third, fifth segment shorter, half as long as third, with

slight endal curve. Third to fifth segments armed with few very short ventral and

distal setae.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 179

Ovigers broken off and missing beyond second and fourth segments,

respectively.

Legs long, slender, tibia 2 longest with first tibia 0,6 as long and femur 0,43

as long, armed with few short setae and long dorsodistal seta over three times

longer than segment diameter of each segment. Tarsus slightly over 0,4 as long

as propodus, both armed with few short dorsal and lateral setae, six equal sole

spines on tarsus and 16 or 17 equal sole spines on propodus, none longer than

segment diameter. Claw about 0,3 as long as propodus (claw tip broken off),

auxiliary claws lacking.

Male. Slightly smaller in all measurements except oviger segments. Fifth

Oviger segment very long, slender, 1,5 times length of fourth, slightly swollen

distally, both armed with few short lateral setae. Sixth segment about half as

long as fourth, with strong proximal curve separated from distal straight section

by small lateral tubercle, armed with lateral and distal setae. Strigilis segments

progressively shorter in length, slender, armed with few ectal setae not as long

as segment diameter and endal denticulate spines in single row in the formula

13: 8:7: 8, with slender terminal claw, slightly curved, not as long as terminal

segment and armed with 13 short endal teeth. Denticulate spines shorter on

proximal part of segments, longer on distal parts, with three slender serrations

per side.

Palp segment lengths differ from female with fourth segment slightly longer

than second, and third 0,87 as long as fourth. Terminal segment, presumably

curved, missing.

Measurements (holotype, in mm)

Trunk length, 4,61; trunk width (across 2nd lateral processes), 1,62; probos-

cis length, 1,62; abdomen, 0,52; third leg—coxa 1, 0,38; coxa 2, 0,99; coxa 3,

0,42; femur, 2,11; tibia 1, 3,01; tibia 2, 4,99; tarsus, 0,38; propodus, 0,84; claw,

0,3 (? broken off).

Remarks

This new species is very like Nymphon microctenatus Barnard, 1946, in

many characters. For purposes of comparison, the type of Barnard’s species was

examined and found, among other differences, to be about three times larger

than these specimens. The major differences are in the chelae teeth, which

number about 213 in the new species and over 300 in Barnard’s species, the

much longer neck, the anterior placement of the oviger bases and ocular

tubercle in the new species, the much less stout trunk and lateral processes that

are separated by much greater distances than in Barnard’s species, the strongly

curved chelae finger tips which overlap in the new species, and the tarsus which

is much shorter than the propodus in the new species.

The striking similarities between the two species are the great number of

chelae finger teeth, the extreme similarity of the palp segments in length, and

the curve of the fifth or terminal segment, the separation of the oviger bases

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

from the first lateral processes in both, and the relative measurements of leg,

oviger and chelifore segments.

Unfortunately, the holotype female of the new species has only one com-

plete leg and the adult male paratype is entirely without legs. We believe the

differences between these specimens and Barnard’s type of N. microctenatus go

beyond variation within a species. However, if subsequent specimens are taken

that are intermediate between these specimens and Barnard’s type then, of

course, the present specimens must be included in his species.

There are some similarities in the new species with N. prolatum Fage, 1942,

a slender species found in West Africa. The general habitus of both species is

similar, except for the ocular tubercle, abdomen and lateral process, which are

very much longer than those of N. serratidentatum.

The presence of several serrate teeth among the otherwise simple curved

teeth on the movable finger of these specimens is, so far we can discern from the

literature, unique among the many species of Nymphon. The chelae teeth of

N. microctenatus are very difficult to see separately in their crowded state, but

none of the proximal teeth has any serrations in the type specimen.

Distribution

Species only found at one station off the Transkei coast at 710-775 m.

Etymology

The species name serratidentatum refers to the uniquely serrate chelae

finger teeth of this new species.

Nymphon sp. indet.

Material

East London area. SAM—A19586, SM 185, 90 m, 1 juv.

Transkei area. SAM—A19587, SM 250, 150-200 m, 1 juv.

Remarks

Neither of these juveniles is sufficiently developed to describe or compare

with other Nymphon species from the above two stations.

DISTRIBUTION AND ZOOGEOGRAPHY

The South African Museum’s Meiring Naude cruises comprise a set of col-

lections made between the southern Mozambique border and just south of East

London, South Africa, and represent the only modern survey of deeper waters

along this coast. Of the 92 benthic stations made during the annual cruises from

1975 through 1979, samples from only 21 (21,7%) of these stations contained

pycnogonids. No pycnogonids were taken during the first cruise, but methods

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 181

improved subsequently and greater numbers appeared in the samples as the

cruises advanced. The number of specimens (455, of which 253 were of one

species) and the small number of stations do not permit us to draw reliable con-

clusions concerning distribution.

A few brief comments can be made about the Meiring Naude collections

with regard to the families represented, depth distribution, geographical bound-

aries, and associated fauna. The majority of pycnogonid species recorded in

these collections belong in three families, while other families are represented

by very few or even no specimens. This situation is different to a deeper-water

collection reported on by Stock (1963) from South Africa, where the predomi-

nant family was the Colossendeidae with six species. Stock’s (1981) deep Walvis

Basin records again emphasized the Colossendeidae (3 species) and two other

families also prevalent in the Meiring Naude material—Nymphonidae

(4 species) and Austrodecidae (2 new species). The Nymphonidae dominate the

Meiring Naude material with 14 species (7 new to science), and most of them

are from the continental slope. Half the species of this family reported on herein

occur in shelf depths of less than 100 m (with some distributions extending well

beyond 100 m), whereas the remaining seven species are found between 100 m

and 850 m; no specimens were taken between 850 m and 1 300 m.

Another very well-represented group in this material is the family Calli-

pallendidae, with 10 species (3 new). It is also represented by the most genera

(six), including the new genus Safropallene.

The family Ammotheidae is represented by five known species with one of

them, Boehmia tuberosa, a relatively rare genus and species. A surprise is the

appearance of new material of Ammothella setacea, taken only for the second

time and redescribed here due to the loss of the type material. It is the most

common pycnogonid taken in this survey, with captures at nine stations in conti-

nental shelf and slope depths of 90 m to 900 m. Indeed, the 253 specimens of

A. setacea comprise over 55 per cent of the entire collection.

The family Pycnogonidae is represented here by four species, all from

bathyal depths, but Pycnogonum forte and P. nodulosum are also known from

shallower depths along the South African coast or in other parts of the Atlantic

and Mediterranean.

There were only two Meiring Naude stations as deep as 1 000 m to 1 300 m,

so the fact that no species of the genus Colossendeis (family Colossendeidae)

was captured is inconclusive. There is evidence (Stock 1963) that these species

live in deeper habitats off the South African coast. This family is represented in

the collections by smaller-sized genera: Rhopalorhynchus gracillimus in rela-

tively shallow depths, and Hedgpethia magnirostris in bathyal depths.

The genus Pantopipetta (family Austrodecidae) is represented by two new

species, adding to the six species already known from southern African waters

(including two from the Walvis Basin). Of the twelve known species, South

Africa seems to be a point of speciation for this genus, although not all six South

African species are endemic. The genus is a predominantly deep-water one, but

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

at least two species (one South African) have been taken in depths as shallow as

66-69 m.

The family Phoxichilidiidae, a predominantly warm, shallow-water group, is

represented here by only one species, Anoplodactylus typhlops, a true deep-

water blind form. The genera Endeis and Phoxichilidium were not recorded at

the depths sampled by the Meiring Naude, but at least two species of Endeis are

known from shallower waters of the South African coast— E. clipeatus (MObius,

1902) and E. mollis (Carpenter, 1894) (Barnard 1954: 129-132).

Tiny specimens of the genus Rhynchothorax (family Rhynchothoracidae)

have yet to be found in South African waters.

We can distinguish three groups of species in the Meiring Naude material,

according to their depth range. There is a continental shelf and slope component

(c. 90 m to 200 m) composed of Ascorhynchus ornatum, Tanystylum thermo-

philum, Rhopalorhynchus gracillimus, Nymphon barnardi, N. crenatiunguis,

N. obesum, N. pedunculatum and N. phasmatodes. A bathyal element, rep-

resented by four species, occurring below 500 m, includes Nymphon granu-

latum, N. microctenatus, N. pilosum and N. serratidentatum. A group of euryba-

thic species, which may be found at shallow as well as bathyal depths, includes

Ammothella setacea, Cilunculus sewelli, Pycnogonum nodulosum, Nymphon dis-

tensum and N. modestum.

The major trends in geographical distribution of southern African littoral

pycnogonids are sufficiently documented in the previous literature (M6bius

1902; Barnard 1954; Stock 1959, 1963), but the Meiring Naude material,

obtained mostly from depths below 500 m, improves our knowledge of the

deeper-water species. There are now approximately 100 species recognized from

the entire area of southern Africa, south of 20°S (Arnaud & Bamber 1987: 71,

table 14). Among the deep-water species of the Meiring Naude material, an

Atlantic component of five species may be recognized: Pallenopsis longirostris,

Pycnogonum crassirostre, P. nodulosum, Callipallene phantoma, and Anoplo-

dactylus typhlops. There is also an Indo-Pacific element of four species: Cilun-

culus sewelli, Pseudopallene gilchristi, Pycnogonum africanum and Rhopalo-

rhynchus gracillimus. Pycnogonum forte, N. crenatiunguis and N. microctenatus

are among the large endemic element of southern Africa. Other endemics are:

Ammothella setacea, Ascorhynchus ornatum, Boehmia tuberosa, Tanystylum

thermophilum, Pallenopsis capensis, Parapallene algoae, Nymphon distensum,

N. modestum, N. phasmatodes, N. pilosum and N. signatum.

There is a true deep-water component of apparently blind species, all of

which are described as new in this report. These are: Nymphon bicornum,

N. granulatum, N. serratidentatum, N. paralobatum, Pantopipetta armata,

Hedgpethia magnirostris and Safropallene longimana.

There is an absence of any Antarctic component, even at bathyal depths of

1 000 m to 1 300 m, in the Meiring Naude material. Several boundaries between

the different parts of the very long coasts of southern Africa have been recog-

nized (Branch & Branch 1981). The cold Benguela Current delimits a cold-

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 183

temperate zone on the west coast, from Cape Town at 33°56’S north to 20°S lati-

tude. A warm-temperate zone occurs from Cape Point to East London (33°S

27°54'E), a subtropical east-coast zone north from East London to Inhambane

(23°51'S 30°30’E), and a tropical east-coast zone from north of Inhambane to

Beira (20°S 35°E). With the Meiring Naude specimens, the Atlantic component

corresponds with the cold-temperate zone, the endemic component mostly

agrees with the warm-temperate zone of the south coast, and the Indo-Pacific

component corresponds with the subtropical east-coast zone.

Comparisons of the Meiring Naude material with that of other abyssal

surveys along the South African coasts is, unfortunately, mostly inconclusive.

The six abyssal stations studied by Stock (1963) between 2 700 m and 3 000 m

reveal a very different type of pycnogonid fauna in these deeper waters of the

west coast of South Africa (off Cape Town and the Cape Peninsula). He found

12 species (six of which were new) belonging to five genera. These were one

Nymphon, one Ascorhynchus, six Colossendeis, one Anoplodactylus and three

species of Pantopipetta. This fauna may not be considered as truly South

African, but rather as a partly cosmopolitan fauna (Atlantic and Indo-Pacific)

for the six Colossendeis species, and a partly deep-sea Atlantic fauna. This deep-

water Atlantic and Cosmopolitan composition holds for Stock’s (1981) paper on

the pycnogonids from the Walvis (or Cape) Basin in 3 350 m to 5 040 m. There

are no affinities with the Meiring Naude fauna at these depths and localities, but

it does reveal that this Walvis Basin fauna is also partly abyssal cosmopolitan

and partly of a deep-water Atlantic component.

Comparison of the Meiring Naude pycnogonid stations with stations

recorded as having possible prey material in the same catch is somewhat more

revealing. It is known that pynogonids prey on hydroids and there is some evi-

dence that they may also feed on the soft parts of bryozoans (Arnaud & Bamber

1987: 45). We note that at almost all pycnogonid capture stations, hydroids

(Millard 1977, 1980) and/or bryozoans (Hayward & Cook 1983) were among the

associated fauna brought up at each station.

On the basis of hydroid faunas, Millard (1978) was able to recognize at least

nine biogeographic elements but, with the relatively small number of stations

containing pycnogonids, we cannot provide such conclusive distribution data,

particularly because of the high number (32%) of new species and other rare or

little known species.

An attempt was made to see whether or not pycnogonids show a tendency

towards greater depths in warmer waters, as Millard (1978: 179) did for

hydroids. For example, Tanystylum thermophilum was dredged on the west

coast at 50-54 m (33°08,6’S 17°57,3’E, sta. WCD203Q, 29 April 1964, unpub-

lished record), and at 17 m and 26-29 m in False Bay on the south coast, but at

150—200 m on the east coast (Transkei, Meiring Naude, SM 250). Pycnogonum

nodulosum has been taken in only 20m at Saldanha Bay on the west coast

(sta. SB306T, 33°03,7'S 17°58,5’E, 30 April 1963, unpublished record), at 9 m

on the south coast, but at 150—200 m on the Transkei and 500 m on the Zulu-

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

land coasts (Meiring Naude, SM 250 and SM 86). Another species, Nymphon

phasmatodes, is known from 48 m, 53 m and 79 m respectively on the west coast

(sta. WCD184, 186 and 189, 33°07,6’E, unpublished records), at 40 m, 61 m and

66 m on the south coast (False Bay, sta. FAL601, 687 and 688, unpublished

records), and at 80-90 m on the east coast (south-west of East London).

Nymphon comes occurs in more shallow waters in False Bay than on the Trans-

kei coast (Meiring Naude, SM 233, 540-580 m).

From these records it is apparent that at least the above listed species do

have a tendency to occupy greater depths on the east coast than on the west

coast, but the phenomenon does not hold true for other species where greater

geographic distribution is known.

Considering the bryozoan fauna, Hayward & Cook (1983: 145, 147)

stressed ‘the accepted faunal similarity of eastern South Africa with the Indo-

Pacific realm’ and the ‘increasing evidence of strong similarities with eastern

Australian and northern New Zealand faunas’. These assumptions do not hold

true for the pycnogonid fauna, as no known Meiring Naude pycnogonid species

appears to have a common distribution with India or the Australian area (nor

even with South America).

The same authors gave an analysis of sediments in relation to the number of

living and dead bryozoan species identified from eleven stations. Pycnogonids

were taken at nine of these stations and it is possible that they contributed to the

number of dead colonies by predation. Hayward & Cook (1983) pointed out

that there was a significant contribution to the sediment made by bryzoan skel-

etal remains at several stations (SM 129, SM 131, SM 151, SM 163/164 and

SM 179). At station SM 163/164 (90 m) they found 43 living and 24 dead spe-

cies, at SM 179 (80 m) 20 living and 4 dead species, SM 180 (80 m) 6 living and

21 dead species, and at SM 185 (90 m) 40 living and 15 dead species.

Both Millard (1978) and Hayward & Cook (1983) emphasized the high level

of endemism in the region between Cape Agulhas and Durban. Considering the

distribution of the Meiring Naude species, this endemism extends to the pycno-

gonid fauna, which in this report includes 14 endemic species from this area.

The 13 new species described here are not included in this count but some may

be found to be endemic when more is known of their distribution.

ACKNOWLEDGEMENTS

We wish to thank the Director and Trustees of the South African Museum

for permission to work on the Meiring Naude collections presented here, and for

the loan of important types and other rare material for comparative purposes.

Our sincerest appreciation is extended to Miss Elizabeth Louw of the South

African Museum for her assistance in providing data, station lists, and encour-

agement during our work on this report, and for her excellent editing of the

manuscript for publication.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 185

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BARNARD, K. H. 1955. Additions to the fauna-list of South African Crustacea and Pycno-

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HeEpDGPETH, J. W. 1948. The Pycnogonida of the western North Atlantic and the Caribbean.

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186 ANNALS OF THE SOUTH AFRICAN MUSEUM

HELFER, H. 1938. Einige neue Pantopoden aus der Sammlung des Zoologischen Museums in

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KRAPP-SCHICKEL, G. & Krapp, F. 1975. Quelques traits de lecologie d’amphipodes et de

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Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data

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Louw, E. 1980. The South African Museum’s Meiring Naude cruises. Part 10. Station data

1977, 1978, 1979. Annals of the South African Museum 81 (5): 187-205.

MitiarpD, N. A. H. 1977. The South African Museum’s Meiring Naude cruises. Part 3.

Hydroida. Annals of the South African Museum 73 (5): 105-131.

MiLiarp, N. A. H. 1978. The geographical distribution of southern African hydroids. Annals

of the South African Museum 74 (6): 159-200.

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Hydroida. Annals of the South African Museum 82 (4): 129-153.

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(Zool. 20): 1-163.

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Stock, J. H. 1952. Revision of the European representatives of the genus Callipallene Flynn,

1929 (Pycnogonida). Beaufortia, Amsterdam 13: 1-14.

Stock, J. H. 1953a. Re-description of some of Helfer’s pycnogonid type-specimens. Beaufor-

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knowledge of the pycnogonid fauna of the East Indian Archipelago. Temminckia 9:

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from Indo-West-Pacific, Australian and New Zealand waters. Videnskabelige Meddelelser

fra Dansk Naturhistorisk Forening i K¢@benhavn 116: 1-168.

Stock, J. H. 1956. Tropical and subtropical Pycnogonida, chiefly from South Africa.

Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i K@benhavn 118: 71-113.

Stock, J. H. 1957. Pantopoden aus dem Zoologischen Museum Hamburg. 2. Mitteilungen

aus dem Zoologischen Museum in Hamburg 55: 81-106.

Stock, J. H. 1958a. The Pycnogonida of the Erythrean and of the Mediterranean coasts of

Israel. Contributions to the knowledge of the Red Sea, 5. Bulletin of the Sea Fisheries

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THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 187

Stock, J. H. 1959. On some South African Pycnogonida of the University of Cape Town

Ecological Survey. Transactions of the Royal Society of South Africa 35 (5): 549-567.

Stock, J. H. 1962. Second list of Pycnogonida of the University of Cape Town Ecological

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Stock, J. H. 1963. South African deep-sea Pycnogonida, with descriptions of five new

species. Annals of the South African Museum 46 (12): 321-340.

Stock, J. H. 1965. Pycnogonida from the southwestern Indian Ocean. Beaufortia 13 (151):

13-33.

Stock, J. H. 1966. Campagne de la Calypso au large des cétes atlantiques de l! Amerique du

sud (1961-62). 4. Pycnogonida. Résultats scientifiques de la campagne de la Calypso, 7.

Annales de I’ Institut Océanographique, Paris 44: 385-406.

Stock, J. H. 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and

Pacific oceans. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kgben-

havn 131: 7-65.

Stock, J. H. 1975. Pycnogonida from the continental shelf, slope, and deep sea of the tropical

Atlantic and East Pacific. Biological results of the University of Miami deep-sea

expeditions, 108. Bulletin of Marine Science 24 (4): 957-1092.

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de Biologie Marine 22 (4): 453-471.

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Witson, E. B. 1881. Report on the Pycnogonida. Reports on the results of dredging .. .

Blake. Bulletin of the Museum of Comparative Zoology, Harvard 8 (12): 239-256.

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

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 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.

FRANCOISE ARNAUD & C. ALLAN CHILD

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES

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