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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 91 BAND 91

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 91 BAND

rE iNUSPEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1983

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CLuVER, M. A. & KING, G. M.

A reassessment of the relationships of Permian Dicynodontia (Reptilia, Therapsida)

and a new classification of dicynodonts. (Published March 1983.).............. (25

Cook, P. L. see HAywarp, P. J.

Hai, M. & Mack, K.

The outline of an eighteenth-century economic system in south-east Africa. (Pub-

hiSie GalaniiaryehO SOs) esse cr) PR yh oe ccs < eie, senh ey Re te CME RT A SNS ts 163

HaAyYwarp, P. J. & Cook, P. L.

The South African Museum’s Meiring Naude cruises. Part 13. Bryozoa II. (Published

INICIO SS re eaters he aa ey eae a dt 4 nrc Ra Eg NO CR er Gh 1

Kinc, G. M. see CLuver, M. A.

Mack, K. see HALL, M.

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Dacwlosicpanday wands COOK; 1983 «2... 2. cea e wecc dae Gee ede we es eae es

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ee! yo

2 a

ey:

Y :

=E ‘

= : :

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’

_ VOLUME 91 PART 1 MARCH 1983 ISSN 0303-2515

S67X

OF THE SOUTH AFRICAN

= MUSEUM

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

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FIsCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

Zool. exp. gén. 74: 627-634.

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

Ann. Mag. nat. Hist. (13) 2: 309-320.

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

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

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 91 _ Band

March 1983 Maart

Part 1 Deel

TARNSON

b 8.8.8:

yas »

OVI

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES

PART 13

BRYOZOA If

P. J. HAYWARD

Pe kb. COOK

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

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ISBN 0 86813 044 3

Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES

PART 13

BRYOZOA II

P. J. HAYWARD

Department of Zoology, University College of Swansea

pe les (CooK

Department of Zoology, British Museum, Natural History

(With 33 figures, 5 tables and 2 appendices)

[MS accepted 8 June 1982|

ABSTRACT

The bryozoan benthos of the shallow continental shelf seas off the eastern South African

coast is rich in species, and taxonomically diverse. Samples from 18 stations established by the

R.V. Meiring Naude between 1977 and 1979 have yielded a total of 38 anascan Cheilostomata,

76 ascophoran Cheilostomata and 16 Cyclostomata. The majority of the species were collected

in depths of less than 100 m. Forty-four new species are described in the following genera:

Amphiblestrum, Copidozoum, Chaperia, Arachnopusia, Micropora, Macropora, Aspidostoma,

Bugulella, Beania, Cribrilaria, Smittina, Smittoidea, Parasmittina, Escharella, Mucropetraliella,

Cleidochasma, Hippoporella, Emballotheca, Fenestrulina, Gigantopora, Adeonella, Sertella,

Iodictyum, Rhynchozoon, Brodiella, Turbicellepora, Hornera. Additionally, Dactylostega gen.

nov. is introduced for several species within the Hiantoporidae.

The South African bryozoan fauna includes a high proportion of endemic species, but also

exhibits a marked faunal affinity with the Indo-West-Pacific region. The antiquity of this link is

suggested by a number of similarities with the Tertiary fossil faunas of Australia and New

Zealand.

CONTENTS

PAGE

NER OCU CHO Mery ee oraci.c- A ens ua eA ae etek BAe ea 2

IETS (ROMS TNE CLES te ter, 5 2d (Oe Bae ey «eT eee ny cotied « Aen a ONS 3)

Systemiaticnccount aes «acct ee tahoe ey al ee citi we ern at 6

OrdermCheilostomatay | os6ace aise soe ob) ates sore Hemme: 6

OrdenmCyclostomatay var, ne. toe eee ye oe ete hue 128

DISCUSSION x cte tw ee ards LOO Clg SO A CN MOA ae Ay h See athe aie 140

SUITAUTTTEN Sg Ree ce hast aan ee aC eee Rea EMU da cfg eed NR OO rT an es 148

PNCKNOWICUSCINEM ES ee a retinscens ec eee GL Rete e ens 149

INGORE CC Sey aN ernst Reel ala Cinna eI A eal cn bbls als Ameer: 149

PNW) TENA OM S Haare eee tay AG sh ai) frase ep eee iene eae Shaye See ene een 155)

Appendix 1. Meiring Naude stations that produced Bryozoa ....... 156

Appendix 2. Indexiof speciesand) Seneral <5. as 4. - 465s esses eee 156

Ann. S. Afr. Mus. 91 (1), 1983: 1-161, 33 figs, 5 tables, 2 appendices

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

The first report on the Bryozoa collected by the South African Museum’s

Meiring Naude cruises (Hayward & Cook 1979) described a fauna of 51

species from a series of stations on the outer continental shelf and the

continental slope of eastern South Africa. These stations ranged in depth from

376 to 1300 m, with most of them being deeper than 500 m. A preponderance

of the species was shown to comprise specialized forms adapted to life on

fine-grained, unstable sediments, although a number of species were more

characteristic of hard-ground benthos and were considered to have been

collected at the lower limits of their bathymetric ranges. A remarkable pro-

portion of the fauna, 23 species, was new to science and, of the remainder, 14

were new records for South Africa. The present report describes the bryozoan

material collected during the Meiring Naude cruises of 1977, 1978, and 1979,

together with an additional sample from the 1976 cruise, and provides a useful

supplement to the first survey in supplying data for the inner shelf areas of the

same region. Thus, of the 18 samples studied here, 8 were collected in depths

of less than 100 m, 1 was collected at 150-200 m, and 6 at less than 700 m.

The samples contained abundant bryozoan material and, in contrast to the

first collection, which included many widely distributed deep-sea species, were

expected to provide a clearer picture of the indigenous South African fauna.

These expectations were exceeded when study of the collections revealed 130

species, including an astonishing total of 44 new species, many of which seem

to be endemic to this area of the south-western Indian Ocean. It is now

apparent that the bryozoan fauna of South Africa comprises a rich and

taxonomically diverse assemblage of species, and that the few previous studies

on this region (see Day et al. 1970) have failed to reveal more than a fraction

of its potential complexity.

The bottom sediments at a number of the stations studied here included a

substantial fraction of bryozoan remains. These fragments included numerous

unrecognized species of Cheilostomata and Cyclostomata that could be neither

identified with any of the species described here nor adequately characterized

from the material available. This report, therefore, cannot be regarded as a

complete survey of the shallow component of the South African bryozoan

benthos. Further collections will, without doubt, produce a yet greater range of

undescribed or poorly known species and genera. Many of the previously

described species reported here for the first time from South African waters are

known to be widely distributed in the Indo-West-Pacific region. Several of the

new species have systematic affinities with Tertiary fossil forms of Australia and

New Zealand. The potential of further research in this region for marine

zoogeographical theory seems exciting.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 8

LIST OF SPECIES

The present collections comprised 130 species of Bryozoa: 38 anascan

Cheilostomata, 76 ascophoran Cheilostomata and 16 Cyclostomata. Cteno-

stomata were not found. In Tables 1-3 the species are listed in systematic order

and their occurrence at each of the eighteen stations is given. The stations are

arranged in order of increasing depth and the notation distinguishes between

live and dead records. Co-ordinates and depths for each of the stations studied

are given in Appendix 1. Data for all stations have been published by Louw

(1977, 1980).

TABLE 1

Anascan Cheilostomata. The species are listed in systematic order, stations in order of

increasing depth. New species are indicated by an asterisk (*). X =represented by living

colonies. + = dead colonies only.

100 m 500m 700m

Depth station 179 180 184 163 163/ 164 185 230) 2504234 233 232 162 103 1234131 129 151

164

Carbasea mediocris Kor aX x

Cupuladria multispinata vo oD it tT

Discoporella umbellata Xia On Gt; t T + t

Setosellina roulei + t +

Heliodoma implicata <<a

Callopora sp. t t sf: aay

Amphiblestrum inermis Ke =x t

*Amphiblestrum pontifex x

*Copidozoum transversum

Crassimarginatella marginalis ate sak Xx

Foveolaria imbricata x + + ? + x + + +

Foveolaria sp. +

Chaperia multifida Sukie Wie Hoch mat x

Chaperia capensis x Ft

Chaperia stephensoni t

*Chaperia familiaris

Chaperia sp.

Notocoryne cervicornis

*Dactylostega prima x

*Arachnopusia corniculata

*Micropora similis it

Steginoporella buskii

*Macropora africana x

Cellaria tectiformis x x + + +

Cellaria punctata t xm x

Cellaria paradoxa +

*Aspidostoma livida

Caberea darwinii x x x «x xX

Eupaxia quadrata x

Menipea crispa x * x

Menipea triseriata x x x xu IX

Menipea ornata

Menipea marionensis xr HX x

Bugulella australis x

*Bugulella problematica x

Beania magellanica x x x

*Beania rediviva x

Bugula dentata x

Species per station AS Gel a SG ior ad 1 di 42 89 1 3

xX + + xX

x —b

x —- tw +

oS OX OK 2K OK SP =ib

x OK eK OK

—-

«x —F

Eats

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Ascophoran Cheilostomata. The species are listed in systematic order, stations in order of

increasing depth. New species are indicated by an asterisk (*). X =represented by living

colonies. + = dead colonies only.

100m 500m 700m

Depth station 179 180 184 163 163/ 164 185 23912501234 233 232 162 103 1234131 129 151

164

Cribrilaria innominata x x

Cribrilaria venusta x

*Cribrilaria africana x x x

Figularia philomela x

Figularia sp. x

Escharoides contorta ie Xe ox 9 ex «x xX if i

*Dimorphocella moderna eek t, .caes. it

Exechonella sp. T tT iT

Pachycleithonia mutabilis He oh

Tropidozoum burrowsi t t

*Smittina sitella t t vy

*Smittina ferruginea x

*Smittoidea circumspecta <x XOX

*Smittoidea errata x x

*Smittoidea calcarata x

Parasmittina tropica x « FT 7 y 6

*Parasmittina novella x

Porella capensis + x x

*Escharella discors if

*Mucropetraliella asymmetrica + + + <n

Arthropoma cecilii

Arthropoma circinatum +

Arthropoma sp. x x

Escharina pesanseris x

Escharina waiparaensis x x

Calyptotheca nivea x Xe i ex x

Calyptotheca porelliformis x x

*Emballotheca ambigua x

Stomachetosella balani x

Cleidochasma porcellanum x

Cleidochasma protrusum +

Cleidochasma cribritheca KK 0X

*Cleidochasma perspicua x

Hippoporidra senegambiensis x

Hippoporella spinigera x x x W it

*Hippoporella labiata x

Hippomenella avicularis + 7

Microporella sp. + x

Flustramorpha flabellaris <i

Flustramorpha marginata f + x x

Flustramorpha angusta X, Kei Xx x tT

*Fenestrulina indigena x XX

Trypostega venusta rE Ka XK x

Gigantopora polymorpha xP x

*Gigantopora foraminosa fi = XO Oe Tt

Adeonella majuscula x

Adeonella cracens i

*Adeonella decipiens > Ee iP PG x x it

*Adeonella confusanea x fin x x

*Adeonella conspicua x

“Adeonella distincta x

*Adeonella infirmata x

*Adeonella abdita XM EX eK EX

*Adeonella gibba x + <r xX x

Table 2 continued on next page.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Depth station

*Adeonella alia

Tessaradoma bispiramina

Tessaradoma circella

Sertella lata

*Sertella verecunda

Schizoretepora tessellata

Reteporella dinotorhynchus

*Todictyum flosculum

*Rhynchozoon documentum

*Rhynchozoon beatulum

*Rhynchozoon incallidum

*Rhynchozoon oscitans

*Rhynchozoon stomachosum

*Rhynchozoon ptarmicum

Brodiella longispinata

*Brodiella ignota

Turbicellepora conica

*Turbicellepora valligera

Celleporaria tridenticulata

Celleporaria capensis

Vittaticella sp.

Anoteropora latirostris

Species per station

100m 500m

700m

179 180 184 163 163/ 164 185 2392504234 233 232 162 103 1234131 129 151

T x

x + +

133 au

—- —- X

164

—F ob

x x

18 35

TABLE 3

—+

Cyclostomata. The species are listed in systematic order, stations in order of increasing depth.

New species are indicated by an asterisk (*). X =represented by living colonies. + = dead

colonies only.

Depth station

Diaperoecia

Mecynoecia clavaeformis

Mecynoecia delicatula

Mecynoecia australis

Plagioecia patina

Liripora lineata

Idmidronea contorta

Idmidronea crassimargo

Idmidronea cf. parvula

Idmidronea cf. biporata

Idmidronea cf. antarctica

Idmidronea cf. atlantica

Crisia elongata

Lichenopora novae-zealandiae

Crisina radians

*Hornera erugata

Species per station

100m 500m

700m

179 180 184 163 163/ 164 185 23942504234 233 232 162 103 123W31 129 151

164

+ FF FX + X X XX X

x xX

—- + xX — x

Oh Pal 5

—b <r

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATIC ACCOUNT

ORDER CHEILOSTOMATA

Family Flustridae d’Orbigny, 1852

Flustridae d’Orbigny, 1852: 324. Smitt, 1868: 357. Ryland & Hayward, 1977: 76.

Carbasea Gray, 1848

Carbasea Gray, 1848: 105, 146. Ryland & Hayward, 1977: 79.

Carbasea mediocris Hayward & Cook, 1979

Fig. 1A—B

Carbasea mediocris Hayward & Cook, 1979: 52, fig. 2A-B.

Material

Stations SM 233, SM 239, SM 250.

Remarks

The present material allows a more complete description of this species.

Live colonies were obtained from each of the three stations and together

include all astogenetic and ontogenetic stages. The specimens were up to

55 mm in length, the slender fronds dividing dichotomously at regular intervals;

there were six to eight longitudinal series of zooids. The ancestrula (Fig. 1B)

was slender and elongate and, including its tubular proximal extension,

exceeded 3 mm in length. The basal portion of the colony presents a most

curious appearance, with short cylindrical rhizoids projecting at right angles

from the lateral borders of the ancestrula and the lowest zooids of the colony,

before flexing abruptly and descending in flat fused bundles to the substratum.

Most of the specimens were brooding embryos; the ovicells, partially immersed

in the zooids distal to the maternal zooids, were up to 0,46 mm long and

equally broad, thinly calcified with a faintly striated surface.

The holotype was collected from 550 m depth (Hayward & Cook 1979),

the present material was obtained at depths of 90 m to 580 m.

Family Cupuladriidae Lagaaij, 1952

Cupuladriidae Lagaaij, 1952: 31. Cook, 1965a: 154; 1965b: 192.

Cupuladria Canu & Bassler, 1919

Cupuladria Canu & Bassler, 1919: 77. Lagaaij, 1952: 32. Cook, 1965b: 197.

Cupuladria multispinata (Canu & Bassler, 1923)

Cupularia multispinata Canu & Bassler, 1923: 78, fig. 13H.

Cupuladria multispinata: Cook, 1965b: 210, pl. 2 (fig. 2A-B), fig. 2d.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 1 A-B. Carbasea mediocris Hayward & Cook. A. Ovicelled zooids. B. The ancestrula, and

basal attachment rhizoids. C. Callopora sp. D-F. Amphiblestrum inermis (Kluge). D. A group of

zooids with large avicularia, including two with characteristic ovicells. E. Zooids from a juvenile

colony, with small avicularia. F. Ovicelled zooids, two with distinctive avicularia. G. Amphiblestrum

pontifex sp. nov. Scale = 0,5 mm for C-G; 1 mm for A-B.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

Stations SM 162, SM 179, SM 180, SM 185.

Description

Cupuladria with large zooids (Lz 0,55-0,76 mm). Colonies with coarsely

tuberculate, alternating, radiating ridges basally. Zooids with well-developed

lateral cryptocyst denticles that end in fine spinules.

Remarks

All the colonies found were dead and worn; they were almost certainly

transported. C. multispinata differs from C. owenii (Gray), which also occurs in

South African waters (Cook 19655: 213), in its larger colonies and larger

zooids. The denticulation on the edge of the cryptocyst and the basal

tuberculations are also coarser than in C. owenit.

Distribution

North-west, west, and south-east Africa, 7-105 m.

Discoporella d Orbigny, 1852

Discoporella d’Orbigny, 1852: 472. Cook, 1965b: 219.

Discoporella umbellata (Defrance, 1823)

Fig. 2

Lunulites umbellata Defrance, 1823: 361, pl. 47 (fig. 1a—b).

Discoporella umbellata: Cook, 1965a: 177, pl. 1 (fig. 7), pl. 3. (figs 1, 3, 5-6), fig. 4;

1965b: 221, pl. 3 (fig. 3), fig. 2h. Hayward & Cook, 1979: 44.

Material

Stations SM 131, SM 162, SM 163/164, SM 179, SM 180, SM 184,

SM 185.

Description

Zooids with complete cryptocyst lamina, pierced by several pairs of

opesiules, and other foramina. Opesia D-shaped, straight proximally. Basal

surface with irregular pores or pits, becoming flat and smooth. At colony

maturity a layer of kenozooids alternating with avicularia grows from the

periphery and spreads over the basal surface, which is usually by then flat, not

curved.

Remarks

Most populations of D. umbellata do not show the basal changes that

occur in these specimens. This morphotype was described as ‘D. umbellata

peyroti-type’ by Cook (1965a: 179), because a similar form of colony growth

—————

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

iew

iew of

B. Basal v

Enlarged frontal v

x 26.

ina

f basal lam

. Frontal view of a colony. x 9,3.

view O

ds and avicularia. x 9,3.

Enlarged

. X61.

ing layer of kenozoo

autozoo

ingrow

iscoporella umbellata (Defrance)

the

Fig. 2.

showing

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

was first described as Cupularia peyroti from the European Tertiary. The cause

of the progressive infilling of the concave basal surface in these colonies is

unknown, but may be similar to that which produces infilling in many colonies

of C. doma (see Cook 1965b: 216). The development of a basal layer of

kenozooids and avicularia, however, seems to occur only in Recent populations

of D. umbellata from South Africa. One of the functions of the basal avicularia

appeared to be to discourage the settlement of larvae of sponges, serpulid

worms and other bryozoans, all of which are often found occupying the basal

concavity in other populations of D. umbellata, and in other species of the

Cupuladriidae (see Cook 1965b: 195). Very young dead colonies were present

at nearly all stations, but living colonies with avicularian setae intact were

collected only at Stations SM 179 and SM 180.

Distribution

The D. umbellata species complex has a wide distribution in tropical and

subtropical regions of the Atlantic Ocean and the western Indian Ocean, but

‘peyroti-type’ colonies are known only from South Africa.

Family Setosellinidae Hayward & Cook, 1979

Setosellinidae Hayward & Cook, 1979: 45.

The family was introduced for genera with free-living colonies of

spirally-budded zooids supported by long setiform avicularian mandibles. The

zooidal opesia is extensive, and large ovicells are present.

Setosellina Calvet, 1906

Setosellina Calvet, 1906: 157. Hayward & Cook, 1979: 48.

Setosellina roulei Calvet, 1906

Setosellina roulei Calvet, 1906: 157; 1907: 395, pl. 26 (figs 5-6). Hayward & Cook, 1979: 48,

figs 1A, 17B, 18B.

Material

Stations SM 103, SM 131, SM 164.

Remarks

In contrast to the large number of specimens previously reported

(Hayward & Cook 1979), a very few ‘dead’ colonies were found in the present

collections.

Heliodoma Calvet, 1906

Heliodoma Calvet, 1906: 157. Hayward & Cook, 1979: 50.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Jil

Heliodoma implicata Calvet, 1906

Heliodoma implicata Calvet, 1906: 157; 1907: 396, pl. 26 (figs 7-9). Hayward & Cook,

1979: 50, figs 17A, 18A.

Material

Stations SM 103, SM 123, SM 131.

Remarks

Some of the specimens from stations SM 103 and SM 123 were alive when

collected; only those from SM 103, which have been reported before (Hayward

& Cook 1979) were numerous.

Family Calloporidae Norman, 1903

Calloporidae Norman, 1903: 587. Ryland & Hayward, 1977: 87.

Callopora Gray, 1848

Callopora Gray, 1848: 109, 146. Ryland & Hayward, 1977: 87.

Callopora sp.

Fig. 1C

Material

Stations SM 163, SM 163/164, SM 164, SM 185.

Description

Colony encrusting. Zooids 0,5-0,7 mm long by about 0,4 mm broad.

Opesia oval, cryptocyst narrow; gymnocyst smooth, of variable extent. Spines

distributed around whole of opesia, up to ?12 in number. No sessile avicularia.

Ovicell small, hyperstomial, perhaps with frontal area.

Remarks

Colonies of an unrecognized species of Callopora were found at each of

the above stations. All were dead, and damaged to a greater or lesser extent.

Although it appears to be undescribed, the poor state of the material precludes

the possibility of an adequate taxonomic description and, accordingly, the

species must remain unnamed until further specimens are collected.

Amphiblestrum Gray, 1848

Amphiblestrum Gray, 1848: 103. Ryland & Hayward, 1977: 103.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Amphiblestrum inermis (Kluge, 1914)

Fig. 1D-F

Membranipora inermis Kluge, 1914: 663, pl. 34 (fig. 6).

Lepralia triangularis O'Donoghue, 1924: 43, pl. 2 (figs 11- 12).

Amphiblestrum triangulare: O'Donoghue, 1957: 74.

Material

Stations SM 163, SM 164, SM 184, SM 185.

Description

Colony encrusting, unilaminar. Zooids flat, hexagonal or irregularly

polygonal, separated by shallow grooves; 0,4-0,5 mm long by 0,36—0,4 mm

broad. Cryptocyst flat, occupying half length of zooid, surrounded by a thin

crenellated rim; two widely spaced, evanescent spines distally; opesia trifoliate,

proportions variable, most frequently with proximal portion constituting

one-third or less of total length. Gymnocyst smooth, largely obscured by a

single adventitious avicularium, orientated transversely and acute to frontal

plane. Rostrum hooked, supporting an acute triangular mandible. Ovicell

prominent, recumbent on succeeding zooid, with a large triangular area of

uncovered granular, entooecium frontally; raised border between entooecium

and ectooecium produced at proximolateral corners of ovicell to form

projecting, proximally directed spikes. Within the colony the size of the

avicularian cystid varies astogenetically and the proportions of the opesia vary

ontogenetically.

Remarks

O’Donoghue (1924) noted that his L. triangularis seemed similar to the

Antarctic Membranipora inermis Kluge. The type specimen (BMNH

1963.3.20.7.) is poor, consisting of fragments of several young colonies; there

are no ovicells and most of the zooids lack avicularia. A second specimen from

Saldanha Bay (BMNH 1936.12.30.281) is equally fragmentary, but the zooids

possess avicularia. It is difficult, therefore, to identify O’Donoghue’s material

with M. inermis Kluge. Good examples of the latter species are known from

South Africa, viz. from False Bay (BMNH 1962.6.4.6.), and from Simon’s Bay

(BMNH 1944.1.8.186); both are well-grown colonies with prominent avicularia,

and the False Bay specimen has numerous ovicells showing the triangular,

granular frontal area, and proximolateral processes, characteristic of Kluge’s

species. Unfortunately neither includes early astogenetic stages and it is not

possible to judge whether the fragmentary specimens of O’Donoghue represent

simply early astogenetic stages of M. inermis.

The Meiring Naude material provides some evidence for this supposition,

but the number of specimens is small and additionally suggests an even broader

range of variation. The material described above (station SM 184, Fig. 1D) is

without doubt closest to Kluge’s species, and includes some early zooids with

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 1S

very small frontal avicularia. A specimen from station SM 163 (Fig. 1E), small

and lacking ovicells, has very small avicularia and the zooids resemble closely

both the early astogenetic stages of the colony from SM 184 and O’Donoghue’s

type specimen of L. triangularis. In a specimen from station SM 185 (Fig. 1F)

the free distal edge of the avicularian cystid is flattened and expanded, forming

a structure similar to that seen in species of Foveolaria; however, the ovicell,

the paired spines, the opesia, and the avicularian rostrum are identical to those

of the material described (SM 184).

Amphiblestrum pontifex sp. nov.

Fig. 1G

Material

Holotype: SAM-A26413, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Description

Colony encrusting. Zooids oval, broad and flat, distinct, separated by

deep grooves. Gymnocyst reduced but visible, continuous laterally with

incurved lateral walls, smooth and hyaline. Cryptocyst extensive, comprising

four-fifths of total zooid length, flat and smooth, encircled by a raised and

thickened mural rim. Opesia subterminal, less than half length of cryptocyst,

and distally less than half its width; shape characteristic, opesiular indentations

small, initially continuous with distal part of opesia, but later isolated from it.

Two widely spaced distal oral spines present in newly developed zooids, later

lost. Avicularium medially situated on gymnocyst, orientated transversely;

cystid globose, rostrum elongate, acute-triangular, its distal end curved to left

or right. Ovicell prominent, spherical, small; smooth surfaced, with a longitudi-

nal median ridge marking the thickened edges of a narrow frontal foramen.

Etymology

Pontifex (L.)—high priest, an allusion to the shape of the opesia.

Remarks

A. pontifex may be distinguished from other species of Amphiblestrum by

the form of the opesia, in particular by the isolation of the opesiular indenta-

tions. Several live colonies encrusting the large specimen of Dimorphocella

were collected.

Measurements (means of 20 values) in mm

Lz Iz

OT 0,48

Copidozoum Harmer, 1926

Copidozoum Harmer, 1926: 226.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Copidozoum transversum sp. nov.

Fig. 3A

Material

Holotype: SAM-A26414, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Other material: station SM 250.

Description

Colony encrusting. Zooids oval, large. Frontal surface almost entirely

membranous: gymnocyst apparent as a small triangular area of calcification on

zooids at a row bifurcation; cryptocyst forming simply a narrow granulated

border, mural rim thin, no spines. Vicarious avicularia conspicuous, cystid

tumid; rostrum orientated transversely, 0,24-0,34 mm long, distal portion slen-

der, with a blunt tip, proximal portion rounded, opesia oval, with a granular

cryptocyst, mandible articulating against paired, thickened condyles. Ovicells

not found.

Etymology

Tranversus (L.)—crosswise, referring to the orientation of the avicula-

rium.

Remarks

Living colonies were collected from both the above stations.

Measurements (means of 20 values) in mm

ez Iz

0,64 0,42

Crassimarginatella Canu, 1900

Crassimarginatella Canu, 1900: 369. Hastings, 1945: 69.

Crassimarginatella marginalis (Kirkpatrick, 1888)

Fig. 3B

Membranipora marginalis Kirkpatrick, 1888: 74, pl. 7 (fig. 2).

Crassimarginatella marginalis: Hastings, 1945: 78, fig. 2B.

Material

Stations: SM 163, SM 163/164, SM 164, SM 239, SM 250.

Remarks

In the size of the zooids, the relatively broad cryptocyst, and the

morphology of the vicarious avicularia, the Meiring Naude specimens closely

resemble the Mauritian type specimen of C. marginalis (BMNH 1888.12.5.8.).

The Meiring Naude specimens tend to be more thickly calcified, but as no live

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 15

0,5

Fig. 3. A. Copidozoum transversum sp. nov. B. Crassimarginatella marginalis (Kirkpatrick). C-—E. Chap-

eria familiaris sp. nov. C. Zooids from a live collected colony, showing the forked proximal spines. D. Two

dead zooids, showing the extent of the occlusor laminae. E. A group of zooids with ovicells in different

stages of development. F. Chaperia capensis (Busk).

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

juvenile colonies were collected, the majority of the material representing dead

colonies, this may be regarded as an ontogenetic effect.

Foveolaria Busk, 1884

Foveolaria Busk, 1884: 68. Harmer, 1926: 246.

Foveolaria imbricata (Busk, 1884)

Fig. 4 =

Amphiblestrum imbricatum Busk, 1884: 65, pl. 15 (fig. 3).

Membranipora imbricata: Marcus, 1922: 16, fig. 9.

Material

Stations SM 129, SM 131, SM 151, SM 163, SM 163/164, SM 179,

SM 180, SM 184, SM 185.

Description

Colony arising from an encrusting sheet of zooids, forming an erect

cylindrical growth. Zooids in alternating linear series around the entire axis of

the branch; broadly diamond-shaped, as wide as long, distal edge raised and

distinctly crenellated. Frontal membrane underlain by a smooth, extensive

cryptocyst, deeply concave; opesia subtriangular, occupying less than one-third

of total zooid length. Frontal surface largely hidden by a tall, broad avicula-

rium, developed on the proximal half of the cryptocyst; cystid cylindrical at

first, broadened distally, with a thin irregular edge. Rostrum situated on the

lateral face of the cystid, acute triangular, apically orientated, with stout

condyles for articulation of the mandible. No spines. Ovicells not observed.

Remarks

Busk’s original description of this species (1884: 65) is confusing. He

considered he was viewing a unilaminar colony encrusting another, erect,

bryozoan species, although his material is clearly an erect cylindrical colony.

The relative proportions of gymnocyst and cryptocyst, the origin of the frontal

avicularium, the nature of the ovicell and, perhaps, colony form are the

features most likely to prove useful in distinguishing Foveolaria (as defined by

Levinsen 1909: 152) from Amphiblestrum (above). At present A. imbricata

Busk seems more correctly placed in Foveolaria than in Amphiblestrum.

Foveolaria sp.

Fig. 5A

Material

Station SM 162.

Description

Colony erect, cylindrical, slender. Zooids in triple whorls, elongate,

rounded distally, tapered proximally; lateral walls flared distally, forming an

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 4. Foveolaria imbricata (Busk). A. An entire colony. X12. B. Detail to

show the avicularia. X 82.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. A. Foveolaria sp.., the dead specimen from SM 162. B. Micropora similis sp. nov. C-F. Macropora

africana sp. nov. C. Zooids from the growing edge. D. The ancestrula, and periancestrular zooids.

E. A vicarious avicularium. F. An ovicelled zooid. G. Exechonella sp. Scale = 0,5 mm for A-F; 1 mm for G.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES i)

arched hood. Cryptocyst extensive, flat, opesia comprising less than one-

quarter of zooid length. A broad-based, possibly cylindrical, avicularian cystid

present on the proximal region of the cryptocyst. Ovicell spherical. No spines.

Remarks

Dead fragments of an unrecognized species of Foveolaria were obtained

from station SM 162. Unfortunately all of the features of the zooids were

damaged to a greater or lesser extent, in particular the larger part of each

avicularium was missing; the condition of the material thus precludes a com-

plete description of the species.

Family Chaperiidae Jullien, 1888

Chaperiidae Jullien, 1888: 61. Brown, 1952: 94.

Chaperia Jullien, 1881

Chaperia Jullien, 1881: 163. Brown, 1952: 94. Gordon, 1982.

The genus Chaperia comprises a large number of species described from

localities throughout the southern hemisphere. The synonymy of many of these

species is confused to the extent that no useful comments may be made

regarding systematic relationships or distributional patterns until the majority

of described species has been critically re-examined. Brown (1952: 94) clarified

the status of Chaperia Jullien, noting that the type species was the originally

designated C. australis Jullien, 1881, and selected a neotype for C. acanthina

(Lamouroux, 1825). Brown also indicated that C. acanthina may prove to be a

senior synonym of C. australis, and it would seem useful to consider here

whether this synonymy may be accepted.

Jullien (1881: 163) introduced C. australis as a new name for Membrani-

pora spinosa Busk, 1879; Busk had referred his material (from Kerguelen) to

Flustra acanthina (Lamouroux, 1825) but had derived the name from the

non-Linnean ‘Flustra épineuse’ used by Lamouroux in the explanation of his

plate (1825, Atlas, pl. 89, figs 1-2). Although Jullien (1881) was describing

specimens of C. australis from South Africa, he was quite explicit in introducing

the name for Busk’s Kerguelen material, and the possible synonymy of C.

acanthina and C. australis thus depends on the identity of Busk’s specimens.

Jullien’s South African specimens pose another problem. Marcus (1922)

described material from South Africa under the name C. acanthina var.

australis Jullien, which is here referred to C. capensis (Busk) (q.v.). The South

African material of C. australis may prove to be identical to Busk’s Kerguelen

Specimens, or it may be identifiable with Marcus’s material; only examination

of Jullien’s specimens will decide the issue, but its result is irrelevant to the

identity of C. australis, which is determined solely by the Busk specimen from

Kerguelen. Fortunately this is still extant (BMNH 1899.7.1.1155, 1156); it

comprises a single, well-grown young colony encrusting a small rhodophyte.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Comparison with the neotype of C. acanthina (BMNH 1930.1.16.26A) and with

other material from the Falkland Islands (BMNH 1935.3.6.59, 316) shows that

Busk’s specimen may be assigned confidently to C. acanthina (Lamouroux).

The zooids of all four specimens are of a similar size (0,5—0,6 mm long by

0,4-0,5 mm broad) with a granular cryptocyst comprising two-fifths of the total

length. The opesia is surrounded by jointed spines of variable thickness, up to

0,6 mm long; these number five or six on the Falkland Islands specimens, and

six or seven in the Kerguelen colony. The occlusor laminae in all cases are

distinct, slightly curved but not converging markedly towards the distal end of

the zooid, the visible length being 0,2 mm.

It may be shown, therefore, that C. australis Jullien is a junior subjective

synonym of C. acanthina (Lamouroux), which accordingly becomes the type

species of Chaperia Jullien. The subsequent synonymy of C. acanthina 1s

adequately provided by Brown (1952: 95); C. acanthina was also discussed by

Harmer (1926: 229), but his material included the Siboga specimens from the

tropical East Indies which appear to differ from Brown’s neotype, most notably

in possessing up to ten small spines. Finally, and inexplicably, Jullien (1888)

figured specimens of a Chaperia from Cape Horn which he attributed to

‘Flustra spinosa’, noting that it was the same as that to which he had in 1881

given the name Chaperia australis. His text, though brief, suggests that he had

by then concluded that C. acanthina and C. australis were identical, but his use

of the spurious ‘Flustra spinosa’ perhaps served to obscure his opinion.

Chaperia multifida (Busk, 1884)

Fig. 6

Membranipora galeata var. multifida Busk, 1884: 64.

Chaperia multifida: Kluge, 1914: 673, text-fig. 44. Marcus, 1922: 7, fig. 3.

Membranipora galeata var. multifida: O'Donoghue & De Watteville, 1935: 205.

Material

Stations SM 163, SM 163/164, SM 179, SM 180, SM 184, SM 1835.

Description

Colony encrusting, forming patches or cylindrical growths around erect

substrata; developing as erect bilaminar sheets or solid branching cylinders.

Zooids hexagonal or irregularly oval, separated by indistinct sutures,

0,44-0,65 mm long by 0,26-0,4 mm broad. Cryptocyst forming a narrow, finely

granular, concave rim, most prominent proximally and tapering rapidly towards

spine bases. Opesia circular or oval, marginally longer than wide in the larger

zooids, occupying three-fifths to three-quarters of total frontal length. Occlusor

laminae distinct, quite divergent distally; arising on each side from a point

below the distal spine and extending to half-way along the lateral margin of the

opesia. Two pairs of distolateral spines, thick, jointed at base with a distinct

coelomic cavity; proximalmost pair broadening rapidly, developed as multi-

branched palmate structures arching over and obscuring most of the frontal

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 6. Chaperia multifida (Busk). A. Portion of a live collected colony (on right)

with spines and pedunculate avicularia intact, and a dead fragment for comparison

(on left). x 29. B. Bleached fragment of a live collected colony. x 79.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

surface of the zooid. Distalmost pair of spines typically erect, cylindrical,

clavate or spatulate, rarely palmate. A single sessile avicularium at the distal

end of each zooid, situated between the distalmost pair of spines; rostrum

triangular, distally orientated, with an incomplete cross-bar and a transversely

oval foramen in the palate; rarely, the rostrum may be broadened to give a

narrowly spatulate shape. Erect, elongate pedunculate avicularia numerous,

most frequently situated adjacent to the sessile avicularium on one or both

sides; sometimes developed between adjacent zooids, arising from the lateral

wall of one of them; one or more of the distal spines may also be replaced by

pedunculate avicularia. Ovicell prominent, hemispherical, with an oval or

irregular frontal area; in fertile zooids the distal pair of spines and the

avicularia are suppressed.

Remarks

The material from SM 179 and SM 185 comprised living colonies retaining

a reddish coloration in alcohol and forming cylindrical growths around hydroid

stems. All the material from the other stations consisted of dead fragments;

some of these were encrusting, unilaminar patches and some bilaminar sheets,

but the majority were fragments of cylindrical or flattened branching colonies

up to 10 mm high. Bleached zooids from SM 179, lacking spines and peduncu-

late avicularia are compared with those of a dead cylindrical fragment in

Figures 6A-B, and it may be seen that, despite the variation in colony

morphology, only one species is represented.

Chaperia capensis (Busk, 1884)

Figs 3k

Amphiblestrum capense Busk, 1884: 67, pl. 23 (fig. 3).

Chaperia acanthina var. australis: Marcus, 1922: 6, fig. 2.

Membranipora galeata var. inermis O'Donoghue, 1924: 38, pl. 1 (fig. 9).

Chapperia acanthina var. australis: O’Donoghue & De Watteville, 1944: 415.

Material

Stations SM 163/164, SM 179, SM 180.

Description

Colony encrusting, forming cylindrical growths around erect substrata,

developing as erect, solid cylindrical structures, branching irregularly. Zooids

broad and flat, typically hexagonal, broader than long, separated by shallow

grooves; 0,5-0,6 mm long by about 0,6 mm broad. Cryptocyst finely granular,

concave, broadest proximally, narrowing towards the distal end. Opesia trans-

versely oval, occupying about three-fifths of total frontal length; occlusor

laminae well developed and distinct, seen to converge distally even in

unbleached undamaged material, extending lateroproximally almost to the

proximal edge of the opesia. One short cylindrical spine present at each distal

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES ue

corner, variable in length but typically no longer than the width of the opesia.

Avicularia and ovicells absent.

Remarks

The proportions of the zooids seem to vary according to the form of

growth of the colony; in regular cylindrical branches they tend to be longer

than broad, but may broaden considerably in encrusting sheets or irregular

branches. The type material from Simon’s Bay, South Africa consists of several

solid cylinders lacking bases. The specimens from Meiring Naude stations

SM 163/164 and SM 179 form hollow cylinders around hydroids; dead frag-

ments from SM 180 consisted mostly of solid cylinders, but some included a

hollow, encrusting cylindrical base.

The material described and figured by Marcus (1922: 6, fig. 2) as C.

acanthina var. australis clearly belongs in the synonymy of Chaperia capensis;

however, as discussed above, the identity of Jullien’s (1881) South African

records of C. australis can be decided only by examination of his specimens. C.

acanthina var. polygonia Kluge (1914: 676, text-fig. 47), from Simon’s Bay,

placed by Marcus (1922) in the synonymy of his C. acanthina var. australis,

differs from C. capensis in possessing five or six distal oral spines and widely

divergent occlusor laminae. It is probably correctly regarded as a variant of C.

acanthina.

Chaperia stephensoni O’Donoghue & De Watteville, 1935

Chaperia stephensoni O'Donoghue & De Watteville, 1935: 205, pl. 5 (fig. 1), pl. 6 (fig. 11).

O’Donoghue, 1957: 74.

Material

Stations SM 163, SM 184.

Remarks

Only small dead fragments of this species were recovered. C. stephensoni

may be recognized by the transversely oval opesia overarched by the distal wall

of the zooid, and by the single distal avicularium, which has a characteristic

elongate lanceolate rostrum curved slightly to the left or right at its tip. Two

spine bases are visible at each distal corner and the sessile avicularium is often

flanked on one or both sides by the swollen bases of erect avicularia.

Chaperia familiaris sp. nov.

Figs 3C-E, 7A

Material

Holotype: SAM-—A26415, station SM 162, 32°55’S 28°31’E, 630 m.

Other material: stations SM 163, SM 180.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. A. Chaperia familiaris sp. nov. x 48. B. Dactylostega prima sp. nov. x 64. C. Dactylos-

tega tubigera (Busk), BMNH 1887.12.9.3462. x 24.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 5)

Description

Colony encrusting, forming small patches, retaining a reddish-brown

colour when preserved in alcohol, but fading when dried. Zooids hexagonal,

occasionally rounded distally, typically as wide as long, but tending to broaden,

separated by distinct sutures. Cryptocyst flat, finely granular, forming a

symmetrical plate of constant width around two-thirds of the periphery of the

Opesia between the proximalmost pair of spines, its free edge distinctly

bevelled. Opesia transversely oval, occupying approximately half the length of

the frontal surface. Occlusor laminae distinct, each extending from a point

one-third of the distance along the distal wall to the proximolateral corner of

the zooid; not markedly convergent distally, although in damaged zooids they

may be seen to join distally at the mid-line of the zooid. Two pairs of

distolateral spines, distinctly shorter than the length of the opesia; distal pair

cylindrical, proximal pair forked and slightly incurved. In brooding zooids only

the proximal pair is present. Ovicell prominent, broader than long, hemispher-

ical or somewhat irregular; ectooecium with a transversely oval frontal

foramen. No avicularia.

Etymology

Familiaris (L.)—familiar, an allusion to the morphological features

characteristic of the genus.

Remarks

Live specimens were obtained from all three stations, those from SM 162

and SM 180 were encrusting the large arenaceous foraminiferan Schizammina

pinnata (Pearcey) (see p. 144). C. familiaris seems to be most similar to C.

capensis (above) but differs in possessing two pairs of spines, most particularly

in the short, forked proximal pair, and in its very characteristic ovicells.

Measurements (means of 10 values) in mm

Lz Iz Lop lop

0,48 0,47 0,30 0,31

Chaperia sp.

Fig. 9A

Material

Stations SM 163, SM 163/164, SM 180, SM 184, SM 185.

Description

Colony forming unilaminar sheets, encrusting or possibly erect, or only

loosely attached to substratum. Zooids flat, typically broader than long,

tapered proximally and rounded distally, 0,6-0,9 mm long by 0,7-1,0 mm

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

broad. Cryptocyst flat or slightly depressed distally, granular; opesia transversely

oval, comprising up to two-thirds total length of zooid, no occlusor laminae

visible. Distal border of zooid with seven to nine closely spaced spine bases. No

avicularia or ovicells. Vertical walls very deep, up to 1,0 mm in some fragments;

in basal view zooid boundaries are marked by distinct ridges. Both vertical and

basal walls densely perforated by large multiporous septula, each in a distinct pit.

Remarks

Detached fragments of this distinctive species were found at each of the

stations indicated. All were dead and worn, and none gave any indication of

what the colony form might have been. It appears to belong to Chaperia but, in

view of the poor state of the material and the uncertainty regarding the status

of at least one South African species of this genus (p. 19), it seems inappropri-

ate to assign a specific name.

Notocoryne Hayward & Cook, 1979

Notocoryne Hayward & Cook, 1979: 54.

Notocoryne cervicornis Hayward & Cook, 1979

Notocoryne cervicornis Hayward & Cook, 1979: 55, fig. 3.

Material

Stations SM 151, SM 163, SM 163/164, SM 180, SM 184, SM 185.

Remarks

The present material shows a greater range of colony size than that

originally described by Hayward & Cook (1979), with specimens of 9,5 mm,

10 mm, and 11 mm in length. In all cases the characteristic faceted club-shape

was constant. None of the colonies was alive when collected.

Family Hiantoporidae MacGillivray, 1895

Hiantoporidae MacGillivray, 1895: 60. Osburn, 1950: 97.

Dactylostega gen. nov.

Colony encrusting, unilaminar, or erect, bilaminar. Cryptocyst developed

aS a narrow rim; gymnocyst reduced, obscured. Avicularia interzooidal, devel-

oped from a series of chambers present between the autozooids; other cham-

bers with simple frontal foramina assumed to be kenozooidal. Secondary

calcification originating from interzooidal chambers, projecting over the frontal

membrane of the autozooids as a series of blunt, irregular spikes infilling

concavities between autozooids and forming an enveloping ooecial cover.

Autozooids communicating with interzooidal chambers via small uniporous

septula. Ovicell hyperstomial, closed by zooidal operculum. Spines absent.

Type species: Dactylostega prima sp. nov.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES ay,

Etymology

Dactylos (Gr.)—finger; stegos (Gr.)—roof, describing the digitate frontal

shield seen in later ontogeny.

Dactylostega prima sp. nov.

Figs 7B, 8A

Material

Holotype: SAM-—A26416, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Other material: stations SM 131, SM 162, SM 163, SM 163/164, SM 164,

SM 179, SM 180, SM 184, SM 185.

Description

Colony encrusting, unilaminar, or forming erect bilaminar sheets. Zooids

oval, deep; boundaries clear at the growing edge of the colony, but obscured by

the development of interzooidal avicularia and ?kenozooidal chambers. Opesia

oval, rimmed by a narrow, granular, basally deflected cryptocyst with distinct

beaded edge; of constant width around whole of opesia. Spaces between zooids

infilled by a continuous series of chambers, some of which develop as avicula-

ria, others forming irregular bodies with a frontal foramen (?opesia) sur-

rounded by a granular area of calcification (?cryptocyst). Avicularia most

frequently situated distolaterally to zooids, but often lateral in position as well,

particularly at the bifurcation of zooid rows. Interzooidal calcification increas-

ing in later ontogeny, appearing continuous over all of the interzooidal cham-

bers and forming a continuous projecting rim around the opesia of each zooid,

the edge developing short irregular processes that extend above the frontal

membrane. Ovicell immersed, hyperstomial; slightly broader than long with a

rather rectangular outline; apparently with a small central ectooecial fenestra in

early ontogeny, later covered by calcification derived from the interzooidal

chambers

Etymology

Primus (L.)—first, denoting the type species.

Remarks

The overarched ‘pericystal’ processes seen in D. prima produce an effect

similar to that of the frontal shield seen in, for example, Arachnopusia.

However, it may be seen from the micrographs (Fig. 7B) that in D. prima this

calcification emanated from the curious interzooidal chambers that characterize

the genus. The possible homology of the central foramen of the chamber and

its surrounding area of granular calcification with the opesia and cryptocyst of

the autozooid, and the small uniporous septula that link each chamber with the

zooids adjoining it, suggest its kenozooidal nature.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. A. Dactylostega prima sp. nov., showing ovicells and kenozooidal chambers

in various stages of development. x 50. B. Cellaria punctata (Busk). X 80.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Zs)

Live encrusting colonies were collected from each of the stations listed

above, and dead fragments of bilaminar sheets were obtained from SM 180 and

SM 164.

Foveolaria tubigera Busk (1884: 68), presently known only from Simon’s

Bay, Cape of Good Hope, shows many similarities to D. prima, in particular

the interzooidal chambers, and clearly belongs in the same genus (see fig. 7C).

Other species referable to Dactylostega are Hincksina nigrans (Hincks) (see

Osburn 1950: 44, pl. 5 (figs 3-4)) and Membraniporidra spissimuralis as

described by Hayami (1975: 102, pl. 13 (fig. 10)). Both these forms occur in the

Northern Pacific, D. nigrans being circumarctic and D. spissimuralis Hayami

(non Canu & Bassler) having been reported from the Pliocene of Japan. A

similar development of supraopesial denticulations and frontal avicularia occurs

in Odontionella cyclops var. tessellata (see Brown 1952).

Measurements (means of 25 values) in mm

Lz Iz

0,62 0,42

Family Arachnopusiidae Jullien, 1888

Arachnopusiidae Jullien, 1888: 62. Moyano, 4970: 260.

Arachnopusia Jullien, 1888

Arachnopusia Jullien, 1888: 62. Moyano, 1970: 260.

Arachnopusia corniculata sp. nov.

Fig. 9B—C

Material

Holotype: SAM—A26417, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 131, SM 163/164, SM 164, SM 250.

Description

Colony encrusting, unilaminar, zooids oval, separated by shallow grooves,

distinct at growing edge, but obscured in later ontogenetic stages. Frontal

membrane occupying most of frontal surface; a minimal area of smooth

gymnocyst proximally, and a narrow, granular cryptocystal rim, occasionally

developing a scalloped edge. Frontal shield developed from the proximal and

lateral edges of the zooid as a variable number of processes, fused medially to

obscure two-thirds of the total length of the frontal surface, leaving a D-shaped

distal aperture and a variable number (typically 3-6) of irregularly shaped

foramina. Area of each foramen subsequently reduced by further calcified

processes, which may fuse to divide it into two. Proximal border of aperture

straight, or developing a short, lobed or multipointed, medial process at an

oblique angle to the frontal plane. Similar processes occasionally present on the

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. A. Chaperia sp. B-C. Arachnopusia corniculata sp. nov. B. Zooids from the growing edge of the

colony, including one with an ovicell. C. Two zooids from a juvenile colony showing stages in the

development of the frontal shield. D-E. Bugulella problematica sp. nov. D. Zooids with distal spines, and

avicularia developed from lateral septula. E. Zooids with spines almost encircling the opesiae; avicularia and

zooids arising from lateral septula. F-G. Beania rediviva sp. nov. F. Showing the rounded distal end of the

autozooid. Note that the rostra of the two distal avicularia are broken short. G. An avicularium in lateral

view.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 31

distal edge of the aperture; oral spines absent. Avicularia adventitious, typically

paired, lateral to aperture; rostrum acute triangular, slightly curved, directed

distally or distomedially. Less frequently, avicularia may occur proximally, on

the edges of the zooid, with variable orientation. Ovicell hyperstomial, partly

immersed, closed by zooidal operculum; as broad as long, smooth and imper-

forate, becoming obscured by calcification derived from the frontal shield of the

distally succeeding zooid. Ancestrula tatiform, 0,4 mm long; opesia oval with a

narrow cryptocyst, bordered by spines. First zooids budded from the ancestrula

have five long oral spines.

Etymology

Corniculata (L.)—horned, referring to the lip of the aperture.

Remarks

The genus Arachnopusia has a wide distribution within the southern

hemisphere, although none of the described species has been reported from

South Africa. A. corniculata may be distinguished readily from other species of

the genus by the relatively large, paired, oral avicularia, and in particular the

laterally curved rostrum, and by the denticulate proximal border of the aper-

ture. The absence of oral spines is also a distinctive feature.

Measurements (means of 25 values) in mm

Lys Iz

0,64 0,27

Family Microporidae Gray, 1848

Microporidae Gray, 1848: 115, 147. Ryland & Hayward, 1977: 112.

Micropora Gray, 1848

Micropora Gray, 1848: 115, 147. Ryland & Hayward, 1977: 112.

Micropora similis sp. nov.

Fig. 5B

Material

Holotype: SAM—A26418, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163, SM 163/164, SM 164, SM 180, SM 233,

SM 239, SM 250.

Description

Colony forming thin unilaminar sheets. Zooids small, broad and flat,

hexagonal, or with proximal half distinctly tapered. Opesia (orifice) twice as

wide as long, proximal border straight, lateral corners rounded to give a more

or less elliptical effect; somewhat more angular in old or ovicellate zooids.

oe ANNALS OF THE SOUTH AFRICAN MUSEUM

Lateral walls finely crenulate, slightly raised above the frontal surface, forming

small scarcely discernible bosses adjacent to the opesia. Cryptocyst finely

granular, punctured by numerous small pores; gently convex medially, distally

becoming concave close to the opesiules and steeply raised at the proximal edge

of the opesia. Opesiules situated close to opesia, distinct. Ovicell recumbent on

succeeding zooid, wider than long and rather depressed, with a transverse

frontal ridge, frequently developed medially into a low umbo. Avicularia

developed sporadically throughout the colony, each situated immediately distal

to an autozooid and orientated transverse to distalproximal axis; rostrum

semi-elliptical or bluntly triangular, oblique to frontal plane of zooid, cross-bar

slender, cylindrical.

Etymology

Similis (L.)—resembling, a reference to the similarity of this species to

other members of the genus.

Remarks

This small species resembles the north-eastern Atlantic Micropora nor-

mani Levinsen, which also possesses interzooidal avicularia. M. similis is

distinguished from this and other species of the genus, however, by its slender

opesia and by the densely punctured frontal wall.

Measurements (means of 15 (zooids) or 10 (avicularia) values) in mm

jz Iz Lav

0,54 0,44 ‘Os

Family Steginoporellidae Hincks, 1884, emend. Bassler, 1953

Steganoporellidae Hincks, 1884: 358.

Steginoporellidae: Bassler, 1953: G171.

Steginoporella Smitt, 1873

Steginoporella Smitt, 1873: 15. Pouyet & David, 1979: 764.

Steganoporella: Cook, 1964a: 45.

Steginoporella buskii Harmer, 1900

Steganoporella buskii Harmer, 1900: 272, pl. 12 (fig. 13), pl. 13 (figs 33-35). Cook, 1964a: 46,

pl. 1 (figs 1-3), fig. 1

Steginoporella buskii: Pouyet & David, 1979: 771, pl. 1 (fig. 9), text-fig. 2.

Material

Stations SM 131, SM 163, SM 163/164, SM 164.

Remarks

Fragments of living colonies were collected from stations SM 163/164 and

SM 164, but the specimens from SM 131 and SM 163 were of dead, transported

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 25)

material. The specimens resemble others from Port Elizabeth (the type locality)

in having a fairly low level insertion of the cryptocyst in the distal wall (see

Cook 1964a). Although S. magnilabris (Busk) is known from east Africa, all

records of it from South Africa are referable to S. buskii.

Family Macroporidae Uttley, 1949

Macroporidae Uttley, 1949: 175. Brown, 1952: 134.

Macropora MacGillivray, 1895

Macropora MacGillivray, 1895: 54. Brown, 1952: 134.

Macropora africana sp. nov.

Fig. 5C-F

Material

Holotype: SAM-—A26419, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163/164, SM 184, SM 185.

Description

Colony encrusting, unilaminar. Zooids broadly hexagonal, convex, sepa-

rated by deep grooves. Primary orifice semi-elliptical, proximal border straight,

closed by a distinctive calcified operculum with a finely granular, punctate

surface. Four or five short, thickened, distal oral spines present but lost in later

ontogenetic stages, only the brown chitinized bases remaining. Frontal wall

finely granular, perforated by numerous small pores; large dietellae visible at

the base of the vertical walls in zooids at the growing edge. Vicarious avicularia

sporadically distributed throughout colony, of a primitive form; cystid of similar

size to an autozooid, rostrum linguiform with a triangular palatal foramen.

Ovicell inflated, large (0,8 mm long), obscuring most of the frontal wall of the

succeeding zooid from which it is derived; with a frontal umbo and a series of

conspicuous radiating striations and marginal slits, closed by zooidal opercu-

lum. Ancestrula similar to later zooids but smaller (0,56 mm long), with seven

oral spines.

Etymology

Africanus (L.)—African.

Remarks

Macropora is an ancient genus known mostly from the Tertiary deposits of

Australia and New Zealand. The type species, M. centralis MacGillivray

(Miocene, Victoria), is possibly synonymous with M. grandis (Hutton) whose

distribution in time extends from the Lower Miocene to the present (Brown

1952). Recent specimens have been reported from New Zealand (Uttley &

Bullivant 1972) and the Philippines (Brown 1952). M. grandis is a larger species

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

than M. africana, with zooids up to 1,2 mm long, and is further distinguished

by a proportionally more elongate operculum and distinct oral shelf, by the

possession of a distinct raised peristome, by a complete absence of oral spines,

and by the absence of a zooid orifice distal to the ovicell.

Measurements (means of 25 values) in mm

EZ Iz

0,78 0,62

Family Cellariidae Hincks, 1880

Cellariidae Hincks, 1880: 103. Ryland & Hayward, 1977: 119.

Cellaria Ellis & Solander, 1786

Cellaria Ellis & Solander, 1786: 18. Ryland & Hayward, 1977: 119.

Cellaria tectiformis Hayward & Cook, 1979

Cellaria tectiformis Hayward & Cook, 1979: 69, fig. 7.

Material

Stations SM 103, SM 131, SM 151, SM 233, SM 234.

Remarks

A single large, living, colony was obtained from each of the two stations

SM 233, and SM 234.

Cellaria punctata (Busk, 1852)

Fig. 8B

Salicornaria punctata Busk, 1852: 366 (partim).

Cellaria gracilis: Marcus, 1922: 19, fig. 11.

Cellaria punctata: Harmer, 1926: 337, pl. 21 (figs 14-16), text-fig. 13a.

Material

Stations SM 163/164, SM 164, SM 180.

Description

Colony forming diffuse, straggling tufts up to 50 mm high. Joints consist-

ing of tangled masses of brown, chitinous tubes, each arising from inconspicu-

ous calcified sockets on the frontal surfaces of the zooids at the distal and

proximal end of each internode, possibly representing kenozooids. Internodes

straight or gently curved, up to 8 mm long, with a maximum width of 0,7 mm,

typically broadening distally, with distinct dilatations along length; comprised

of five to seven longitudinal series of zooids. Autozooids regularly

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 35)

hexagonal, separated by distinct sutures; cryptocyst granular, concave, the

central area particularly depressed and delimited by a prominent ridge. Opesia

situated in distal third of zooid, semicircular; distal border raised, forming a

distinct cowl, with finely beaded edge; proximal border with a projecting,

rounded lip, and a pair of thick lateral denticles. Avicularia vicarious, infre-

quent, typically present at distal end of each internode, close to joint; as large

as autozooid, with a broadly triangular rostrum occupying half frontal surface,

supporting a distally directed triangular mandible. Proximal border of avicula-

rian opesia with an anvil-shaped lip, distal corners of which fuse with sides of

opesia to delimit paired lateral opesiules; palate scarcely developed, with a

large rounded foramen. Fertile zooids constitute the characteristic dilatations,

typically seven in a whorl; ovicellar orifice inconspicuous, largely occluded by a

proximal projection from the succeeding zooid.

Remarks

This species was described by Busk (1852) from Queensland and rede-

scribed by Harmer (1926), who examined material from a number of Indo-

Pacific localities and included Marcus’s (1922) South African specimens and

Thornley’s (1905, given as 1895 in error) Ceylon material of ‘Cellaria johnsoni’

in his synonymy. The present material shows some differences from the

specimens studied by Harmer (for example, Holborn Island, Queensland,

BMNH 1928.9.13.87), most notably in zooid numbers. In the Holborn Island

specimen each internode comprises four to six longitudinal series of zooids; in

the Meiring Naude material the proximal end of each internode has five series

of zooids, broadens rapidly to six, and has seven in the fertile swellings. The

avicularian rostrum is more rounded in the Australian material, but zooid

morphology and size are comparable with the South African specimens. A

“specimen from the Red Sea (BMNH 1963.8.10.37 pt.) is identical to the

Meiring Naude specimens.

Measurements (means of 25 values) in mm

Ly Iz

0,48 0,29

Cellaria paradoxa Hayward & Cook, 1979

Cellaria paradoxa Hayward & Cook, 1979: 71, fig. 8.

Material

Station SM 103.

Remarks

Worn internodes of this species were present in the coarser sediment

fraction from station SM 103. A somewhat similar species with dimorphic

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

zooids was described from the Eocene of North and South Carolina as C.

bifaciata by Canu & Bassler (1920: 274, pl. 40 (figs 14-17)).

Family Aspidostomatidae Jullien, 1888

Aspidostomatidae Jullien, 1888: 77. Harmer, 1926: 322.

Aspidostoma Hincks, 1881

Aspidostoma Hincks, 1881: 159. Harmer, 1926: 323.

Aspidostoma livida sp. nov.

Fig. 10

Material

Holotype: SAM-—A26420, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Other material: stations SM 131, SM 164, SM 184, SM 239.

Description

Colony erect, bilaminar, reticulate, forming broad, plate-like lobes, larg-

est fragment obtained measuring 45 X50 mm; alcohol-preserved material

retaining a deep blue-grey colour. Fenestrulae up to 4 mm long, apparently

regularly distributed, about four occurring in each cm? of colony surface.

Zooids hexagonal, convex, separated by distinct grooves, thickly calcified with

a coarsely granular surface. Frontal wall convex proximally, dipping distally

towards a small deep-set opesia; proximal edge of opesia (delimiting the

‘polypide tube’) forming a thickened axehead-shaped lip. A thick median

ridge extends for a short distance proximally from the outer edge of the

opesial lip. Distal end of zooid raised as a prominent hood, frequently deve-

loped on each corner into short flattened processes. Operculum transversely

oval, thickly calcified, white. Ovicell globose, rather flattened frontally, with a

coarsely granular surface; opening via a hooded aperture distal to the zooid

operculum. Interzooidal avicularia sporadically distributed over the colony;

rostrum acute triangular, distolaterally directed; cross-bar incomplete, palate

with an elliptical central foramen. Fenestrulae rimmed by single series of large

kenozooids.

Groups of frontally budded zooids appear to inaugurate new laminae. The

form of the colony could not be discerned, but the large bilaminar plates were

flat or gently dished, rather than convoluted, and new plates appear to develop

perpendicular to their predecessors. Unlike the fenestrate colonies of Dimor-

phocella moderna (see p. 48) from the same station, A. livida shows no

formation of central ribs of calcification.

Etymology

Lividus (L.)—bluish, referring to the colour of the colony.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

il ofa

ds and

Deta

icelled zoo

ing two ov

laria. <x 40

includ

and avicu

lls,

x 40.

f a colony,

ing Ovice

ice

Portion o

Develop

damaged ov

nov.

x< DS

da sp

ids

Aspidostoma liv

ight) two kenozoo

Fig. 10.

(bottom r

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The material from station SM 239 comprised a large number of reticulate

fragments which may have represented just one large colony, no basal attach-

ment was found. The only erect Recent species of Aspidostoma develop slender

cylindrical colonies, although Waters (1905: 243) described specimens of A.

giganteum (Busk) that displayed a loosely anastomosing form. Erect branched

species of Aspidostoma were described from the Tertiary of New Zealand by

Brown (1952).

Measurements (means of 25 values) in mm

1e7; Iz

0,89 O75

Family Scrupocellariidae Levinsen, 1909

Scrupocellariidae Levinsen, 1909: 130. Ryland & Hayward, 1977: 128.

Caberea Lamouroux, 1816

Caberea Lamouroux, 1816: 128. Ryland & Hayward, 1977: 128.

Caberea darwinii Busk, 1884

Caberea darwinii Busk, 1884: 29, pl. 32 (fig. 6c—f).

Caberea darwinii: Hastings, 1943: 374, pl. 5 (figs 1-3), text-figs 21, 22A—C, 23A—D, 24A.

Material

Stations SM 163/164, SM 179, SM 184, SM 185, SM 239.

Remarks

The geographical distribution of Caberea darwinii has been discussed at

length by Hastings (1943). It extends from New Zealand westward to the

Patagonian Shelf and is widespread in Antarctic and Subantarctic waters; it has

been recorded from Kerguelen, Prince Edward Island, and Marion Island, but

has not been reported before from South Africa.

Eupaxia Hasenbank, 1932

Eupaxia Hasenbank, 1932: 321, 363.

Eupaxia quadrata (Busk, 1884)

Cellularia quadrata Busk, 1884: 18, pl. 5 (fig. 5).

Eupaxia incarnata Hasenbank, 1932: 363, fig. 30A—C.

Eupaxia quadrata: Hayward & Cook, 1979: 63, fig. 6C-F.

Material

Station SM 232.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 39

Remarks

This species was redescribed and figured by Hayward & Cook (1979). A

single fragment of a live colony was collected during the present survey.

Menipea Lamouroux, 1812

Menipea Lamouroux, 1812: 183. Harmer, 1923: 339. Hastings, 1943: 331.

Menipea crispa (Pallas, 1766)

Cellularia crispa Pallas, 1766: 71.

Menipea crispa: Marcus, 1922: 11. Hastings, 1943: 332. O’Donoghue, 1957: 75, figs 1-2.

Material

Stations SM 163/164, SM 164, SM 179.

Remarks

M. crispa appears to have a limited geographical distribution, from

Saldanha Bay (South Africa) to Madagascar (Hastings 1943). Live specimens

were collected from each of the three Meiring Naude stations, which ranged in

depth from 50 to 90 m.

Menipea triseriata Busk, 1852

Menipea triseriata Busk, 1852: 22, pl. 23 (figs 2-4).

Menipea triseriata: Harmer, 1923: 342, pl. 17 (fig. 18), pl. 19 (figs 40-42). O’Donoghue,

IETS WO

Material

Stations SM 163, SM 164, SM 179, SM 180, SM 185.

Remarks

This distinctive species is known only from South Africa. The present

material includes several very large, luxuriantly branched colonies.

Menipea ornata (Busk, 1852)

Cellularia ornata Busk, 1852: 20, pl. 20 (figs 3-4).

Menipea ornata: Harmer, 1923: 340.

Menipea flabellum: Marcus, 1922: 13, fig. 7.

Cellularia infantae O’Donoghue, 1924: 30, pl.1 (fig. 6). O'Donoghue & De Watteville,

1935: 207; 1937: 12.

Material

Station SM 185.

Remarks

The confused synonymy of this species has been clarified by Harmer

(1923) and by Hastings (1943: 332).

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Menipea marionensis Busk, 1884

Menipea marionensis Busk, 1884: 21, pl. 4 (figs 3, 3a).

Menipea marionensis: Harmer, 1923: 341, pl. 17 (fig. 22), pl. 19 (figs 43-45).

Material

Stations SM 179, SM 180, SM 185.

Remarks

This species appears to be known only from the Cape of Good Hope. The

present specimens were obtained living, at depths of 80 and 90 m.

Family Bicellariellidae Levinsen, 1909

Bicellariellidae Levinsen, 1909: 93. Ryland & Hayward, 1977: 146.

Bugulella Verrill, 1879

Bugulella Verrill, 1879: 472. Maturo & Schopf, 1968: 36.

Bugulella australis Hayward & Cook, 1979

Bugulella australis Hayward & Cook, 1979: 64, fig. 6A-B. Millard, 1980: 143.

Material

Station SM 233.

Remarks

The material comprised a tangled mass, probably representing several

colonies, and was living when collected. Following its recent description in the

first report of the Meiring Naude Bryozoa (Hayward & Cook 1979), substantial

material was found in the Galathea deep-sea collections from a station in the

Tasman Sea at 610 m (Hayward 1981).

Measurements (means of 20 values) in mm

IZ Lop lop

0,74 0,28 0,18

Bugulella problematica sp. nov.

Fig. 9D-E

Material

Holotype: SAM-A26421, station SM 233, 32°15,2’S 29°09,8’E, 540-580 m.

Description

Colony erect, straggling, delicate; composed of branching uniserial chains

of zooids forming a dense tangled tuft. Zooids elongate, club-shaped: oval

distally, tapered proximally to a slender tubaeform shape; thinly calcified and

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 41

translucent. Opesia oval, with a narrow cryptocyst border; typically with three

pairs of slender spines regularly spaced around distal end, rarely up to ten

spines distributed around whole of opesial border. Each zooid budding up to

three new zooids: one distal, one on each side from a point level with the

proximal half of the opesia. Distal bud constant, colony developing as a

uniserial chain, some or all of whose members produce secondary chains, on

one or both sides, perpendicular to the first; tertiary chains may form in the

same way. The lateral budding points, marked by large, distinct septula, may

instead give rise to irregularly tubular kenozooids that link with septula of

zooids in neighbouring branches; rarely, such kenozooids develop at the distal

budding point. Avicularia pedunculate, with short, semi-elliptical mandibles;

single, more usually paired, arising from lateral budding points and thus

precluding development of side chains where they occur. Rarely, a single

avicularium develops distally, frontal to the proximal tubular portion of the

next zooid. Ovicell prominent, terminal; spherical, with finely tessellated sur-

face, closed by zooidal operculum.

Etymology

Problematicos (Gr.)—problematical.

Remarks

The budding pattern of Bugulella problematica, and the paired lateral

avicularia, serve to distinguish it from B. australis (above), in which lateral

branches are initiated by a second distal bud, which becomes fused with a

tubular structure (?kenozooid) arising from the lateral septulum of its twin. The

avicularium of B. australis is constantly distal in position, and the zooids tend to

be larger than those of B. problematica. However, zooids of the latter species

may be found with up to ten opesial spines, typical of B. australis, and the

ovicell is practically identical in both species. Specimens of the two species were

found intermingled in the same sample and some doubt must remain that the

apparently significant difference in budding pattern may prove to be an

astogenetic effect.

Measurements (means of 20 values) in mm

Lz Lop lop

0,51 O25 0,15

Family Beaniidae Canu & Bassler, 1927

Beaniidae Canu & Bassler, 1927: 14. Ryland & Hayward, 1977: 150.

Beania Johnston, 1840

Beania Johnston, 1840: 272. Ryland & Hayward, 1977: 150.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Beania magellanica (Busk, 1852)

Diachoris magellanica Busk, 1852: 54, pl. 67 (figs 1-3).

Beania magellanica: Hastings, 1943: 414, figs 34C, 35G.

Material

Stations SM 163, SM 179, SM 239.

Distribution

Widespread; reported from the Mediterranean to the Falkland Islands,

from Australia to Japan, and throughout the Indian Ocean. Recorded from

South Africa by Marcus (1922) and O’Donoghue (1957).

Beania rediviva sp. nov.

Fig. 9F-G

Beania erecta: Hasenbank, 1932: 342, fig. 1SA—C.

Material

Holotype: SAM—A26422, station SM 250, 31°59,3’S 29°22,5’E, 150-200 m.

Etymology

Redivivus (L.)—renewed, alluding to the newly recognized identity of the

species.

Remarks

The Antarctic-Subantarctic Beania erecta Waters was redescribed by

Hastings (1943: 416), who excluded Hasenbank’s (1932) record from Agulhas

Bank from her synonymy. Hasenbank’s figure showed several significant differ-

ences from typical B. erecta: the distal ends of the zooids- were smoothly

rounded, lacking the paired lateral oral projections seen in B. erecta, and the

avicularia were very large, with a slender, strongly hooked rostrum comprising

almost half the total length. These features are shown particularly well in the

present specimen (Fig. 9G) and it is appropriate to introduce a new name for

Hasenbank’s species.

Family Bugulidae Gray, 1848

Bugulidae Gray, 1848: 110, 146. Ryland & Hayward, 1977: 151.

Bugula Oken, 1815

Bugula Oken, 1815: 89. Ryland & Hayward, 1977: 151.

Bugula dentata (Lamouroux, 1816)

Acamarchis dentata Lamouroux, 1816: 135, pl. 3 (fig. 3a—b).

Bugula dentata: O’Donoghue, 1924: 33; 1957: 82. Harmer, 1926: 439, pl. 30 (figs 5-6), pl. 32

(figs 21-25).

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 43

Material

Station SM 180.

Remarks

This well-characterized species was reported from South Africa by

O’Donoghue (1924), who collected it on the lower shore at Agulhas Light,

Cape Province. All South African records were documented in a later paper

(O’Donoghue 1957).

Family Cribrilinidae Hincks, 1880

Cribrilinidae Hincks, 1880: 182. Hayward & Ryland, 1979: 56.

Cribrilaria Canu & Bassler, 1929

Cribrilaria Canu & Bassler, 1929: 33. Hayward & Ryland, 1979: 62.

Cribrilaria innominata (Couch, 1844)

Lepralia innominata Couch, 1844: 114, pl. 22 (fig. 4).

Cribrilaria innominata: Harmelin, 1970: 84, figs 1d-f, 2, pl. 1 (figs 4-6). Hayward & Ryland,

M/S 564> figs 17:

Material

Stations SM 163, SM 163/164.

Remarks

This widespread species is characterized by its umbonate frontal shield,

and the presence of a large, conspicuous, suboral lacuna. Live colonies were

collected at both of the above stations.

Cribrilaria venusta (Canu & Bassler, 1925)

Fig. 11A

Puellina venusta Canu & Bassler, 1925: 22, pl. 2 (fig. 5).

Cribrilaria venusta: Harmelin, 1976a: 180, pl. 2 (figs 3-5).

Material

Station SM 239.

Remarks

This is a distinctive species recognized initially by its broad and relatively

flat frontal shield composed of numerous slender costae. These are usually little

thickened, though occasionally developing a series of peripheral knobs. The

first pair of costae proximal to the orifice fuse and thicken medially to form a

triangular umbo and there are no suboral pores; conversely the pores between

the umbonate first pair of costae and the second pair are typically large and

distinct. There are five oral spines. The avicularian rostrum is slender, acumi-

nate, and frequently with a gentle lateral curve.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. A. Cribrilaria venusta (Canu & Bassler). B. Figularia sp., two zooids, with an ovicell from a

missing zooid. C. Cribrilaria africana sp. nov., a specimen from SM 164. D. Vittaticella sp. E-F. Giganto-

pora foraminosa sp. nov. E. Portion of a branch, showing progressive elaboration of the spiramen.

F. Outline diagram of primary orifice. Scale = 0,5 mm for A—D, F; 1 mm for E.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 45

Cribrilaria venusta has only recently been redescribed by Harmelin

(1976a), who summarized its known distribution. The present material is

closely similar in zooid size (0,46-0,6 mm xX 0,38-0,44 mm) and morphology to

specimens collected from the western end of the English Channel. Although

the present record marks a significant extension of its geographical range, the

distribution of C. venusta, like those of other species of Cribrilaria, is

imperfectly known and will probably prove to be very broad.

Cribrilaria africana sp. nov.

Fig. 11C

Material

Holotype: SAM—A26423, station SM 164, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163, SM 250.

Description

Colony encrusting. Zooids oval, rather flat; costate frontal shield

comprising larger part of frontal surface, with only a narrow rim of gymnocyst.

Costae with small marginal tubercles, but without a central umbo. Orifice with

straight, smooth proximal edge, six or seven closely spaced oral spines. Suboral

region (between orificial bar and first pair of frontal costae) broad, triangular

and flat, with up to five large pores, and many small pores. Avicularium small,

Squeezed in between successive zooids, or occasionally developed on gymno-

cyst, semi-pedunculate; rostrum slender, c.0,1 mm long. Ovicell small, spher-

ical, hyaline, with a small, median umbo, frequently with an avicularium closely

adjacent to it on each side.

Etymology

Africanus (L.)—African.

Remarks

This species is readily distinguishable from those reviewed by Harmelin

(1976a) by the conspicuous suboral pores and the small semi-pedunculate

avicularium.

Measurements (means of 16 values) in mm

187, Iz

0,39 0,29

Figularia Jullien, 1886

Figularia Jullien, 1886: 608. Hayward & Ryland, 1979: 70.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figularia philomela (Busk, 1884)

Cribrilina philomela Busk, 1884: 132, pl. 17 (fig. 6).

Figularia philomela: Hayward & Cook, 1979: 76, fig. 9B.

Material

Station SM 163.

Remarks

The present material, comprising several small encrusting live colonies,

was collected in waters far shallower than those sampled by the first series of

Meiring Naude stations.

Figularia sp.

Fig. 11B

Material

Station SM 163.

Remarks

The material comprised two fragments of live colonies, the largest

including five complete zooids, together with portions of four others. The

zooids were large and flat (0,8-0,9 mm x 0,7-0,8 mm); the costate frontal

shield was small, constituting less than half of the total frontal surface, although

costae and intercostal pores were distinct. Two or more large pseudopores were

present on each of the costae, and the rest of the frontal surface, and the

surface of the ovicell, were covered with similar pseudopores, indicated by

brown chitinized cuticle. This species does not seem to have been described

before, but the paucity of material precludes a complete description being

presented here.

Family Exochellidae Bassler, 1953

Exochellidae Bassler, 1953: G205. Hayward & Ryland, 1979: 78.

Escharoides Milne Edwards, 1836

Escharoides Milne Edwards, 1836: 218. Hayward & Ryland, 1979: 78.

Escharoides contorta (Busk, 1854)

Eschara contorta Busk, 1854: 89, pl. 108 (figs 1-3).

Mucronella contorta: Busk, 1884: 155, pl. 20 (fig. 9). O’Donoghue & De Watteville, 1937: 18.

Escharoides contorta: O’Donoghue, 1957: 88, figs 10-11.

Material

Stations SM 151, SM 162, SM 163, SM 163/164, SM 164, SM 180,

SM 184, SM 185; SM 239.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 47

Remarks

This species is known only from South Africa. It was particularly

abundant in the present collections and live colonies were collected from most

of the stations listed. Colony form varied greatly, from simple encrusting sheets

to broad bilaminar plates and slender, branching cylindrical growths.

Family Adeonidae Jullien, 1903

Adeonidae Jullien in Jullien & Calvet, 1903: 53. Cook, 1973: 246.

Dimorphocella Maplestone, 1903

Dimorphocella Maplestone, 1903: 140.

Description

Colony erect, branching, bilaminar. Autozooids with sinuate secondary

calcified orifice. Frontal shields umbonuloid, zooids surrounded by marginal

frontal septula. Brooding zooids dimorphic, large, with wide, non-sinuate

orifices; frontal shields with several spiramina. Adventitious avicularia uni-

lateral or paired, acute; interzooidal marginal avicularia sometimes present,

mandibles slung on paired condyles.

Remarks

The genus is inferred to have umbonuloid ontogeny of zooid frontal

shields, as in Adeona and Adeonellopsis (see Cook 1973). The enlarged

brooding zooids and condylate avicularia are also characteristic of the family.

Dimorphocella differs from the other genera in the strongly sinuate autozooid

orifice, and the apparent lack of frontal spiramen pores in autozooids.

Dimorphocella is known from the Tertiary of Australia, but has not been

reported as Recent (see below). The genus was introduced somewhat

informally by Maplestone (1903), who stated, ‘I propose this genus for a form

presently to be described, and Adeonella triton McG’. The species then

described, D. pyriformis Maplestone (1903: 141, pl. 16 (fig. 1)), was illustrated

with strongly sinuate autozooid orifices, and paired, medially orientated

avicularia. The brooding zooids had an enlarged distal cavity and a central,

circular, porous frontal area, and paired avicularia orientated distally.

Adeonella triton MacGillivray (1895: 90, pl. 19 (fig. 23)) was originally

described with similar characters, except that the autozooidal avicularia were

unilateral and the brooding zooids were not distally expanded. A. triton, as

later described by Maplestone (1903, pl. 16 (fig. 2)) had paired, distally

orientated avicularia and a denticulate process on the proximal side of the

brooding zooid orifice, but appears to be the same species. Although Canu &

Bassler (1920: 571, fig. 170A—B; 1929: 384) indicated A. triton as ‘genotype’ of

Dimorphocella, Maplestone’s (1903) choice was definite, if unorthodox, and D.

pyriformis was later listed as type species by Bassler (1935: 95; 1953: G213,

fig. 161, 5). Canu & Bassler (1920) included Dimorphocella portmarina

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maplestone (1913: 359, pl. 28 (fig. 5)), a Recent Australian species, in

Dimorphocella, but this form has conspicuous autozooidal spiramina and is

referable to Adeona (see Canu & Bassler 1929).

Waters (1881: 340, pl. 18 (fig. 85)), described another Tertiary Australian

species as Schizoporella submersa. No brooding zooids were described and

none have been found in specimens in the British Museum (Natural History)

collections (D32901-2, D34733, see fig. 14A). The autozooidal characters of S.

submersa are similar to those of D. pyriformis. S. submersa as described by

MacGillivray (1895: 82, pl. 11 (figs 8-9)) may not be the same species (see

Brown 1958: 60).

Dimorphocella moderna sp. nov.

Figs 12-13

Material

Holotype: SAM-A26424, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Other material: stations SM 164, SM 179, SM 180, SM 185, SM 250.

Description

Colony erect, bilaminar, fenestrate, deeply pigmented, formed of slightly

curved, frequently anastomosing plates. Central part of plates becoming

thickened and forming branched strengthening ribs. Zooids arranged in

alternate bifurcating and recombining series, forming fenestrulae that are

bordered by large interzooidal avicularia. Autozooids with a complete series of

marginal frontal septula, sometimes with additional frontal septula. Frontal

shields otherwise imperforate except for a suboral spiramen which is occluded

early in ontogeny. Secondary calcified orifice with a deep proximal sinus.

Brooding zooids very large, raised distally, with a wide orifice, denticulate

proximally. Central frontal area with several spiramen pores. Adventitious

avicularia usually unilateral, occasionally absent in autozooids, in all cases

orientated distally. Large, interzooidal fenestral avicularia acute, with raised

rostra, mandibles hinged on prominent paired condyles, orientated distally.

Etymology

Modernus (L.)—of the present, referring to the Recent occurrence of the

species.

Remarks

D. moderna is represented by worn fragments from all the stations listed,

except from SM 239. This station provided abundant material that was alive

when collected. Although no complete colony is present, and the substratum

and mode of attachment is unknown, the larger plates indicate that colonies

probably exceed 100 mm in height or diameter. Each plate is slightly curved,

and the zooids on the outward-facing convex side are occluded by extrazooidal

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

12. Dimorphocella moderna sp. nov. A. Frontal surface of a branch, showing

dimorphic orifices. x 48. B. Basal surface. x 36.

ANNALS OF THE SOUTH AFRICAN MUSEUM

ion of two branches. X 27.

ion of a branch showing

fus

Sect

il showing

. X 66.

Deta

lar

kening

icarlous avicu

nov.

ith v

hocella moderna sp

f a branch w

imorp

in oO

ig. 13.

B. Marg

, x 24°5:

frontal thic

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 51

calcification earlier in astogeny than those of the concave surface (see Fig.

12A-B). Although the budding pattern of plates is primarily in one plane,

secondary plates arise later in astogeny at right angles to the primary plates,

and anastomose with them. These secondary plates originate from a frontally

budded group of three to seven zooids that later form a bilaminar expansion.

The central rib of each plate is formed by localized thickening of the frontal

shields of zooids, which becomes extrazooidal. The thickening also occurs on

the convex surface in advance of that on the concave surface (Fig. 13C), but

at the base of the colony is equally developed on both surfaces, forming a

cylinder 6mm in diameter. The frontal shields may exceed 2,80 mm in

thickness, but the zooidal cavity is not occluded. Neighbouring unthickened

zooids, even at the base of the colony, possess lophophores, and are inferred

to have been capable of feeding. The fenestrulae are oval and are formed by

the regular bifurcation and recombination of zooid series. They are more

frequent in occurrence than those of Aspidostoma livida (see p. 36). Some

growing edges that had been damaged before collection show that regenerated

zooids (with lighter pigmentation) re-form fenestrulae in the same position as

before. There is some variation in orifice width among autozooids (see Fig.

13A). This appears to be related to the bifurcation and recombination of

series, but by analogy with Adeonella (see Harmer 1957), may also reflect a

sexual function.

The umbonuloid nature of the frontal calcified shield may be seen at the

growing tip of a few young secondary branches of anastomosing plates, which

have been protected. The exposed growing edges of the primary plates have

been damaged. The minute frontal spiramen is apparently occluded early in

ontogeny. This may be correlated with the presence of a sinus in the secondary

calcified orifice. This, by analogy with similar sinuate cryptocystidean forms

(e.g. Laminopora), would allow passage of water into the ascus during

protrusion of the lophophore.

Colonies are the substratum for several encrusting bryozoans, cirripedes,

foraminifera, serpulids, etc., and have numerous small ophiuroids in the

cavities formed by the anastomoses of plates.

No Recent species truly referable to Dimorphocella has been reported

before (see above), and the genus does not now seem to be represented in

Australian seas. Fossil Australian colonies were erect and branched, but D.

moderna resembles the fenestrate colonies of Adeona, which are found from

the Australian Tertiary-to-Recent. These colonies reach a height and diameter

of 25 cm, and are attached and supported by flexible rooting structures formed

by alternating cuticular and calcified kenozooidal elements. Large colonies of

Adeona also develop strengthening ribs by extrazooidal frontal calcification. At

present, the attachment of both D. moderna and the fenestrate Aspidostoma

livida, which were abundant at station SM 239, is unknown.

D. moderna closely resembles D. triton, differing in its colony form and

the presence of fenestral avicularia.

a2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (means of 20 values) in mm

7, Iz Lbr.z Ibr.z

0,61 0,35 0,69 0,53

Lor lor Lbr.or l.br.or

0,15 0,13 0,06 OMA

Lad.av. Lint.av

0,24 0,45

Lm Im

0,14 0,25

Family Exechonellidae Harmer, 1957

Exechonellidae Harmer, 1957: 651.

Exechonella Duvergier, 1924

Exechonella Duvergier, 1924: 18, see Cheetham, 1966: 62. Canu & Bassler, 1927: 4. Cook,

1967: 337.

Exechonella sp.

Fig. 5G

Material

Stations SM 164, SM 185, SM 250.

Remarks

Only small and badly damaged colonies of this species were found. The

zooids were up to 1,3 mm long by 0,9 mm broad, pyriform in shape with the

distal end formed into a tubular peristome, broken short in all cases. The frontal

wall consists of an umbonuloid shield perforated by large round pores, rimmed

by concentric calcification, indicating that they are progressively infilled.

Avicularia, spines and ovicells were not evident. Sufficient detail of the zooid

morphology was preserved to suggest that this is a species of Exechonella (Cook

1967), but the material was too damaged to permit identification to species.

Family Watersiporidae Vigneaux, 1949

Watersiporidae Vigneaux, 1949: 15, 20.

The family includes genera with finely pseudoporous frontal shields and

opercula with well-developed sclerites. Avicularia are usually absent, but

polymorphic zooids may occur (Cook 1979). Ovicells are present in some

species of Pachycleithonia but absent in other genera.

Pachycleithonia Canu & Bassler, 1930

Pachycleithonia Canu & Bassler, 1930: 25. Cook, 1983b.

Colonies encrusting. Zooids large; orifice with almost rectangular sinus

and often massive condyles. Frontal cuticle and opercula dark brown or purple.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 53)

The affinities and general characters of the genus, which has a worldwide

tropical and subtropical distribution, have been discussed by Cook (19835).

Ovicellate brooding zooids have not been described in the type species, P.

nigra Canu & Bassler, from the Galapagos Islands, but distinctive, reticulate,

hyperstomial ovicells are known in P. mutabilis and P. africana.

Pachycleithonia mutabilis (Canu & Bassler, 1929)

Fig. 14B

Galeopsis mutabilis (?partim) Canu & Bassler, 1929: 273, pl. 28 (figs 4-5), ?fig. 111A-B, ?non

pl. 28 (fig. 6).

Gigantopora mutabilis: Harmer, 1957: 883, pl. 40 (fig. 8).

Pachycleithonia mutabilis: Cook, 1983b.

Material

Stations SM 129, SM 131.

Description

Pachycleithonia with elongated tubular peristome and large peristomial

spiramen. Zooids communicating by distal and lateral septula situated at the

base of the vertical walls, surrounded by calcified buttresses. Frontal septula

large: one pair on each side of the orifice, another pair at each lateral corner of

the zooid. Ovicell hyperstomial not closed by the zooidal operculum; formed by

a very thinly calcified, or wholly cuticular inner capsule derived from the

maternal zooid, surrounded by a reticulate ooecial cover derived from the

frontal shield of the distal zooid.

Remarks

The Meiring Naude material consists only of three small fragments, one of

which has a single ovicell. Like many of the species described here, P. mutabilis

is a relatively shallow-water form and these specimens represent transported

debris. The elongated peristome, with its prominent spiramen, is easily

damaged and is not present in all zooids.

Both Harmer (1957) and Cook (1983b) have noted that the material

originally described by Canu & Bassler (1929) from the Philippine Islands

appears to have comprised two species. One of these (Canu & Bassler 1929,

pl. 28, figs 4-5) resembles the material described by Harmer (1957) from

Indonesia, and also the Meiring Naude fragments; the other (pl. 28, fig. 6) has

larger zooids and was described as having minute lateral oral avicularia.

However, this latter photograph had been retouched, and it appears that the

‘avicularia’ are, in fact, the elongated cavities (areolae) above a distal pair of

frontal septula. A. H. Cheetham (1979 in litt.) examined the original specimen

and commented, “They do appear to be aerolae, and ... can be seen to lie

above large septular openings’. The opercula figured by Canu & Bassler (1929,

fig. 111A—B) might have originated from either or both of their specimens. The

ANNALS OF THE SOUTH AFRICAN MUSEUM

ithonia mutabilis

Pachycle

x 64.

: B.

18.46.

Xx 63

BMNH 1882

tmorphocella submersa (Waters)

(Canu & Bassler)

ig.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 55)

opercula of the Siboga specimens, and of a specimen from the Mascarene

Islands (BMNH 1882.10.18.46), closely resemble those of Watersipora arcuata

Banta (see Ryland 1974, fig. 3B), and those of the west African species

P. africana Cook (19836). P. africana is very similar to P. mutabilis in several

other characters, and also has reticulate ovicells. Unlike P. mutabilis, however,

the ovicell is closed by the operculum, and although the peristome is elongated

it is not tubular and a spiramen rarely develops.

Distribution

P. mutabilis is known from the Philippines and Indonesia, Mauritius and

south-east Africa.

Family Euthyrisellidae Bassler, 1953

Euthyrisellidae Bassler, 1953: G226.

Tropidozoum Harmer, 1957

Tropidozoum Harmer, 1957: 1106. Cook, 1975: 161. Cook & Chimonides, 1981b: 64.

Colonies rooted, cellariiform, internodes connected by cuticular keno-

zooidal joints. Zooid frontal surfaces occupying only part of the curved face of

the internode; the other side consisting of a column of extrazooidal coelom,

limited by a cuticular wall. Frontal shields depressed, with foramina; hypostegal

coelom on both sides of the calcification. Brooding zooids large, with inflated

basal walls and dimorphic orifices. Avicularia absent.

The characters and relationships of this interesting genus are discussed by

Cook & Chimonides (19815).

Tropidozoum burrowsi Cook & Chimonides, 1981

Tropidozoum sp. Cook, 1975: 165, pl. 1 (NB explanation of pls. 1 and 3 transposed), fig. 2B.

Tropidozoum burrowsi Cook & Chimonides, 1981b: 65, figs 5—6, 10.

Material

Stations SM 131, SM 151.

Description

Tropidozoum with narrow basal coelom. Zooids with small frontal

foramina; orifice with a deep triangular sinus. Orifice of brooding zooid wide,

without a sinus.

Remarks

T. burrowsi is known to live in relatively shallow water (15 m), as does the

closely related Indonesian species, T. cellariiforme Harmer. The worn

internodes from stations SM 131 and SM 151 (780 m and 900 m respectively)

have obviously been transported. Previous records from South Africa (BMNH

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

1949.11.10.184, off Durban, 90 m) were also of transported internodes, and it

would appear that living colonies will eventually be found only in shallow or

coastal waters.

Distribution

Madagascar and South Africa.

Family Smittinidae Levinsen, 1909

Smittinidae Levinsen, 1909: 335. Hayward & Ryland, 1979: 98.

Smittina Norman, 1903

Smittina Norman, 1903: 120. Hayward & Ryland, 1979: 98.

Smittina sitella sp. nov.

Fig. 15D-E

Material

Holotype: SAM-A26425, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163, SM 163/164, SM 185, SM 239.

Description

Colony encrusting. Zooids oval, convex, separated by distinct grooves.

Primary orifice slightly wider than long, proximal border largely occupied by a

very broad lyrula, condyles small and generally inconspicuous. Peristome well

developed, deep, distal border encroaching on to frontal surface of succeeding

zooid, typically bearing one or two lateral lobes and incorporating medioproxi-

mally a conical avicularian cystid. Rostrum situated on distal face of cystid,

elongate, triangular, acute to frontal plane. Adventitious avicularia also present

adjacent to orifice, typically paired, occasionally single; cystid short, conical,

rostrum directed laterally, or proximally, or oblique to either direction. Frontal

wall finely granular with a few widely spaced pores, each becoming deeply

immersed and conspicuous in later ontogeny. With the development of the

suboral avicularium the pores proximal to the orifice become occluded and in

highly calcified zooids frontal perforation appears to be limited to marginal

areas. Ovicell hyperstomial, prominent, spherical, with regularly spaced pores.

Ectooecium produced as a conspicuous frontal lip.

Etymology

Sitella (L.)—a little bucket, referring to the shape of the ovicell.

Measurements (means of 20 values) in mm

Lz Iz

0,67 0,44

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 3//

Fig. 15. A-C. Smittina ferruginea sp. nov. A. Zooids from the growing edge. B. Ovicelled zooids, with

additional avicularia. C. Outline diagram of primary orifice. D-E. Smittina sitella sp. nov. D. A group of

zooids, with ovicells and lateral avicularia. E. Outline diagram of primary orifice. F. Smittoidea errata sp.

nov. G-—H. Smittoidea circumspecta sp. nov. G. A group of zooids, with varying development of the suboral

avicularia. H. Outline diagram of primary orifice. I-J. Celleporaria capensis (O’Donoghue & de

Watteville). I. Three autozooids and a vicarious avicularium. J. A young zooid with oral spines intact.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Smittina ferruginea sp. nov.

Fig. 1SA—C

Material

Holotype: SAM—A26426, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Description

Colony encrusting, multilaminar, forming broad irregular sheets. Zooids

large, hexagonal, rather flat, separated by shallow grooves. Primary orifice

subterminal, wider than long; lyrula broad and low, occupying greater part of

proximal border, condyles short, quadrate, conspicuous. No oral spines.

Peristome low, scarcely projecting above frontal surface of zooid; distal border

formed from frontal calcification of succeeding zooid, incorporating proximally

the cystid of a small, median suboral avicularium. Rostrum of avicularium

triangular or semi-elliptical, partly enclosed within peristome, acute to frontal

plane and directed proximally. Additional avicularia may be developed lateral

to the peristome, single or paired; rostrum slender, elongate, straight or gently

curved, typically directed proximally or obliquely proximally, rarely obliquely

distal. Frontal wall finely granular, closely perforated by numerous round

pores, each becoming immersed in a distinct pit as calcification continues.

Ovicell hyperstomial, prominent, spherical, finely granular and regularly

perforated by numerous small round pores; ectooecium forming a frontal lip

continuous with the rim of the peristome.

Etymology

Ferruginus (L.)—rust-coloured.

Remarks

Several large well-grown colonies were collected from station SM 239,

encrusting the living colony of Aspidostoma livida. The preserved material

retained a dull purplish brown colour with scattered patches of deeper reddish

brown. These patches corresponded to sites of active frontal budding, marking

the development of new laminae, and the pigment was seen to be concentrated

in the polypides of the young zooids, and in a narrow fringe around the

undifferentiated growing edge of each lamina.

Measurements (means of 25 values) in mm

Lz Iz

0,87 0,58

Smittoidea Osburn, 1952

Smittoidea Osburn, 1952: 408. Hayward & Ryland, 1979: 108.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 3

Smittoidea circumspecta sp. nov.

Fig. 15G-H

Material

Holotype: SAM—A26427, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 184, SM 185, SM 239.

Description

Colony encrusting, forming small round white patches. Zooids small,

hexagonal, broad and convex, separated by deep grooves. Primary orifice

longer than wide, distinctly narrowed proximally, with broad, basally deflected

condyles; proximal border with a slender lyrula, fragile and frequently missing.

Five or six slender closely spaced oral spines on the distal and lateral borders of

the orifice; peristome scarcely developed, forming at the most a low lateral wall

on each side. Suboral avicularium rarely single, typically twinned, frequently

tripled or quadrupled; cystid short, cylindrical and erect, rostrum facing

distally, perpendicular to frontal plane, mandible semicircular. The avicularian

complex forms a two-, three-, or four-lobed digitate process immediately

proximal to, and largely obscuring, the orifice. Frontal wall finely granular,

with a single series of conspicuous marginal pores. Ovicell spherical, recumbent

on succeeding zooid, with numerous small round pores.

Etymology

Circumspectus (L.)—guarded, an allusion to the oral avicularia.

Remarks

The elongate orifice, the closely spaced oral spines and the cluster of

suboral avicularia impart a highly characteristic appearance to this species,

which allows it to be readily distinguished from all other species of Smittoidea.

Measurements (means of 20 values) in mm

Lz Iz

0,46 0,37

Smittoidea errata sp. nov.

Fig. 15F

Smittoidea ?hexagonalis: Hayward & Cook, 1979: 89, fig. 12A-B.

Material

Holotype: SAM-—A26428, station SM 163/164, 33°04,6’S 28°06,6’E, 90 m.

Other material: station SM 239.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Colony encrusting. Zooids quadrate, flat or slightly convex, separated by

distinct grooves. Primary orifice broader than long, with a short square lyrula

occupying about half of the proximal border; condyles distinct, triangular,

basally deflected. Three short distal oral spines present in newly budded zooids;

peristome a low thickened rim enclosing orifice distally and _ laterally,

proximally incorporating a small suboral avicularium. Mandible semicircular,

acute to frontal plane, directed proximally. Occasionally replaced by an

enlarged spatulate avicularium, directed proximolaterally (Hayward & Cook

1979). Frontal wall fine grained, with a series of marginal pores. Ovicell

hyperstomial, recumbent on succeeding zooid; oval, thinly calcified, with about

twenty small irregularly shaped frontal pores.

Etymology

Erratum (L.)—a mistake, referring to the previous misidentification of this

species.

Remarks

The single specimen of this species collected from station SM 86 during

the first Meiring Naude cruise (Hayward & Cook 1979) was erroneously

ascribed to ‘Smittina’ hexagonalis O’Donoghue 1924, which, in fact, lacks a

suboral avicularium. Three small living colonies were present in the second

collection, two of which were incubating embryos.

Measurements (means of 15 values) in mm

Lz Iz

0,67 0,51

Smittoidea calcarata sp. nov.

Fig. 16A—-B

Material

Holotype: SAM—A26429, station SM 233, 32°15,2’S 29°09,8’E, 540-580 m.

Description

Colony encrusting, unilaminar, delicate and hyaline. Zooids flat and

broad, irregularly polygonal, separated by shallow grooves. Primary orifice

wider than long, proximal border with a tapered peg-like lyrula, condyles

sharp, slender and conspicuous; distal border almost straight, two short distal

oral spines visible in peripheral zooids, occluded by the development of

succeeding zooids. Peristome short, erect, distal portion formed from the

frontal calcification of the succeeding zooid, with a narrow, parallel-sided

median fissure proximally. Avicularium situated immediately proximal to

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 61

Fig. 16. A-B. Smittoidea calcarata sp. nov. A. Two zooids from close to the growing edge. B. An ovicelled

zooid. C—D. Parasmittina tropica (Waters). C. Zooids with entire peristomes, showing different types of

avicularia. D. Older zooids, with thickened peristomes. E-H. Parasmittina novella sp. nov. E. A group of

_zooids, with different types of avicularia. F. A zooid bearing an enlarged avicularium with flared serrate

rostrum. G. Zooids with smaller avicularia, with serrate rostra. H. Outline diagram of primary orifice.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

peristomial fissure, cystid low, rostrum elliptical, directed proximally, mandible

semicircular. Frontal calcification thin, fine grained and sutured, with a single

series of marginal pores. Ovicell oval, depressed frontally, developing a slight

lip continuous with the zooid orifice; surface finely granular, with scattered,

small, irregular pores.

Etymology

Calcar (L.)—a spur, referring to the sharp condyles within the primary

orifice.

Remarks

A single large colony, measuring approximately 20 mm X 10 mm encrust-

ing Dimorphocella, was collected from station SM 233.

Measurements (means of 15 values) in mm

1EZ Iz

1205 0,79

Parasmittina Osburn, 1952

Parasmittina Osburn, 1952: 411. Hayward & Ryland, 1979: 114.

Parasmittina tropica (Waters, 1909)

Fig. 16C—D

Smittia tropica Waters, 1909: 174, pl. 17 (figs 10-14).

Smittina tropica: Harmer, 1957: 934, pl. 64 (figs 23-28).

Parasmittina tropica: Cook, 1968: 215.

Material

Stations SM 163, SM 163/164, SM 164, SM 184, SM 185, SM 239.

Description

Colony encrusting. Zooids elongate, oval or irregular, rather flat. Primary

orifice wider than long, a prominent quadrate lyrula occupying most of the

proximal border, tapered distally to a greater or lesser extent; condyles large

and conspicuous. Four short distal oral spines present in newly budded zooids.

Peristome developing early in ontogeny, encircling primary orifice, and

extending on to frontal surface of ovicell in fertile zooids; tubular, its edge

variably lobed or produced. Proximomedially, the inner edge of the peristome

develops a pair of longitudinal ridges delimiting a central channel, which in

damaged specimens gives the illusion of a sinuate orifice. Frontal wall nodular,

with a single series of large and distinct marginal pores. Adventitious

avicularium arising laterally at base of peristome, directed proximally, laterally

or medially, typically single, occasionally, paired; rostrum slender, elongate, up

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 63

to 0,15 mm long. Ovicell spherical, with numerous small pores; recumbent on

succeeding zooid and eventually deriving an ooecial cover from it.

Remarks

Parasmittina tropica is readily distinguished from other species of this

genus by the form of the peristome and by the characteristically slender

avicularia. Some specimens included zooids bearing a second type of avicula-

rium, broader proximally than the usual type (Fig. 16C), but as the latter

frequently occurred on the same zooids also, this variation does not seem to be

significant. The lyrula also varied greatly in width, but again such variation was

observed within single colonies. In the structure of the peristome and the

presence of slender parallel-sided avicularia all the numerous specimens here

ascribed to P. tropica are closely similar.

Distribution

Described originally from the Red Sea coast of the Sudan (Waters 1909),

this species has subsequently been reported from west Africa (Cook 1968) and

the Mediterranean (Hayward 1974), and was accorded a wide distribution in

the Indo-West-Pacific region by Harmer (1957). Soule & Soule (1973) have

shown that some of Harmer’s material comprises a distinct species, P. serrula

Soule & Soule, and, consequently, the distribution of P. tropica may be more

restricted than Harmer supposed.

Measurements (means of 20 values) in mm

Lz Iz

0,54 0,28

Parasmittina novella sp. nov.

Fig. 16E-H

Material

Holotype: SAM—A26430, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Description

Colony encrusting, multilaminar. Zooids oval, rectangular or irregular,

convex, separated by shallow grooves. Primary orifice wider than long, lyrula

short, anvil-shaped, occupying half width of proximal edge, condyles blunt,

broad and conspicuous; two to four short, evanescent, distal oral spines, bases

occluded by development of peristome. Distal portion of peristome developed

from frontal calcification of succeeding zooid; prominent laterally and proxi-

mally, thin and erect, with a deep, parallel-sided medioproximal fissure;

extending on to frontal surface of ovicell in fertile zooids, forming a complete

ring. Free edge of peristome even or lobed. Frontal calcification at first smooth

and hyaline, with a single series of small inconspicuous marginal pores; later

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

thicker and rather nodular. Avicularia lateral or proximolateral to peristome,

single, paired or occasionally tripled, directed proximally; rostrum slender,

parallel-sided and straight, or gently curved, up to 0,17 mm long, palate with a

triangular foramen, pivotal bar thin, without a columella. In many, but not all,

instances the distal edges of the rostrum are slightly flared and distinctly serrate

(Fig. 16G). A larger avicularium may be developed on one side of the zooid,

with an elongate, proximally directed rostrum up to 0,4 mm long; the rostrum

is variably flared distally with fine or coarse serrations. Ovicell small, spherical,

recumbent on distally succeeding zooid, surface finely granular, with numerous

small pores.

Etymology

Novus (L.)—new.

Remarks

Parasmittina novella differs most markedly from P. tropica (Waters) and

from the eastern Pacific P. serrula Soule & Soule in the size of its zooids; for

the latter species Soule & Soule (1973) gave mean dimensions of Lz 0,368 mm,

Iz 0,284 mm, while the zooids of P. novella were 0,6-1,0mm long and

0,38-0,6 mm wide. Although the two species are similar in possessing avicularia

with flared serrate rostra, those of P. serrula seem to be less variable in all

respects than those of P. novella. Both species resemble P. tropica in the form

of the peristome, but in P. novella this is a far more delicate structure, lacking

the proximal longitudinal thickening and sinuate appearance of Waters’s

species.

Measurements (means of 40 values) in mm

Lz Iz

0,74 0,44

Porella Gray, 1848

Porella Gray, 1848: 127, 148. Hayward & Ryland, 1979: 116.

Porella capensis O'Donoghue, 1924

Fig. 17A-B

Porella capensis O’Donoghue, 1924: 45, pl. 2 (fig. 14).

Material

Stations SM 163, SM 164, SM 239.

Remarks

Several small colonies of this species were found, occurring as irregular

encrusting patches on organic calcareous substrata. The characteristic peri-

stome clearly develops, as O’Donoghue (1924) described, as separate elements,

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 65

Fig. 17. A-B. Porella capensis O'Donoghue. A. Zooids with typical development of peristome; note the

lyrula. B. The ancestrula. C. Arthropoma circinatum (MacGillivray). D. Arthropoma sp. E-F. Escharella

discors sp. nov. E. Part of a colony, cleaned to show the lyrula below the peristomial denticle. F. Outline

diagram of primary orifice. G. Escharina waiparaensis Brown. H. Calyptotheca nivea (Busk). I. Emballo-

theca ambigua sp. nov. J—K. Calyptotheca porelliformis (Waters). J. Two young zooids. K. Two ovicelled

zooids. Scale = 0,5 mm for A-H, J—K; 1 mm for I.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

a broad flared proximolateral portion and two distolateral portions. Small

adventitious avicularia may occur on the distolateral parts of the peristome rim,

but the specimens were too worn or damaged for further details to be observed.

The lyrula, not described by O’Donoghue, is quadrate and quite distinct.

Family Escharellidae Levinsen, 1909

Escharellidae Levinsen, 1909: 314. Hayward & Ryland, 1979: 136.

Escharella Gray, 1848

Escharella Gray, 1848: 125, 148. Hayward & Ryland, 1979: 136.

Escharella discors sp. nov.

Fig. 17E-F

Material

Holotype: SAM—A26431, station SM 163/164, 33°04,6’S 28°06,6’E, 90 m.

Description

Colony encrusting, forming small irregular white patches. Zooids small,

oval, convex, separated by deep grooves. Primary orifice wider than long,

proximal border with a broad anvil-shaped lyrula occupying the whole of its width;

encircled by a tall, thickened, cylindrical peristome, with eight slender spines

disposed around its distal and lateral edges. Proximal edge of peristome peaked

medially and bearing a bifurcate denticle on its inner face. Frontal wall thickly

calcified, finely granular, marginal pores small and rather conspicuous. Ovicell

spherical, tilted basally, the oval orifice opening into the top of the peristome.

Etymology

Discors (L.)—different.

Remarks

The morphology of the primary orifice and the number of oral spines are

the most useful characters for distinguishing the different species of Escharella.

E. discors has a further distinction in the large and conspicuous peristomial

denticle, which in uncleaned specimens may be confused with the lyrula.

Measurements (means of 20 values) in mm

ez Iz

Ori 0,31

Family Petraliellidae Harmer, 1957

Petraliellidae Harmer, 1957: 692.

The family includes a closely related group of genera, all of which are

characterized by zooids with large orifices, and frontal shields with pseudopores

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 67

and marginal septula. Multiporous septula in the basal walls give rise to

anchoring rhizoids; large distinctive, hyperstomial ovicells occur (see Cook &

Chimonides 1981a).

Mucropetraliella Stach, 1936

Mucropetraliella Stach, 1936: 363, 372. Harmer, 1957: 709. Cook & Chimonides, 1981a: 118.

Characterized principally by the presence of a suboral complex consisting

of a central lyrula and a mucro associated with an avicularium.

Mucropetraliella asymmetrica sp. nov.

Fig. 18

Material

Holotype: SAM—A26432, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Other material: stations SM 163, SM 164, SM 179, SM 180, SM 185.

Description

Colonies unilaminar, loosely encrusting, anchored by rhizoids originating

from basal pore plates. Primary orifice wide, with two to three spines. Lyrula

short and wide, sometimes extended laterally; lateral denticles pointing

proximally. Lateral sinuses unequal and asymmetrical, the larger one occurring

basally to the rostrum of the avicularium. Suboral mucro long and very stout;

proximal part of the orifice and the frontal shield considerably raised. Suboral

avicularium small, completely hidden at the base of the mucro, which is

directed frontally and distally; rostrum curved, directed laterally, palate vertical

to frontal plane, mandible rounded, hinged on a stout curved bar and directed

medially. One or two pairs of small lateral oral avicularia present, also with

rounded mandibles, directed laterally at an acute angle to frontal plane. In a

few zooids other small rounded avicularia occur at the base of the mucro; one

of the oral avicularia may be enlarged, with a conspicuous swollen cystid, the

rostrum raised, with a serrated edge, elongate and slightly spatulate, often

orientated proximally. Ovicells rather elongated, finely tuberculate, with one or

two small frontal avicularia.

Etymology

Asymmetros (Gr.)—without symmetry, referring to the suboral sinuses.

Remarks

The lyrula and suboral avicularium are difficult to see as they are situated

vertically below the mucro, which may reach a length of 1,2 mm and frequently

obscures the orifice completely. M. asymmetrica closely resembles M. watersi

Harmer (1957: 721, pl. 46 (fig. 9), fig. 67, from the East Indies), which was

introduced for Petralia vultur var. armata Waters (1913: 518, pl. 70 (figs 15-20),

from east Africa). Harmer noted that synonymy of the two species might be

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Mucropetraliella asymmetrica sp. nov. A.A group of

zooids in frontal view to show primary orifice and lateral

avicularia. x 39. B. Fragment of a colony in distal view, showing

elevation of the mucrones and associated avicularia. x 30. C. A

developing ovicell. x 61.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 69

doubted as Waters did not mention oral spines in his description. Comparison

of the Siboga specimens described by Harmer (1957) with one from the coast of

Kenya (BMNH 1975.4.16.5) shows them to be closely similar. The zooids of M.

watersi are less robust than those of M. asymmetrica and somewhat smaller

(Lz 0,85-1,0 mm, Iz 0,55—0,65 mm, lor 0,24-0,31 mm); the primary orifice has

four to six spines and symmetrical, equally developed lateral sinuses. The

proximal part of the orifice is not greatly raised and the suboral complex is

easily visible. Enlarged laterally or proximally directed avicularia are common

and ovicells are generally as wide or wider than long (cf. Harmer 1957).

Although most of the specimens of M. asymmetrica are fragmentary and

often worn, one colony from station SM 239, and another very young colony

(with Micropora similis) on a lamellibranch shell from station SM 164 were

alive when collected.

Measurements in mm

(range of 10 values)

Lz Iz lor

0,85-1,30 0,60-0,90 0,30-0,42

(means of 3 values)

Lov lov

0,55 OS

Family Schizoporellidae Jullien, 1883

Schizoporellidae Jullien, 1883: 527. Hayward & Ryland, 1979: 166.

Arthropoma Levinsen, 1909

Arthropoma Levinsen, 1909: 332. Hayward & Ryland, 1979: 190.

Arthropoma cecilii (Audouin, 1826)

Flustra cecilii Audouin, 1826: 239, pl. 8 (fig. 3).

Schizoporella cecilii: O'Donoghue, 1924: 42.

Arthropoma cecilii: Hayward & Ryland, 1979: 190, fig. 79.

Material

Station SM 185.

Remarks

A single live colony of this species was found. A cecilii has a wide

distribution in temperate, subtropical and tropical waters of the Atlantic,

Indian and western Pacific oceans.

Arthropoma circinatum (MacGillivray, 1869)

Fig. 17C

Lepralia circinata MacGillivray, 1869: 134.

Arthropoma circinatum: Harmer, 1957: 1003, pl. 72 (figs 29-30). Powell, 1967: 256, text-figs

29-30.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

Station SM 163.

Description

Colony encrusting. Zooids hexagonal, convex, separated by deep grooves;

0,46-0,5 mm long by 0,34-0,4 mm broad. Primary orifice D-shaped, proximal

border straight or slightly concave, with a short, quadrate median sinus. Six

oral spines present. Frontal wall finely granular, with distinct marginal pores,

extending towards the midline of the zooid proximolateral to orifice; elsewhere,

minute scattered pores are occasionally visible in the frontal calcification.

Peristome forming a slender crescentic lip proximal to the sinus, variably

developed. Ovicell hyperstomial, prominent, spherical; surface finely granular,

imperforate.

Remarks

A single dead colony was found, few of the zooids of which were

completely undamaged. The morphological features of this specimen were very

similar to those of specimens from New Zealand (BMNH 1899.5.1.1017) and

Bass Strait (BMNH 1899.5.1.1018), although the New Zealand specimen has

avicularia identical to those figured by Powell (1967, fig. 30) for ‘A. circinatum

form B’. No complete avicularia were found in the Meiring Naude specimen,

but a damaged zooid at the periphery of the colony bore a structure on the

frontal wall which seemed to represent a developing ‘form B’ avicularium.

Distribution

Arthropoma circinatum seems to be widespread in the western Pacific,

from New Zealand to Japan (Harmer 1957; Powell 1967), and has been

reported from Tristan da Cunha (Busk 1884) and southern California (Osburn

1952). The record from Ceylon listed by Harmer (1957: 1004) is based on a

specimen from the collection of L. R. Thornely that, on examination, proved

to be a species of Schizomavella.

Arthropoma sp.

Fig. 17D

Material

Stations SM 163, SM 185.

Remarks

Very small colonies of this species were recovered from two stations.

Although superficially similar to A. cecilii, the zooids differed in possessing five

or six distal oral spines, and a smaller orifice with a more slender sinus. However,

the material was fragmentary and none of the colonies represented a complete

astogenetic series. Consequently, although its affinities with Arthropoma seem

clear, this species must remain unnamed until further material is collected.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Tak

Escharina Milne Edwards, 1836

Escharina Milne Edwards, 1836: 230. Hayward & Ryland, 1979: 192.

Escharina pesanseris (Smitt, 1873)

Hippothoa pesanseris Smitt, 1873: 43, 76, pl. 7 (figs 159-160).

Escharina pesanseris: Harmer, 1957: 998, pl. 67 (figs 12-14, 18-19).

Material

Station SM 250.

Remarks

A single live colony of this distinctive species was collected; it is

recognized by the characteristic duck-foot shape of the avicularian mandible

(Harmer, 1957: pl. 67 (fig. 12)). Escharina pesanseris has been reported from

numerous localities in the Caribbean, the subtropical-tropical Atlantic, and the

Indo-West-Pacific region. It is known from Ceylon and Madagascar but does

not seem to have been recorded before from South Africa.

Escharina waiparaensis Brown, 1952

Fig. 17G

Escharina waiparaensis Brown, 1952: 229, figs 163-165. Powell, 1967: 275, pl. 6 (fig. a),

text-fig. 45.

Material

Stations SM 164, SM 239.

Description

Colony encrusting. Zooids broad and flat, rounded distally, separated by

raised, compressed sutures. Primary orifice wider than long; anter semiorbicu-

lar, with a slightly convex proximal border, poster slender, slit-like, broadening

proximally. Four short distal oral spines present. Frontal wall coarsely granular,

perforated by numerous closely spaced, minute pores. Avicularia paired, lateral

to orifice, level with the sinus; cystid small and rounded, supporting an

elongate, distally directed, setiform mandible. Ovicells were not found.

Numerous small basal pore chambers present.

Remarks

E. waiparaensis was described by Brown (1952) from the Miocene and

Pliocene of New Zealand; recent specimens were reported by Powell (1967)

from Three Kings Islands, northern New Zealand. Brown’s material lacked

spines, but Powell described two distal oral spines and two lateral ‘eminences’,

possibly representing thickened spine bases. The Meiring Naude material,

which included several living colonies, had four lightly calcified oral spines in

the youngest zooids, but the ontogenetically earliest zooids lacked spines

TP ANNALS OF THE SOUTH AFRICAN MUSEUM

altogether. In all respects the present material was identical with Powell’s

specimens from Three Kings Islands (BMNH 1964.8.12.59C, 59D).

Measurements (means of 20 values) in mm

lez Iz

ORAZ 0,63

Calyptotheca Harmer, 1957

Calyptotheca Harmer, 1957: 1008.

Calyptotheca nivea (Busk, 1884)

Fig. 17H

Schizoporella nivea Busk, 1884: 163, pl. 17 (fig. 1). Marcus, 1922: 25, fig. 15.

Schizoporella tenuis: O'Donoghue & De Watteville, 1935: 214.

Emballotheca nivea: O’Donoghue & De Watteville, 1944: 424. O’ Donoghue, 1957: 87.

Material

Stations SM 163, SM 163/164, SM 164, SM 179, SM 185, SM 232.

Description

Colony encrusting, multilaminar. Zooids quadrate, broad, flat or slightly

convex, separated by distinct raised sutures; 0,5—0,8 mm long by 0,38—0,52 mm

broad. Primary orifice subterminal, orbicular, as wide as long, or slightly wider

than long; poster forming a shallow U-shaped sinus below conspicuous

quadrate lateral condyles. Frontal wall regularly and closely perforated by

numerous round pores, each in a shallow pit, giving a rugose appearance.

Avicularia adventitious, small, typically paired, situated lateral to orifice, close

to suture of zooid; mandible semi-elliptical, short, variably orientated. Ovicell

hyperstomial, closed by zooidal operculum; oval, flattened frontally, regularly

perforated and typically crossed by sutures. Orifice of ovicelled zooid broader

than that of non-fertile zooid, with a wider and more shallow sinus.

The shape of the primary orifice showed some variation between the

different specimens, and earlier astogenetic stages tended to have a more

narrow sinus. In the colony from station SM 185 the zooids had developed

short cylindrical umbones proximal to the orifice, like those of the specimen

figured by Marcus (1922). The small avicularia are often very numerous, but

tend to be distributed along the interzooidal sutures.

Distribution

Calyptotheca nivea seems to be known only from the southern and eastern

coasts of South Africa. Reports of its occurrence elsewhere in the Indo-West-

Pacific region have been shown by Harmer (1957) to be attributable to other

species of Calyptotheca.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 73

Calyptotheca porelliformis (Waters, 1918)

Fig. 17J-K

Schizoporella porelliformis Waters, 1918: 15 (footnote), pl. 2 (figs 19-21).

Calyptotheca porelliformis: Harmer, 1957: 1008, 1020.

Material

Stations SM 163, SM 185.

Description

Colony encrusting, multilaminar. Zooids broad, quadrate, flat or slightly

convex, separated by distinct raised sutures; 0,56-0,76 mm long by 0,4-0,6 mm

broad. Primary orifice as broad as long, anter and poster of equivalent length;

anter D-shaped, poster shallowly concave, condyles prominent, blunt and

downcurved. Frontal wall rugose, regularly and closely punctured by numerous

round pores, each in a pit; a low, nodular ridge develops around proximal half

of orifice. Avicularia adventitious, minute, oval, with semi-elliptical mandible;

infrequent, situated lateral to orifice, close to sutures, single or paired, but

often absent altogether. Ovicell recumbent on succeeding zooid, hyperstomial,

closed by zooidal operculum; oval, flat frontally, closely punctured and very

rugose. Anter of ovicelled zooids much shorter than that of non-ovicelled

zooids.

Distribution

This species does not seem to have been reported since its original

description by Waters (1918) from Port Elizabeth.

Emballotheca Levinsen, 1909

Emballotheca Levinsen, 1909: 89, 333. Harmer, 1957: 1086.

Emballotheca ambigua sp. nov.

Fig. 171

Material

Holotype: SAM-A26433, station SM 250, 31°59,3’S 29°22,5’E, 150-200 m.

Description

Colony erect, dichotomously branching; branches cylindrical, somewhat

uneven or curved, about 2,5 mm thick, composed of whorls of four zooids.

Zooids large and broad, convex, separated by faint sutures. Frontal calcifica-

tion thick, finely granular, with numerous small closely spaced pores and a

single series of more distinct marginal pores. A conspicuous thick light brown

epitheca present. Primary orifice broader than long; anter approximately

semicircular, poster forming a shallow semi-elliptical trough between short

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

blunt lateral condyles. In fully developed, undamaged zooids the condyles

appear twinned (Fig. 171). Ovicell very large, immersed and scarcely protrud-

ing from branch surface, obscuring most of the distally succeeding zooid, but

not disturbing the four-whorled arrangement; orifice of ovicelled zooid

enlarged, more nearly quadrate, with the poster considerably broadened.

Etymology

Ambiguus (L.)—uncertain, referring to the systematic placing of the

species.

Remarks

The material comprised two fragments, each 15 mm long; both were living

when collected and perhaps represent parts of a single colony. The very large

ovicells, dimorphic orifices and prominent condyles of this species suggest it is

allied to the species of Emballotheca described by Harmer (1957). However, it

should be noted that the distinction between this genus and Calyptotheca

Harmer is unclear (see Dumont 1981), and the systematic placing of E.

ambigua must, therefore, be regarded as tentative.

Measurements (means of 20 values) in mm

EZ Lor lor

1,42 0,35 0,36

Family Stomachetosellidae Canu & Bassler, 1917

Stomachetosellidae Canu & Bassler, 1917: 44. Hayward & Ryland, 1979: 208.

Stomachetosella Canu & Bassler, 1917

Stomachetosella Canu & Bassler, 1917: 45. Hayward & Ryland, 1979: 208.

Stomachetosella balani (O’Donoghue & De Watteville, 1944)

Schizoporella balani O’Donoghue & De Watteville, 1944: 426, pl. 16 (figs 15-16).

Stomachetosella balani: O’Donoghue, 1957: 87.

Schizoporella balani: Hayward, 1980: 705, fig. 3G.

Material

Station SM 185.

Remarks

This characteristic South African species was recently redescribed and

figured by Hayward (1980). Its affinities with the boreal Atlantic species

currently assigned to Stomachetosella seem doubtful, and the relationship of all

Recent species to the type species, S. crassicollis Canu & Bassler, requires

re-examination. S. balani appears more similar to the Philippine species,

Schizoporella perforata Canu & Bassler (1929: 318), than to other species of

Stomachetosella.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 75

Family Cleidochasmatidae Cheetham & Sandberg, 1964

Cleidochasmatidae Cheetham & Sandberg, 1964: 1032.

Systematic problems involving species within this family are reviewed on

p. 104.

Cleidochasma Harmer, 1957

Cleidochasma Harmer, 1957: 1032. Cook, 1964b: 11.

Cleidochasma porcellanum (Busk, 1860)

Lepralia porcellana Busk, 1860: 283, pl. 31 (fig. 3).

Cleidochasma porcellanum: Cook, 1964b: 11, pl. 1 (fig. 4), pl. 2 (figs 1-2), fig. 4A-E.

Material

Station SM 163.

Description

Colony encrusting, zooids with imperforate, semi-transparent porcellan-

ous frontal shields and four small marginal septula. Orifice rounded distally,

with a small rounded sinus delineated by large proximally directed condyles.

Avicularia small, often paired, lateral and suboral, mandible rounded or

subtriangular, directed laterally; if not present, avicularium replaced by one of

the marginal septula. Ovicells hyaline and prominent at first, hyperstomial, not

closed by the operculum, becoming partially immersed. Proximal edge of

ovicell with paired lateral indentations and an area of thin calcification.

Remarks

A single, fairly large colony, comprising approximately 1250 zooids, was

found. It was alive when collected, and includes a well-preserved growing edge.

The earlier stages of colony growth are obscured by small groups of frontally

budded zooids. The orifice is more elongated than those of east African

specimens (BMNH 1976.7.20.7, Wasin, Zanzibar) and of the west African

specimens illustrated by Cook (19645).

C. porcellanum is a widely distributed and very variable species-complex

which includes C. bassleri (Calvet), as described by Harmer (1957). Records

are circumtropical and subtropical, with a depth range of 1-220 m.

Cleidochasma protrusum (Thornely, 1905)

Gemellipora protrusa Thornely, 1905: 119, pl. 7.

Cleidochasma protrusum: Harmer, 1957: 1040, pl. 71 (figs 1-4), fig. 112. Hayward & Cook,

1979389:

Material

Station SM 131.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

A single, small dead colony was found. The characteristic morphotype of

the South African populations of C. protrusum was noted by Hayward & Cook

(1979).

Cleidochasma cribritheca (Busk, 1884)

Fig. 19A

Gemellipora cribritheca Busk, 1884: 176, pl. 33 (fig. 5).

Material

Stations SM 163, SM 163/164, SM 164.

Description

Colony encrusting, unilaminar, appearing white and porcellanous. Zooids

oval to hexagonal, convex, separated by shallow grooves; 0,4—-0,6 mm long by

0,2-0,3 mm broad. Primary orifice longer than wide, anter suborbicular, poster

elongate, V-shaped; condyles prominent, proximally directed. Orifice becoming

immersed as calcification thickens, but without a defined peristome; no oral

spines. Frontal wall at first smooth, with numerous small frontal pores;

becoming more rugose in later ontogenetic stages, with pores sunk in small pits.

Avicularium single, proximal to sinus, on a small tumid cystid; mandible

semi-elliptical or semicircular, directed proximally. Ovicell recumbent on

succeeding zooid, oval, flattened frontally and perforated by numerous small

pores; later enveloped distally and laterally by a thickened ooecial cover.

A colony from SM 163 (Fig. 19A) included early astogenetic stages. These

were partly obscured by later zooids but showed clearly two small zooids, each

with numerous small oral spines. These may represent a twinned ancestrula;

alternatively, the ancestrula may be obscured by the two largest zooids and the

small zooids may be simply the first to be budded from the ancestrula.

Distribution

This species is known only from South African waters.

Cleidochasma perspicua sp. nov.

Fig. 19B

Material

Holotype: SAM—-A26434, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Description

Colony encrusting. Zooids hexagonal, convex, separated by shallow

grooves. Primary orifice cleithridiate: anter orbicular, poster wider than long,

V-shaped, condyles large and distinct, blunt, basally deflected. Peristome

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES ia

Fig. 19. A. Cleidochasma cribritheca (Busk). B. Cleidochasma perspicua sp. nov. C-D. Hippoporella

spinigera (Philipps). C. Portion of a young colony. D. Outline diagram of primary orifice. E-F. Hippo-

porella labiata sp. nov. E. Portion of a young colony. F. Outline diagram of primary orifice.

G. Hippomenella avicularis (Livingstone). H. Celleporaria tridenticulata (Busk).

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

developed as a low rim distally and laterally, frequently with two or more short

umbones laterally; a third small umbo may be developed proximal to the

orifice. Frontal wall thick, vitreous, smooth, with very few sparsely distributed

and indistinct marginal pores; thick, translucent secondary calcification infills

the grooves between zooids in later ontogenetic stages, pore openings migrate

medially as this encroaches and the passage of the pore appears tubular

(Fig. 19B). Avicularium adventitious, single, proximolateral to orifice; cystid

tumid, rostrum slender, acute triangular, directed obliquely laterally, palate

with a triangular foramen, cross-bar stout with a thickened, quadrate

columella. The avicularia are sporadically developed, frequently missing from

many zooids. Ovicell recumbent on succeeding zooid, as wide as long, flattened

frontally and with a distinct labellum; calcification smooth and imperforate,

developing a small median umbo. Large basal pore chambers present.

Etymology

Perspicuus (L.)—transparent, an allusion to the frontal calcification.

Remarks

The holotype comprises a single unilaminar colony, 5 mm” in area. It was

alive when collected and many of the ovicells contained embryos. C. perspicua

is most similar to C. porcellanum (Busk), which appears to have a circumtropi-

cal distribution. In particular, the primary orifice of this species is very similar

to that of a variant of C. porcellanum from Ceylon, illustrated by Cook (19645,

text-fig. 4). However, the large tumid avicularian cystids and the conspicuous

frontal labellum of the ovicell are sufficient to distinguish C. perspicua from

other species of Cleidochasma. For further discussion of this species see p. 104.

Measurements (means of 20 values) in mm

Lz Iz

0,45 0,4

Hippoporidra Canu & Bassler, 1927

Hippoporidra Canu & Bassler 1927: 21, 31. Cook, 1964b: 22. Taylor & Cook, 1981: 244.

Hippoporidra senegambiensis (Carter, 1882)

Cellepora senegambiensis Carter, 1882: 416, pl. 16 (fig. 1A—-V).

Hippoporidra senegambiensis: Cook, 1964b: 29, pl. 3 (figs 3-4), figs 7B—C, 8A—D; 1968: 196,

4pl. 8 (fig. C).

Material

Station SM 185, one young colony on Turritigera shell.

Description

Colonies encrusting gastropod shells, usually those inhabited by pagurid

crabs. Zooids multilamellar, produced by frontal budding, with two to four

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 79

series of frontal marginal septula and orifices with rounded sinus and distinct

condyles. Colony becoming mamillate by budding of groups of prominent,

large (?male) zooids with small orifices and tuberculate peristomes. Inter-

zooidal and adventitious avicularia triangular or rounded. Ovicelled zooids

occurring in hollows among the mamillae, ovicells with a small frontal area.

Remarks

H. senegambiensis is common in west African waters, to a depth of 100 m.

The single specimen found, which encrusts a small Turritigera shell 10 mm in

length, is a young colony without ovicells that was alive when collected.

Another species of Hippoporidra, H. picardi, has been reported from South

Africa from depths of 45 to 200m. H. picardi has larger, more recumbent

zooids, with orifices having a small rounded sinus.

Hippoporella Canu, 1917

Hippoporella Canu, 1917: 36. Harmer, 1957: 1096. Hayward & Ryland, 1979: 218.

Hippoporella spinigera (Philipps, 1899)

Fig. 19C-D

Escharoides spinigera Philipps, 1899: 440, 448, pl. 43 (fig. 12).

Hippoporella spinigera: Harmer, 1957: 1100, pl. 73 (fig. 13).

Mucronella serratilabris O’Donoghue, 1924: 48, pl. 3 (fig. 18).

Material

Stations SM 131, SM 151, SM 163, SM 164, SM 250.

Description

Colony encrusting, forming small circular, silvery patches. Zooids oval to

hexagonal, small and rather broad; separated by shallow grooves, later

obscured by calcification. Primary orifice broader than long, appearing rather

quadrate; distal edge arched, proximal edge almost straight, blunt lateral

condyles present. Seven long, slender, closely grouped distal oral spines.

Peristome developed proximally as a blunt, prominent mucro, with a broad,

quadrate, finely serrated lip on its distal edge. Adventitious avicularia paired,

lateral to orifice; mandible acute triangular, directed laterally or distolaterally.

Additional avicularia present on frontal wall: most frequently elongate, situated

on midline of zooid, with slender triangular mandible directed proximally;

more rarely, one or two avicularia, similar to oral type but slightly larger,

situated elsewhere on frontal wall, with variable orientation. Frontal wall

smooth, imperforate except for inconspicuous marginal pores. Basal pore

chambers present. Ovicell hyperstomial, recumbent on succeeding zooid,

spherical, with a large quadrate orifice, often developing a small umbo.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

H. spinigera is characterized by the small size of its zooids, the distinctive

peristomial mucro and the closely grouped slender oral spines. Originally

described fom the Loyalty Islands (Philipps 1899), it has since been reported

from Hong Kong, the Philippines, and Sumbawa (Harmer 1957), and a single

specimen was recorded from South Africa by O’Donoghue (1924), as

Mucronella serratilabris. Some Australian Tertiary species such as Mucronella

mooraboolensis MacGillivray (1895: 100, pl. 13 (fig. 9)) and Rhynchopora

spinifera MacGillivray (1895: 102, pl. 13 (fig. 19)) may be related to H.

spinigera.

Measurements (means of 10 values) in mm

Lz Iz

0,32 O27)

Hippoporella labiata sp. nov.

Fig. 19E-F

Material

Holotype: SAM—A26435, station SM 250, 31°59,3’S 29°22,5’E, 150-200 m.

Description

Colony encrusting. Zooids hexagonal, convex, small; separated by distinct

grooves. Primary orifice lepralioid: anter longer than wide, with a finely

denticulate edge, proximal border of poster gently concave; prominent, blunt

lateral condyles present. Seven or eight short oral spines closely spaced around

distal border of aperture. Frontal wall thick, vitreous, with a nodulated surface,

imperforate except for indistinct marginal pores; a short cylindrical umbo

developing just proximal to orifice and arching towards it. Avicularia

adventitious, situated lateral to orifice, typically paired, mandible semi-

elliptical, directed obliquely laterally; often replaced on one or both sides by a

second type with an elongate, parallel-sided, distally-rounded mandible. Ovicell

hyperstomial, pyriform, imperforate, with a distinct frontal labellum; becoming

submerged in secondary calcification derived from distally succeeding zooids.

Small basal pore chambers present.

Etymology

Labiatus (L.)—lipped, referring to the prominent suboral umbo.

Remarks

This species has a remarkable superficial resemblance to H. multidentata

(Thornely), from Ceylon, described and figured by Harmer (1957: 1099,

pl. 73 (figs 9-12)). Comparison of the present material with Thornely’s type

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 81

specimen (BMNH 1906.12.3.4) shows several important differences. The orifice

of H. labiata is proportionately less broad than in H. multidentata; the ovicell is

longer and more distinctly pyriform than the rather squat ovicell of H.

multidentata. Finally, the avicularia of H. labiata are very much larger than

those of H. multidentata which, moreover, are consistently monomorphic. The

systematic status of this species is considered at greater length on p. 104.

Measurements (means of 20 values) in mm

Lz Iz

0,45 0,39

Hippomenella Canu & Bassler, 1917

Hippomenella Canu & Bassler, 1917: 41. Brown, 1949: 513.

Hippomenella avicularis (Livingstone, 1926)

Fig. 19G

Lepralia tuberculata var. avicularis Livingstone, 1926: 93, pl. 5 (figs 1-3).

Hippomenella spatulata Harmer, 1957: 1095, pl. 72 (figs 27, 31).

Material

Stations SM 131, SM 163.

Description

Colony encrusting. Zooids large, oval, convex, separated by deep

grooves. Primary orifice with a semi-orbicular anter separated from a narrower

U_ shaped poster by prominent, blunt, basally deflected condyles. Orifice rim

with six or seven spine bases. Frontal calcification thick and smooth, with

numerous, very small inconspicuous pores; these are distributed around the

periphery of the zooid, their frontal openings tend to shift centripetally as the

frontal calcification thickens (giving a striated appearance to the wall), but the

central area remains imperforate. Small adventitious avicularia present on all

zooids, lateral to orifice and typically paired, mandible semi-elliptical, disto-

laterally directed. Identical avicularia sporadically present frontally, up to five

on a single zooid, orientation variable but directed away from the central area

of the zooid. One or both of the lateral-oral pair may be replaced by an

elongate avicularium with a slender spoon-like mandible. Small basal pore

chambers present.

Remarks

This species was represented by several small dead colonies; ovicells were

present but were so badly damaged that their structure could not be discerned.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

H. avicularis has been rarely reported and appears to be known from just

three localities in the western Pacific (Harmer 1957).

Measurements (means of 16 values) in mm

Lz Iz

0,92 OZ

Family Microporellidae Hincks, 1879

Microporellidae Hinks, 1879: 156. Hayward & Ryland, 1979: 220.

Microporella Hincks, 1877

Microporella Hincks, 1877: 526. Hayward & Ryland, 1979: 220.

Microporella sp.

Fig. 20A

Material

Stations SM 163/164, SM 239.

Description

Colony encrusting. Zooids oval to hexagonal, broad, flat or slightly

convex, separated by shallow grooves. Primary orifice subterminal, semicir-

cular, with four or five short, distal oral spines. Frontal wall finely granular,

perforated by numerous small pores; ascopore situated just proximal to orifice

in distal third of zooid. Avicularium single, developed midway along the length

of the zooid; mandible slender, setiform, up to 0,2 mm long, directed obliquely

distally. No ovicells present.

Remarks

The familiar Microporella ciliata appears to have an almost cosmopolitan

distribution, but the degree of morphological variation which it displays over its

entire geographical range requires examination. The present material, consist-

ing of two small colonies, is similar to M. ciliata, but in comparison with

specimens from British waters appears rather larger, with more robust

avicularian mandibles. No complete astogenetic series was represented in the

Meiring Naude specimens, and neither colony possessed ovicells. consequently,

the material is left unnamed until better specimens should be collected.

Measurements (means of 20 values) in mm

eZ Iz

0,57 0,42

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 83

Fig. 20. A. Microporella sp. B-C. Flustramorpha marginata (Krauss). B. Ovicelled zooids. C. Young

zooids, showing shape of primary orifice. D. Flustramorpha flabellaris (Busk). E. Fenestrulina indigena sp.

nov., ancestrula and first two zooids. F. Flustramorpha angusta Hayward & Cook, young zooids showing

shape of primary orifice. G. Fenestrulina indigena sp. nov., a group of zooids from close to the colony edge.

H. Flustramorpha angusta Hayward & Cook, ovicelled zooids.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Flustramorpha Gray, 1872

Flustramorpha Gray, 1872: 168. Busk, 1884: 135.

All three species of Flustramorpha known from South Africa were present

in the Meiring Naude samples. The genus is a homogenous one and the three

species are often difficult to distinguish; it seems useful, therefore, to describe

briefly the features considered here to be of importance. F. flabellaris (Busk)

typically develops a tufted colony of short, broad lobes, but its most distinctive

feature is the avicularium that, unlike those of the other two species, has a

short acuminate mandible, and a palate that is almost parallel to the frontal

plane of the zooid. In F. marginata (Krauss) and F. angusta Hayward & Cook

the avicularium has a setiform mandible, and the palate is orientated at an

oblique, or even perpendicular, angle to the frontal plane.

The avicularian mandibles of F. angusta were not preserved in the type

specimen. They are similar to those of F. marginata, setiform and up to

0,75 mm long; they are broadened basally close to the point of articulation

(Fig. 20H). The ovicell of F. angusta is proportionately smaller, narrower and

more depressed than that of F. marginata, its width being considerably less

than that of the maternal zooid. Further, whereas in F. marginata the ovicell is

prominent at all stages of growth, in F. angusta it is progressively obscured by

secondary calcification which typically overhangs its aperture on each side. The

orifice of F. angusta is semi-elliptical, with a straight proximal border

(Fig. 20F), whereas that of F. marginata has distinct lateral condyles that

impart to it a characteristic outline at all ontogenetic stages (Fig. 20C).

Finally, it may be the case that the colony form of these latter two species

are a further useful specific character. The specimens of F. angusta were all

slender, unbranched, strap-like colonies, up to 50 mm long, with a maximum

width of 3,5 mm. The material of F. marginata was mostly fragmentary, but

included two regularly bifurcating colonies, 30 mm and 40 mm high, with a

maximum branch width of 4 mm.

Flustramorpha flabellaris (Busk, 1854)

Fig. 20D

Eschara flabellaris Busk, 1854: 91, pl. 107 (figs 7-10).

Microporella flabellaris: Marcus, 1922: 28, fig. 16.

Material

Stations SM 179, SM 180.

Flustramorpha marginata (Krauss, 1837)

Fig. 20B-C

Flustra marginata Krauss, 1837: 35, fig. 3.

Flustramorpha marginata: Busk, 1884: 135, pl. 20 (fig. 8). Hayward & Cook, 1979: 80.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Material

Stations SM 163/164, SM 184, SM 185, SM 239.

Flustramorpha angusta Hayward & Cook, 1979

Fig. 20F, H

Flustramorpha angusta Hayward & Cook, 1979: 80, fig. 11E.

Material

Stations SM 131, SM 163, SM 163/164, SM 184, SM 1835.

Fenestrulina Jullien, 1888

Fenestrulina Jullien, 1888: 37. Hayward & Ryland, 1979: 224.

Fenestrulina indigena sp. nov.

Fig. 20E-G

Material

Holotype: SAM-A26436, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: station SM 163/164.

Description

Colony encrusting. Zooids oval to hexagonal, convex, separated by distinct

grooves. Primary orifice semicircular, with a thin raised rim, relatively

small—constituting less than one-tenth of the total zooid length; six or seven

short distal oral spines present. Frontal wall thin and hyaline, with a few

radiating sutures; marginal pores distributed in a single series, small and rather

inconspicuous. Ascopore situated just proximal to orifice in the distal half of

the zooid, small, transversely oval; a short umbo develops proximal to the

ascopore but does not support it. Ovicell recumbent on succeeding zooid,

globular, as wide as long; smooth surfaced, with faint marginal flutings.

Ancestrula tatiform, oval, 0,32 mm long, with twelve marginal spines.

Etymology

Indigenus (L.)—native to.

Remarks

Fenestrulina indigena may be distinguished from other species of the genus

by the number of oral spines, and by the characteristic frontal umbo, which

does not seem to be in any way associated with the ascopore.

Measurements (means of 20 values) in mm

Lz Iz

0,60 0,43

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Hippothoidae Levinsen, 1909

Hippothoidae Levinsen, 1909: 274. Hayward & Ryland, 1979: 246.

Trypostega Levinsen, 1909

Trypostega Levinsen, 1909: 280. Hayward & Ryland, 1979: 258.

Trypostega venusta (Norman, 1864)

Lepralia venusta Norman, 1864: 84, pl. 10 (figs 2-3).

Trypostega venusta: Hayward & Ryland, 1979: 258, fig. 111.

Material

Stations SM 163, SM 163/164, SM 164, SM 239.

Distribution

Trypostega venusta has a circumtropical, warm-temperate distribution. It

has been reported from the Red Sea, Mauritius, and east Africa (Harmer 1957)

but has not been recorded before from South Africa.

Family Gigantoporidae Bassler, 1935

Gigantoporidae Bassler, 1935: 32. Harmer, 1957: 878.

Gigantopora Ridley, 1881.

Gigantopora Ridley, 1881: 47. Harmer, 1957: 879.

Gigantopora polymorpha (Busk, 1884)

Gephyrophora polymorpha Busk, 1884: 167, pl. 34 (fig. 2).

Adeonella ponticula O'Donoghue, 1924: 54, pl. 4 (fig. 23).

Gigantopora polymorpha: Brown, 1952: 208, figs 145-146. Hayward & Cook, 1979: 81.

Material

Stations SM 179, SM 180, SM 185.

Remarks

Numerous live colonies of this characteristically South African species

were collected at stations SM 179 and SM 185.

Gigantopora foraminosa sp. nov.

Fig. 11E—F

Material

Holotype: SAM-A26437, station SM 163/164, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 131, SM 163, SM 163/164, SM 164, SM 184.

Description

Colony erect, rigid, cylindrical, branching irregularly; attached by an

encrusting base, up to 21 mm high in present material, the branches broadening

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 87

distally. Zooids in alternating longitudinal series, disposed all around the branch

axis; rectangular, flat or slightly convex, separated by distinct raised sutures.

Primary orifice longer than wide; anter D-shaped, poster concave, constituting a

broad, shallow, U-shaped sinus between small lateral condyles. Frontal wall

closely perforated by numerous round pores, each in a distinct pit. Peristomial

complex occupying distal half of zooid; initiated by the development of paired

lateral oral avicularia with elongate, acute triangular rostra, directed medially.

These arch over the primary orifice and fuse medially; proximal to the area of

fusion, a transversely oval spiramen communicates with the space above the

primary orifice, distally a rim of calcification delimits a semicircular secondary

orifice. Border of secondary orifice frequently peaked medially on either or both

the distal and proximal edge. The gap between the two avicularian cystids is

infilled as the opening of the spiramen is elaborated; this develops a lobed distal

hood, above which two further lacunae are seen, separated medially by a

longitudinai ridge. In later ontogenetic stages these lacunae are completely

obliterated. Ovicell large, prominent but partially immersed; broadly oval and

densely punctured by small pores, becoming rather rugose. Secondary orifices of

ovicellate zooids broader than those of non-ovicellate zooids. Vertical walls of

zooids with large and distinct multiporous septula.

Etymology

Foraminosus (L.)—full of holes.

Remarks

This species is distinguished from G. polymorpha by its generally larger

dimensions, by its relatively longer primary orifice, and by the characteristic

twinned lacunae below the oral avicularia. A similar fossil species, Porina

cribraria, was described from the Australian Tertiary by MacGillivray

(1895: 104, pl. 14 (fig. 25)).

Measurements (mean values) in mm

Zooids Ovicell

n Lz Iz n Lov lov

25 1207 0,6 8 0,48 0,66

Family Adeonellidae Gregory, 1893

Adeonellidae Gregory, 1893: 241. Cook, 1973: 246.

Adeonella Busk, 1884

Adeonella Busk, 1884: 183. Cook, 1968: 180.

The genus Adeonella is abundantly represented in eastern South African

waters and a total of 15 species may now be recognized. A. regularis (Busk,

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

1884) seems to have been the first species to be described from this area; the

type specimen is no longer extant, however, and the status of this species

remains to be clarified. Similarly, A. pygmaea Levinsen (1909), the identity of

which even Levinsen was uncertain about, has not been reported since the

original, rather brief account. O’Donoghue (1924) described four new species

of Adeonella from South Africa; A. coralliformis, A. expansa, and A. ligulata

are well-defined species, the type specimens of which have been examined

during the preparation of this account. A. ponticula O’Donoghue (1924) has

proved to be identifiable with Gigantopora polymorpha (Busk) (Hayward &

Cook 1979); A. pectinata Busk, described from a single specimen collected by

the Challenger off Cape York, Queensland, was reported by O’Donoghue

(1924) from two South African localities but his specimens prove to represent

A. gibba sp. nov. Adeonella meandrina O’Donoghue & De Watteville (1944)

has been shown (Cook 1973) to be referable to the umbonuloid genus

Adeonellopsis, and A. jellyae Levinsen (1909) may be assigned to Laminopora

Michelin, 1842 (Cook 1983a).

The first Meiring Naude collections included two undescribed species of

Adeonella, A. majuscula and A. cracens (Hayward & Cook 1979). A. falcicula

and A. cultrata were then described (Hayward 1981) from South African

specimens collected by the Galathea deep-sea expedition. The present

collections yielded eight new species of Adeonella, and it would seem useful at

this stage to present a Key to the presently known South African species of the

genus. The morphological features of most use in distinguishing between the

different species are: the shape of the primary orifice, the shape and position of

the spiramen, the size, orientation, and position, relative to the secondary

orifice, of the peristomial avicularia. Certain later ontogenetic features seem to

be consistent in some species, and the form of the colony may prove also to be

an important character.

PROVISIONAL KEY TO THE SOUTH AFRICAN SPECIES OF ADEONELLA

—

Primary orifice orbicular, oval or quadrate, without asinus —..................... 2,

—) Pomary ontice withia distince Sinus a eee ere eee one err eene 6

2. Primary orifice longer than broad, quadrate. Spiramen broadly oval, developing a

proximal concavity in later ontogeny; flanked by one or two small medially directed

avicularia. Other avicularia rare. Marginal vicarious avicularia absent

A. infirmata sp. nov.

— Primary orifice oval or rounded. No concavity proximal to spiramen _—..................

3. Primary orifice almost circular. Spiramen oval, situated close to distal end of

peristome, flanked by small paired medially directed avicularia, a second pair often

present on distal rim of peristome. Marginal vicarious avicularia absent

A. falcicula Hayward (1981)

— Primary orifice transversely oval. Spiramen flanked by paired or single avicularia.

Enlarged vicarious avicularia present on colony margins ==———S—........... sss vee 4

4. Spiramen distinctly oblong, its primary outline contrasting with, and visible beneath,

the rounded opening developed in later ontogeny. Small paired avicularia developed

lateral to spiramen, extending distally towards, but not reaching the proximal border

of the secondary ontfiices) 1 ee ee A. cracens Hayward & Cook (1979)

— Spitamen round or ovalat all ontogenetic staces! eae renee ene eee 5

18).

14.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 89

Spiramen large, round; situated in the middle of the zooid, well proximal to the

secondary orifice. A single avicularium present in early ontogeny, elongate, extending

obliquely medially, between the spiramen and the secondary orifice. Smaller frontal

avicularia abundant in later ontogenetic stages ..... A. majuscula Hayward & Cook (1979)

Spiramen large, oval; situated in distal half of zooid, close to base of peristomial

calcification. A single frontal avicularium developed just proximal to spiramen,

extending obliquely distally towards, and almost reaching, the proximal border of the

secondary orifice. Other frontal avicularia apparently absent ... A. cultrata Hayward (1981)

Spiramen, in early ontogenetic stages, overarched from one side by a distinct hood, so

that the plane of its aperture is perpendicular to the frontal plane of the zooid. A

simalludistallyadimectediaviculaniumradjacentto spiramen) =.) 4.4 ee 7

Spiramen, in early ontogenetic stages, with a low distal arch, and frequently with a

distinct concavity proximal to it, or normal to frontal plane, but without a laterally

GeVelOpe Cah OO CBee cath ira thee daa th ace MHRNG ore GNOME RS ARO Ae os AY hint 9

Primary orifice as broad as, or broader than long. A single, median, proximally

directed avicularium developing proximal to, and partly obscuring, the spiramen in

later ontogeny, Colony storming slender blades = -.55 5.) ..4.... 5556.14: A. gibba sp. nov.

Primary orifice longer than broad. No median, proximally directed avicularium

Sinus of primary orifice forming a short symmetrical U. Small avicularia frequent on

frontal wall and around peristome in later ontogeny. Colony forming slender blades

A. abdita sp. nov.

Sinus of primary orifice forming a broad, shallow U. Small avicularia infrequent in

later ontogeny; typically one extending obliquely between spiramen and secondary

orifice, and one orientated transversely along proximal border of secondary spiramen.

Colonystonnimne broadhat platesy=n eas aae es] ae: A. expansa O’ Donoghue (1924)

. Spiramen large, oval or circular, with the sinus visible through it in at least the earliest

CMIOLUSDEWS STSS TO A ee tae eg aie earn Me car! ce eee ge ee Se 10

Spiramenismalls circular, sinus mot visible ase sna ese enone ne ene e eee 14

. Primary orifice with slender, pointed sinus. A single, median, proximally directed

avicularium developed proximal to the spiramen in later ontogeny. Colony forming

SOMO ORCEDIATC Si ieee Lit owe clan Packs, « soul a een eats Sis ae RR oh A. alia sp. nov.

Primary orifice with a V- or U-shaped sinus. Colony typically forming slender blades.. 11

. Sinus narrow, parallel-sided. Peristomial avicularia typically paired, arising distal to

spiramen and extending medially above distal border of secondary orifice

A. decipiens sp. nov.

Sinus forming a symmetrical, broader U-shape. Peristomial avicularia single or paired 12

Primary orifice broader than long. Peristomial avicularium single, short; situated

adjacent to spiramen, extending obliquely distally towards proximal border of

SeECONG AIVAOMIN CCH res te ee hess Gea wee hs We oon a4 Cees are to Sauamun aps een A. conspicua sp. nov.

Primary orifice as long as, or longer than broad. Peristomial avicularia typically paired 13

Peristomial avicularia paired, arising distal to spiramen and directed medially, the

apices of the rostra almost converging distal to the secondary orifice. Frontal wall

CONVEXoproximalitOrspiramMeny qs arssns 25s aes bs: A. coralliformis O'Donoghue (1924)

Peristomial avicularia paired, arising adjacent to, or just distal to the spiramen and

directed medially, extending between the spiramen and the proximal border of the

secondary orifice. Frontal wall concave proximal to spiramen; later filled by a

pLOxdnialllvadinectedraviCulaniUmMls yaa 4 cle oie at ksh oc ae ore A. confusanea sp. nov.

Primary orifice as broad as long, anter and poster of almost equal length. Spiramen

situated in distal half of zooid but distant from the proximal border of the secondary

©) Telli 6c MMR MP ME re URE ACMA cogs at ae or eae pened Mas Costbe os pes A. distincta sp. nov.

Primary orifice much broader than long; poster very shallow, constituting less than

one-third of orifice length. Spiramen small, situated close to proximal border of

SECONGARVAOTINCE Rasch. tn aa ahaa tha aises Gane dh Ss eh ates A. ligulata O’Donoghue (1924)

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

Adeonella majuscula Hayward & Cook, 1979

Adeonella majuscula Hayward & Cook, 1979: 82, fig. 1OE—-H.

Material

Station SM 233.

Remarks

Part of a single large colony, 75 mm high, was collected from a depth of

540-80 m.

Adeonella cracens Hayward & Cook, 1979

Adeonella cracens Hayward & Cook, 1979: 85, fig. 10I-L. Hayward, 1981: 44, Fig. 22A.

Material

Station SM 103.

Remarks

Dead fragments only were recovered from the above station.

Adeonella decipiens sp. nov.

Fig. 21A—-E

Material

Holotype: SAM-A26438, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 131, SM 163/164, SM 179, SM 180, SM 184,

SM 185.

Description

Colony erect, branching, rigid; branches bilaminar, slender, up to 5 mm

wide. Zooids oval to hexagonal, broadening distally, tapered proximally,

separated by indistinct grooves which are increasingly obscured in older zooids.

Primary orifice semi-orbicular, sinus slender, U-shaped; peristome thickened,

prominent, tending to project from branch surface; inner proximal edge with a

blunt projecting knob (Fig. 21B), later hidden as secondary calcification

progresses, secondary orifice eventually orbicular. Frontal wall convex, finely

granular, with numerous small pores, indistinct in newly developed zooids,

more pronounced in later ontogenetic stages. Spiramen oval and large, situated

at base of peristome so that calcification between it and the secondary orifice

forms merely a slender bridge in young zooids; despite further thickening, the

proximal border of the primary orifice is visible in all but the most heavily

calcified zooids. Adventitious avicularia paired, lateral to the orifice, rostrum

acute triangular, directed distomedially along the distal border of the secondary

orifice. In later ontogenetic stages these avicularia may be obliterated and

replaced by a second generation with a similar orientation, or more proximally

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 91

Fig. 21. A-E. Adeonella decipiens sp. nov. A. Four typical zooids. B. Zooids from a growing edge,

showing characteristic shape of early peristome. C. Older zooids with hooded spiramina. D. View of a

branch edge, showing vicarious and adventitious avicularia. E. Outline diagram of primary orifice.

F-J. Adeonella confusanea sp. nov. F. Young zooids, with concave frontal walls. G. Later zooids, with

thickened frontal walls. H. View of a branch edge, with vicarious avicularia. I. Zooids from near the base

of the colony, with secondarily developed frontal avicularia. J. Outline diagram of primary orifice.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

situated with the rostra extending between the spiramen and the proximal

border of the secondary orifice; rarely, the avicularia may be developed

frontally, proximal to the spiramen. Vicarious avicularia developed along the

branch edge, with enlarged, distally directed rostra, interspersed with what may

be small kenozooids, or autozooids with obliterated orifices; both these and the

vicarious avicularia may bear small adventitious avicularia.

Etymology

Decipia (L.)—deception, a reference to the similarity between this species

and A. coralliformis.

Remarks

At the distal tips of branches the peristomes and avicularia project

markedly from the surface. With continued calcification both are progressively

immersed, the spiramen becomes deeply sunk, more rounded and smaller. A.

decipiens is superficially similar to A. coralliformis O’Donoghue but is readily

distinguished by the small size of its zooids and by the primary orifice, which is

almost circular in A. coralliformis. The material included numerous living and

dead fragments, up to 22 mm long.

Measurements (means of 25 values) in mm

Lz Iz

0,50 0,36

Adeonella confusanea sp. nov.

Fig. 21F—J

Material

Holotype: SAM-A26439, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Other material: stations SM 163, SM 163/164, SM 179, SM 185.

Description

Colony erect, branching, rigid, attached by an encrusting base; branches

bilaminar, up to 39 mm long in present material, with a maximum width of

4mm. Zooids oval to rectangular, hexagonal in central part of branch, convex,

separated by deep grooves. Primary orifice semi-orbicular, with a short

U-shaped sinus; peristome with a broad blunt denticle within proximal border,

visible in earlier ontogenetic stages, developing a transversely oval secondary

orifice. Avicularia typically paired, occasionally single, arising lateral to

spiramen, rostrum acute triangular, directed distomedially between spiramen

and secondary orifice, close to the proximal border of the latter. Frontal wall

granular, convex and rather rugose, with numerous small pores; spiramen

circular, situated close to proximal base of peristome, sinus visible through it in

a i ee ee

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 23)

youngest zooids. Later in ontogeny, additional avicularia may be developed

elsewhere on the frontal wall; most frequently, a single avicularium is

developed medially, proximal to spiramen. Proximally directed vicarious

avicularia present in a single linear series along the branch edge.

Etymology

Confusaneus (L.)—mixed, referring to characters shared with other

species (below).

Remarks

The proximally directed frontal avicularium, which develops in later

ontogenetic stages, is also seen in A. abdita (p. 97). However, that species may

be distinguished from A. confusanea by the shape of the primary orifice (Fig.

23D), and by the broad, flat, plate-like branches of the colony.

Measurements (means of 25 values) in mm

Lz Iz

0,48 O27

Adeonella conspicua sp. nov.

Fig. 22A—D

Material

Holotype: SAM-A26440, station SM 179, 33°30,3’S 27°22,1'E, 80 m.

Description

Colony erect, branching, rigid. Zooids oval to hexagonal, or irregular,

separated by distinct grooves. Primary orifice almost semicircular, with a short,

wide and shallow sinus. Peristome thickened, nodular, but relatively depressed,

secondary orifice semicircular. Frontal wall convex, nodular, with small but

distinct pores; spiramen circular, situated close to base of peristome. Sinus of

primary orifice visible through it in newly developed zooids. Avicularia

typically single, rarely paired, situated lateral to spiramen; rostrum triangular,

relatively short, at an acute angle to frontal plane, directed distomedially or,

rarely, proximally. In zooids towards the basal region of the colony, avicularia

may be developed elsewhere on the frontal wall. Marginal vicarious avicularia

very characteristic, rostrum hooked and projecting from the edge of the

branch; interspersed with smaller types, or bearing them.

Etymology

Conspicuus (L.)—manifest, referring to the characters of the species.

Remarks

A single juvenile colony was found, 9 mm high and bearing three short

lobes with a maximum width of 2 mm. Despite this shortage of material, the

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 22. A-D. Adeonella conspicua sp. nov. A. Young zooids at a branch bifurcation, note prominent

hooked avicularian rostra. B. Older zooids with nodular frontal walls. C. The branch edge, with vicarious

avicularia. D. Outline diagram of primary orifice. E-H. Adeonella distincta sp. nov. E. Four typical

zooids. F. The branch edge, with vicarious avicularia. G. Old zooids from the colony base. H. Outline

diagram of primary orifice. I-K. Adeonella infirmata sp. nov. I. Young zooids from close to the colony

margin, note kenozooids on right. J. Old zooids with thickened frontal walls. K. Outline diagram of

primary orifice.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 95

specimen seems sufficiently distinctive to warrant the introduction of a new

specific name.

Measurements (means of 25 values) in mm

Lz Iz

0,47 ORs

Adeonella distincta sp. nov.

Fig. 22E—H

Material

Holotype: SAM—A26441, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Description

Colony erect, branching, rigid, attached by encrusting base; branches

bilaminar, slender, up to 35 mm long in present material, with a maximum

width of 2 mm. All branches in the same plane. Zooids elongate, rectangular,

separated by deep grooves, becoming irregular in outline in later ontogeny.

Primary orifice broadly drop-shaped: anter semicircular, poster forming a broad

and symmetrical U-shape. Peristome developing an orbicular secondary orifice.

Avicularium single or, rarely, paired, situated lateral to spiramen and directed

distally; rostrum acute triangular, not quite reaching the proximal border of the

secondary orifice. Frontal wall convex, finely granular, with numerous small

pores; spiramen more or less centrally placed, midway along the length of the

zooid, at first longitudinally oval, becoming quite circular as frontal calcification

thickens. Vicarious avicularia developed along branch edges, interspersed with

small kenozooids, which may or may not bear small adventitious avicularia.

Etymology

Distinctus (L.)—different.

Remarks

The orientation of the frontal avicularium may vary within the colony, and

in some zooids the rostrum may extend medially between the spiramen and the

secondary orifice. In the oldest parts of the colony avicularia may be developed

proximal to the spiramen, but there are rarely more than three per zooid. This

species is most similar to A. cracens Hayward & Cook (1979) but is distinguished

by the smaller size of its zooids and by the shape of the primary orifice, which in

A. distincta is sinuate but in A. cracens forms a simple transverse ellipse.

Fragments of three live colonies of this species were collected.

Measurements (means of 25 values) in mm

Lz Iz

0,59 0,30

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Adeonella infirmata sp. nov.

Fig. 22I-K

Material

Holotype: SAM—A26442, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Description

Colony erect, branching, rigid, attached by an encrusting base; branches

bilaminar, up to 45 mm high in present material, with a maximum width of

5mm. Zooids elongate, hexagonal or rectangular, rounded distally and often

tapered proximally; smallest zooids in middle region of branch. Primary orifice

bell-shaped, proximal border shallowly convex below indistinct condyles.

Peristome with orbicular secondary orifice. Avicularium single or paired,

arising laterally at a level midway between the spiramen and the secondary

orifice; rostrum acute triangular, short, directed distally or distomedially,

typically extending to proximolateral corners of secondary orifice. Frontal wall

convex, finely granular, evenly perforated by numerous small pores; spiramen

situated close to proximal border of secondary orifice, large, transversely oval,

arched, the proximal border of the primary orifice visible in all but the most

heavily calcified zooids. Small adventitious avicularia may develop elsewhere

on the frontal wall in the later stages of ontogeny, with variable orientation.

Vicarious avicularia present in single linear series along branch edge, the rostra

of which frequently project noticeably from the branch edge. These may

alternate with small kenozooids.

Etymology

Infirmis (L.)—weak, referring to the rather delicate form of the colony.

Remarks

Zooid rows frequently terminate at the edges of the branch in small

kenozooids which often bear adventitious avicularia. The longitudinal branch

keel typical of some of the species described here (p. 99) is only poorly

developed in A. infirmata; it seems to be formed from thickened frontal

calcification rather than from inflated avicularian cystids, although small

adventitious avicularia are more frequent in the middle regions of the branch.

Fragments of five living colonies were collected.

Measurements (means of 25 values) in mm

ez Iz

0,71 0,30

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 97

Adeonella abdita sp. nov.

Fig. 23A-E

Material

Holotype: SAM—A26443, station SM 239, 32°14,8’S 29°00,8’E, 90 m.

Other material: stations SM 163, SM 163/164, SM 164, SM 1835.

Description

Colony erect, branching, rigid, attached by an encrusting base; branches

bilaminar, up to 30 mm long in present material, with a maximum width of

5mm. Zooids elongate, rectangular or oval, frequently tapered proximally;

separated by distinct grooves, becoming obscured in older parts of colony.

Primary orifice as wide as long, with inconspicuous, blunt lateral condyles

demarcating a broad, shallow proximal sinus. Peristome with a transversely

oval secondary orifice, the inner proximal edge developing a blunt denticle.

Frontal wall convex, granular, closely punctured by numerous small pores.

Spiramen medially situated in the distal third of the zooid, orbicular, oval or

elongate, overarched from the left or right by a distinct hood, so that its

Opening appears to be perpendicular to the frontal plane of the zooid. A single

avicularium adjacent to spiramen, rostrum acute triangular, directed distally;

becoming quite immersed, below the level of the spiramen hood. A second

avicularium may be developed on the other side of the spiramen, preventing

complete development of its hood; occurs rarely in some colonies, more

frequently in others. Additional avicularia may develop laterally, between the

Sspiramen and the secondary orifice, smaller than the frontal type, single or

paired, with rostrum directed towards the peristome. In the oldest parts of the

colony these may be very frequent, with four or five developed around the

distal end of the zooid, all directed towards the peristome. Frontal avicularia

are progressively immersed and eventually completely obscured; peristomial

avicularia then proliferate, but there is no regeneration of regularly cule rueites

frontal avicularia (cf. A. gibba, p. 99).

In well-grown colonies each branch has a median longitudinal keel,

formed by the development of numerous adventitious avicularia, each with a

particularly voluminous cystid. Large vicarious avicularia present in single

linear series along branch edges.

Etymology

Abditus (L.)—hidden, referring to the spiramen.

Remarks

Adeonella expansa O’Donoghue shows a similar development of the

spiramen to that of A. abdita, but differs in that the adjacent avicularium is

very small, being scarcely longer than the spiramen itself. Further, unlike the

present species, the peristomial avicularia are almost medially situated, with

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. A-E. Adeonella abdita sp. nov. A. Young zooids showing characteristic form of spiramen and

adjacent avicularium. B. Older zooids, with numerous additional avicularia. C. Zooids from the branch

keel, showing immersed spiramina and prominent avicularian cystids. D. Outline diagram of primary orifice.

E. View of the branch edge, showing vicarious avicularia. F-J. Adeonella gibba sp. nov. F. Young zooids

showing characteristic form of spiramen and adjacent avicularium. G. Later zooids with nodular frontal

walls. H. The branch edge showing vicarious avicularia. I. Old zooids from the colony base, with

proximally directed frontal avicularia. J. Outline diagram of primary orifice.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 99

transversely orientated mandibles. The colony of A. expansa develops broad

plate-like branches.

Measurements (means of 25 values) in mm

Lz Iz

0,76 0,34

Adeonella gibba sp. nov.

Fig. 23F—J

Adeonella pectinata: O’Donoghue, 1924: 51.

Material

Holotype: SAM—A26444, station SM 179, 33°30,3’S 27°22,1’E, 80 m.

Other material: stations SM 163, SM 163/164, SM 179, SM 180, SM 185.

Description

Colony erect, branching, rigid, rising from an encrusting base; branches

bilaminar, up to 4 mm wide. Zooids oval to rectangular, convex; separated by

deep grooves, less apparent in oldest parts of colony. Primary orifice as wide as

long, proximal half constituting a broad shallow sinus. Peristome with an

orbicular secondary orifice, inner proximal border with a broad shelf-like

denticle. Frontal wall granular, closely perforated by numerous round pores

which become accentuated as calcification continues; marginal series particu-

larly prominent in later ontogenetic stages. A pronounced umbo frequently

present on proximal frontal wall. Spiramen medially situated in distal half of

zooid; relatively large and rather elongate, partly hidden by a lateral hood, the

aperture thus appearing perpendicular to frontal plane of zooid. A single lateral

avicularium developed, sporadically, just proximal to aperture, rostrum

extending obliquely distally on to the peristome; frequently absent. Larger,

vicarious, avicularia present in single linear series along branch edges, and

often intercalated in zooid rows at the margins of the branch; mandible acute

triangular, distally directed.

As secondary calcification proceeds the spiramen becomes deeply

immersed, and the frontal wall quite concave; a second frontal avicularium is

then budded, obscuring the first and partly filling the concavity. This new

avicularium is orientated proximally in all zooids; the extended opening of the

Spiramen is just visible at its distal end. A median longitudinal ridge may be

apparent along the branches of the oldest part of the colony, formed by the

development of extra adventitious avicularia; elsewhere, however, the avicula-

ria do not proliferate as they do in A. abdita (above).

Etymology

Gibbus (L.)—protuberant, referring to the frontal umbones of later

zooids.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The secondary frontal avicularium, with its constant orientation, is a

characteristic feature of this species and serves to distinguish it further from

Adeonella abdita.

Measurements (means of 25 values) in mm

IL Iz

(SY 0,29

Adeonella alia sp. nov.

Fig. 24A—D

Material

Holotype: SAM-—A26445, station SM 179, 33°30,3’S 27°22,1’E, 80 m.

Description

Colony erect, branching, rigid, attached by an encrusting base; branches

bilaminar, broad and flat with lobed edges, up to 20 mm wide. Zooids rounded

distally, tapered proximally. Primary orifice with semicircular anter and slender

V-shaped poster, condyles strongly marked. Peristome with a semicircular or

semi-elliptical secondary orifice, the inner proximal border thickened and

raised medially to form a broad denticle with a serrated margin. Spiramen

situated close to base of peristome, its orifice vertical to frontal plane of zooid;

proximal edge incomplete in younger zooids, later quite rounded. Frontal wall

finely granular, with large, deeply sunk pores giving it a rugose appearance.

Avicularium single or paired, arising adjacent to spiramen, elongate rostrum

directed distomedially towards the proximal border of the secondary orifice.

When only a single avicularium is present a short conical umbo may be

developed on the opposite side of the spiramen. In early ontogeny the middle

area of the frontal wall, proximal to the spiramen, is deeply concave;

subsequently an extra adventitious avicularium develops here, with a prominent

tumid umbo and an elongate, proximally directed rostrum. Other, smaller,

avicularia may develop elsewhere on the frontal surface, and along the branch

edges, in the oldest parts of the colony.

Etymology

Alius (L.)—another.

Remarks

The single colony found was 40 mm high, with broad lobed branches;

either flat, or concave, or distinctly twisted about the longitudinal axis. The

concave frontal wall, and the subsequent development of a proximally directed

avicularium, recall the morphology of A. gibba (above). However, A. alia is

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 101

Fig. 24. A-D. Adeonella alia sp. nov. A. Young zooids, with typical concave frontal walls. B. Later

zooids, with characteristic development of frontal avicularia. C. Two zooids in oblique view to show

peristomial denticle. D. Outline diagram of primary orifice. E. Reteporella dinotorhynchus Hayward

& Cook. F-G. Sertella lata (Busk). F. Portion of a colony showing ovicelled zooids. G. Outline

diagram of primary orifice. H. Schizoretepora tessellata (Hincks).

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

distinguished from that species by the shape of the primary orifice, the form of

the spiramen, and by the characteristic form of its colony.

Measurements (means of 20 values) in mm

EZ Iz

OFS2 0,29

Family Tessaradomidae Jullien, 1903

Tessaradomidae Jullien in Jullien & Calvet, 1903: pl. 14. Hayward & Ryland, 1979: 242.

Tessaradoma Norman, 1869

Tessaradoma Norman, 1869: 309. Lagaaij & Cook, 1973: 494. Hayward & Ryland, 1979: 242.

Tessaradoma bispiramina Hayward & Cook, 1979

Tessaradoma bispiramina Hayward & Cook, 1979: 90, fig. 13A—D.

Material

Stations SM 103, SM 151, SM 233, SM 234.

Remarks

Living colonies were collected from stations SM 233 and SM 234, within

the bathymetric range observed by Hayward & Cook (1979).

Tessaradoma circella Hayward & Cook, 1979

Tessaradoma circella Hayward & Cook, 1979: 91, fig. 13E—H. Hayward 1981: 48, fig. 25.

Material

Stations SM 103, SM 233.

Remarks

The astogeny of the basal attachment ring of Tessaradoma circella and the

morphology of the ancestrula have recently been described by Hayward (1981).

Family Sertellidae Jullien, 1903

Sertellidae Jullien in Jullien & Calvet, 1903: 57. Hayward & Ryland, 1979: 260.

The genera and species here assigned to the Sertellidae do not, perhaps,

constitute a natural assemblage, yet display a confusing similarity in many of

their morphological features. Lagaaij (1952: 109-110) followed accepted taxo-

nomic practice in proposing the use of Sertellidae Jullien (1903) for the genera

then grouped in the old family Reteporidae Smitt (1868), the type genus of

which (Retepora Lamarck, 1801) appears to have no recognizable or acceptable

type species. Levinsen (1902, 1909) had expanded the Reteporidae to include

encrusting, non-fenestrate genera such as Rhynchozoon and Schizotheca.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 103

Despite the objection of Harmer (1933), this arrangement is now generally

accepted (e.g. Osburn 1952; Powell 1967).

The substantial number of genera of both erect and encrusting forms

presently included in the Sertellidae appears to have several important

morphological characteristics in common. The primary orifice, though widely

variable in shape, typically has a denticulate or beaded distal rim (‘vestibular

arch’), and distinct condyles; the ovicell is imperforate, often prominent, and in

most species is provided with a conspicuous frontal lip or labellum. In several

genera (Sertella, Schizotheca, Triphyllozoon, for example) the ovicell is further

characterized by a frontal fissure of variable size and extent. The peristome is

often well developed, often with a notch, fissure or ‘pseudospiramen’

proximally, and frequently incorporating a suboral avicularium. Adventitious

and vicarious avicularia occur in most genera.

The genera of erect species seem to constitute a fairly homogenous group,

although some species diverge from the common pattern. Schizoretepora

tessellata (Hincks), for example, has a smooth orifice rim and a minimal

development of the peristome. The growth form of this species (see p. 108),

and of the bilaminate species of Reteporella described by Hayward & Cook

(1979), perhaps suggest a link with the encrusting genera Schizotheca and

Rhynchozoon. The ovicell of the former, moreover, is very similar to those of

S. tessellata and Reteporella. Rhynchozoon is characterized principally by its

distinctly beaded orifice rim, by the ovicell, which has a frontal ‘area’ of

uncalcified ectooecium and often possesses a short labellum, and by the

peristomial complex, which includes a hammer- or anvil-shaped uncinate

process projecting distally over the orifice. The uncinate process is variably

developed; in the new species here assigned to Rhynchozoon it varies from an

inconspicuous structure in R. incallidum to the massive development seen in R.

oscitans and R. stomachosum. In R. ptarmicum there is no uncinate process;

this fact caused some difficulty in deciding to which genus the species should be

assigned, but in comparison with certain other species in the present collections

prompted consideration of the systematic relationships of the Sertellidae with

other family groups.

The genus Brodiella was introduced by Uttley & Bullivant (1972) for

Schizoporella longispinata Busk. This well-defined species was first described

(Busk 1884) from the Straits of Magellan, was reported from two localities in

the Chatham Islands, New Zealand (Uttley & Bullivant 1972), and was found

in the present collections. The affinities of Brodiella with Rhynchozoon seem

clear; in both genera the zooid has a conspicuously beaded orifice and the

Ovicell typically develops a distinct labellum. On this basis Gautier (1962)

referred the warm-temperate North Atlantic species, Lepralia armata Hincks,

to Rhynchozoon and it is now clear (D. P. Gordon 1978, in litt.) that the

species is correctly placed in Brodiella. Curiously, Uttley & Bullivant (1972)

remarked upon the similarity of the two genera, yet assigned Brodiella to the

Schizoporellidae. Although the Meiring Naude specimens of B. ignota sp. nov.

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

did not include ovicelled zooids, its morphology seems quite consistent with the

generic diagnosis of Brodiella. The genus is here placed within the Sertellidae,

emphasizing its relationship to Rhynchozoon. However, it must then be

recognized that there is a need to consider the systematic affinities of two large

and heterogenous genera, Cleidochasma Harmer and Hippoporella Canu, both

currently assigned to the family Cleidochasmatidae Cheetham & Sandberg

(1964). Species of both genera were present in the Meiring Naude collections.

Cleidochasma perspicua sp. nov. (p. 76) conforms most nearly to the

diagnosis of Cleidochasma, although its ovicell, which has a well-marked

frontal labellum, is clearly more similar to those of Rhynchozoon and

Brodiella than the spherical, tuberculate ovicells of most species of Cleido-

chasma. A second species, C. contractum (Waters), figured by Cook

(1964b: 15, fig. SA), also has the same, typically Sertellid, ovicell morphology

and, moreover, a finely denticulate orifice rim. Hippoporella labiata sp. nov.

(p. 80) constitutes a more severe test of current systematics, and its inclusion

in Hippoporella can only be regarded as a doubtful compromise. The

morphology of the ovicell and the denticulate orifice rim again suggest an

affinity with the encrusting genera placed among the Sertellidae, but the

species does not conform strictly with the taxonomic diagnosis of any of them.

The lepralioid orifice and prominent suboral umbo suggest that it might be

temporarily accommodated in Hippoporella, particularly as this genus includes

many tropical species (e.g. Harmer 1957: 1096. Cook 19646: 8) that seem to

have little in common with the boreal-arctic type species, H. hippopus

(Smitt). The Australian species Schizoporella pulchra MacGillivray (1891: 81,

pl. 9 (fig. 7)) illustrates the apparent convergence of characters in tropical,

particularly Indo-West-Pacific, species of these different genera. S. pulchra

forms encrusting, multilaminar colonies (e.g. BMNH 1897.5.1.758); the zooid

has a broadly cleithridiate orifice with a finely denticulate distal rim. There

are single or paired lateral oral avicularia, with short, distally directed,

semicircular mandibles; one of these is frequently supplanted by an enlarged

avicularium with a slender spatulate mandible. These avicularia seem to be

interzooidal in origin and analogous to those seen in Brodiella, yet the tall

basally jointed spines and proximal peristome rim characteristic of that genus

are lacking in S. pulchra. The ovicell is prominent, frontally flattened, with a

large area of uncalcified ectooecium and in the older parts of the colony the

frontal walls of many zooids are obscured by massive, frontally budded,

vicarious avicularia with broad scaphoid mandibles. These features are most

usually associated with Rhynchozoon.

It seems clear, in conclusion, that any further investigation into systematic

relationships within the Sertellidae must be combined with a thorough review

of the morphology and systematics of many tropical species at present placed

among the Cleidochasmatidae. The need for such a study is highlighted by the

anomalous positions of several species reported upon here, but is beyond the

scope of the present work.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 105

Sertella Jullien, 1903

Sertella Jullien in Jullien & Calvet, 1903: 57. Hayward & Ryland, 1979: 260.

Sertella lata (Busk, 1884)

Fig. 24F—G

Retepora lata Busk, 1884: 115, pl. 27 (fig. 1).

Material 3

Stations SM 163, SM 179, SM 185.

Description

Colony generally thick and robust; trabeculae composed of three to seven

longitudinal series of zooids, fenestrulae oval, small, 0,5-1,0 mm long. Primary

orifice of zooid about twice as broad as long, lacking the usual denticulate rim;

condyles basally deflected and not visible in frontal view. Peristome thin, with a

central fissure and a small proximal pseudosinus, a single short spine present on

each edge of peristome. Adventitious avicularia numerous, distributed over the

entire frontal surface of the colony, mandible either semi-elliptical or acute

triangular, mostly less than 0,1 mm long; rarely, an enlarged avicularium

occurs, with a triangular mandible up to 0,15 mm long. Ovicell prominent,

pear-shaped, convex, with a longitudinal frontal fissure and a very long

labellum extending deep into the peristome. Basal surface of colony densely

papillate, with a few scattered avicularia; frontal calcification becoming

similarly papillate in later ontogenetic stages.

Remarks

The most characteristic feature of Sertella lata is the pear-shaped ovicell

with its very long labellum. This feature was not illustrated by Busk (1884,

pl. 27 (fig. 1)) who shows the ovicell orifice opening just above the level of the

peristome. However, examination of the type specimen shows this detail to be

incorrect, and in all respects it is in close correspondence with the Meiring

Naude material. This species is known only from South Africa.

Sertella verecunda sp. nov.

Figs 25, 26A

Material

Holotype: SAM—A26446, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Description

Colony robust, holotype 6 mm high with a spread of 15 mm. Trabeculae

stout, comprising five to eight longitudinal series of zooids, fenestrulae

irregularly oval, up to 2 mm long. Zooids hexagonal or irregular, small, about

0,4 mm long by 0,2 mm broad; frontal calcification tessellated, with a few large

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

distinct marginal pores. Primary orifice transversely oval, deeply immersed and

seen only at broken edges. Peristome tubular and deep, secondary orifice

scarcely raised above frontal surface of zooid, a single pair of lateral oral spines

present in early ontogeny; proximal border with a rounded notch, later closed

forming a circular pseudosinus, persisting through later ontogenetic stages.

Avicularia sporadically developed on frontal wall, one, two or more per zooid,

either short, with a rounded rostrum bearing a semicircular mandible, a thick

cross-bar and stout columella, or more elongate, almost bispatulate, with a

semi-elliptical mandible, a centrally placed cross-bar and a delicate columella.

The short type has a denticulate distal margin to the rostrum. Large vicarious

avicularia distributed around edges of fenestrulae; rostrum quadrate, parallel-

sided, palatal foramen triangular, cross-bar slender with a delicate columella.

Ovicell pyriform, smooth surfaced, with an elongate central fissure and a short

labellum; obscured by a thickened ooecial cover early in ontogeny. Basal

surface of colony with both types of adventitious avicularia, often numerous.

Etymology

Verecundus (L.)—bashful, an allusion to the hidden primary orifice.

Remarks

S. verecunda may be distinguished from other species of Sertella by its

pronounced pseudosinus, and the absence of peristomial avicularia, by the two

types of adventitious avicularia, and by the characteristic quadrate fenestral

avicularia.

Schizoretepora Gregory, 1893

Schizoretepora Gregory, 1893: 224. Harmer, 1933: 619.

Schizoretepora tessellata (Hincks, 1878)

Fig. 24H

Retepora tessellata Hincks, 1878: 358, pl. 19 (figs 9-12). Busk, 1884: 112, pl. 27 (fig. 8).

O’Donoghue & De Watteville, 1935: 210; 1937: 15. O’Donoghue, 1957: 91.

Material

Stations SM 163, SM 163/164, SM 180, SM 1835.

Description

Colony rather delicate; trabeculae composed of four to six longitudinal

series of zooids, or up to eight at points of trabecular fusion, fenestrulae

elongate-oval, up to 1,6 mm long. Zooids hexagonal or irregular, rather flat,

separated by distinct sutures at first, boundaries later indistinct. Primary orifice

semi-elliptical, longer than wide; distal border smooth, proximal border gently

concave, with a short notch-like sinus. Peristome developing initially as a thin

erect flange on each side of the orifice, not developed proximally; later

107

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

ia and

with

lar

ing avicu

showi

ip of a branch,

f a colony,

ing

x 100.

ion o

icells.

Port

ids from the grow

Ils

ing OV

nov

Zoo

develop

x 100

ina

Fig. 25. Sertella verecunda sp

pseudospiram

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

immersed and somewhat unclear, but proximal side of orifice still distinct. Up

to six distal and lateral oral spines present, only the proximal pair of which

persist as secondary calcification proceeds. Frontal calcification finely granular

with a small number of marginal pores. Avicularia variably developed;

frequently proximolateral to orifice, small, with a semi-elliptical mandible; also

elsewhere on frontal wall, small, with more elongate, semi-elliptical or acute

triangular mandibles. Gigantic avicularia typically frequent; cystid obscuring

most of the zooid bearing it, rostrum elongate, acute triangular, up to 0,3 mm

long, perpendicular to frontal plane of zooid. Ovicell prominent, spherical,

with a conspicuous oval orifice situated well above peristome rim. Basal surface

sutured, with numerous small avicularia and typically a gigantic avicularium at

the proximal end of each fenestrula.

Remarks

Schizoretepora tessellata is unusual among the Sertellidae in possessing the

facility to grow as a typical fenestrate colony, or as a folded bilaminar sheet.

This facility may be expressed in a single colony, with substantial bilaminate

colonies producing peripheral fenestrulae (e.g. BMNH 1962.6.4.17pt., Millers

Point, Cape Town). Described originally from South Australia, it was reported

from Simon’s Bay (Cape of Good Hope) by Busk (1884). Comparison of

Australian and South African specimens with the Meiring Naude samples shows

some variation in the width of the primary orifice both between and within each

of the three series of specimens. Additionally, some of the Australian

specimens develop long (0,8 mm) antenniform spines. However, the avicularia

and ovicells are closely comparable in all instances.

Reteporella Busk, 1884

Reteporella Busk, 1884: 126. Harmer, 1934: 572.

Reteporella dinotorhynchus Hayward & Cook, 1979

Fig. 24E

Reteporella dinotorhynchus Hayward & Cook, 1979: 95, fig. 14A—D.

Material

Stations SM 163, SM 163/164, SM 184, SM 1835.

Remarks

The material from station SM 185 comprised a more complete ontogenetic

sequence than was provided by the first Meiring Naude samples, and it is clear

that the original description of this species must be amplified. In particular, it is

evident that a small adventitious avicularium develops on the peristome

adjacent to the lateral notch. This is missing in even the least worn fragments,

and is obscured by the peristome in the oldest parts of the colony. Developing

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 109

ovicells were found in the present specimens, but no complete examples

remained; the ovicell seems to have a substantial frontal fissure.

Iodictyum Harmer, 1933

Todictyum Harmer, 1933: 624; 1934: 537.

Iodictyum flosculum sp. nov.

Fig. 27A—C

Material

Holotype: SAM-—A26447, station SM 164, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163, SM 185.

Description

Colony white, forming a delicate cup shape, up to 8mm high with a

spread of 6 mm in present material; trabeculae composed of three alternating

series of zooids, increasing to four or five at trabecular fusion, fenestrulae

elongate-oval, up to 0,8 mm long. Zooids quadrate, rather flat, separated by

distinct raised sutures; calcification smooth, imperforate except for a few rather

large round marginal pores, usually situated at the proximal end of the zooid.

Primary orifice semicircular, distal edge not denticulate, proximal edge slightly

concave, with large blunt lateral condyles. Peristome characteristic of genus:

developed early in ontogeny and completely hiding orifice; erect, broadly

flared, the free edge drawn into a number of delicate spikes, aligned with a

second series of smaller spikes around the inner rim of the peristome. The

inner denticulations of the peristome are continuous with delicate ridges which

extend down its interior surfaces; proximally, two of the ridges delimit a

U-shaped channel which appears as a distinct pseudosinus in damaged zooids.

With increasing calcification the peristome is immersed, the outer spikes are

lost and the aperture appears as a simple denticulate opening. Avicularia

infrequent, orientated oblique to proximal edge of peristome, rostrum

elongate, subtriangular, hooked at tip. Ovicell not found. Basal surface of

colony faintly papillate, crossed by conspicuous sutures, lacking avicularia.

Etymology

Flosculus (L.)—a little flower.

Remarks

This delicate species represents the first occurrence of the genus Jodictyum

remote from the western Pacific.

Measurements (means of 20 values) in mm

ILyZ Iz

0,42 0,19

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Rhynchozoon Hincks, 1895

Rhynchozoon Hincks, 1895: V. Hayward & Ryland, 1979: 271.

This large and difficult genus is widely distributed in both temperate and

tropical waters. Most bryozoan faunas of the continental shelf seas include one

or more species of Rhynchozoon, yet few of these are adequately characterized

and taxonomic confusion obscures the identity of most of them. In almost all

species of Rhynchozoon the primary orifice of the zooid becomes immersed in a

secondarily developed peristomial complex which is frequently ornamented

with a variety of knobs and processes, and there is often a proliferation of small

adventitious avicularia. These later stages of ontogeny are often subject to

great variation within a single colony and cannot be used as reliable specific

characters. The shape of the primary orifice and the morphology of the suboral

avicularium, together with its uncinate process, should be used as the most

important features for discriminating between species and it is clear that when

this is done a greater diversity of species is revealed than is apparent from the

literature (e.g. Hayward 1974). These features, together with selected second-

ary characters, show that the South African fauna includes a rich variety of

species, none of which from among those represented here may be identified

with either of the two species described by O'Donoghue & De Watteville

(1935), or with any of the Indo-West-Pacific species reported by Harmer

(1957).

Rhynchozoon documentum sp. nov.

Fig. 26B-C

Material

Holotype: SAM—-A26448, station SM 163/164, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163, SM 164.

Description

Colony encrusting, multilaminar, forming small nodular sheets. Zooids

distinct only at growing edge, oval and convex; frontal wall smooth, with large

marginal pores. Primary orifice broader than long; anter transversely oval, with

finely denticulate rim, poster forming a short, U-shaped sinus, oral spines

lacking. Suboral avicularium large, distinct in early ontogenetic stages,

mandible elongate elliptical, uncinate process well developed and conspicuous,

delimiting laterally a closed pseudosinus. Peristome developing a thickened rim

obscuring primary orifice and suboral avicularium; proximally a median fissure

is flanked by short conical processes, in later ontogeny further short columnar

processes are developed around the whole of the secondary orifice. Frontal

avicularia often numerous, small, with semicircular or semi-elliptical mandibles.

Etymology

Documentum (L.)—an example.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

~ 03

s @r r @

Fig. 26. A. Sertella verecunda sp. nov., the edge of a fenestrula (left) with a vicarious avicularium.

x 96. B—C. Rhynchozoon documentum sp. nov. B. Zooids at a growing edge, note primary orifice

C. Later zooids, each with one frontal avicularium and

and conspicuous areolae. X 76.

conspicuous areolae. x 64,3.

nD ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 27. A-C. Jodictyum flosculum sp. nov. A. Young zooids showing typical flared spinose peristomes.

B. Old zooids with worn peristomes; a single suboral avicularium. C. a complete juvenile colony, possibly

including the ancestrula. D-F. Rhynchozoon incallidum sp. nov. D. Zooids from the growing edge.

E. Later zooids, with ovicells and developed peristomes. F. Outline diagram of primary orifice.

G-H. Rhynchozoon oscitans sp. nov. G. Four young zooids; note conspicuous uncinate process. H. Outline

diagram of primary orifice. I-K. Rhynchozoon stomachosum sp. nov. I. Two young zooids. J. Later

zooids, with typical development of the umbo, and two vicarious avicularia. K. Outline diagram of the

primary orifice.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 113

Remarks

In later ontogenetic stages the marginal pores of the zooids are

particularly distinct, appearing as elongate tubular structures within the rather

vitreous calcification, and appear to be associated with the production of

numerous small adventitious avicularia. This particularly conspicuous morpho-

logical feature seems to be a fairly constant characteristic of this species.

Measurements (means of 20 values) in mm

GZ Iz

0,39 0,32

Rhynchozoon beatulum sp. nov.

Fig. 28

Material

Holotype: SAM—A26449, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: stations SM 163/164, SM 184, SM 185.

Description

Colony encrusting, forming small irregular sheets, apparently unilaminar.

Zooids oval to hexagonal, strongly convex; frontal wall finely granular,

becoming increasingly vitreous in later ontogenetic stages, marginal pores

conspicuous. Primary orifice wider than long; anter orbicular, with finely

denticulate rim, poster broad, shallowly concave, two short, widely-spaced,

distal oral spines present in newly budded zooids. Suboral avicularium large

and distinct, rostrum hooked, with elongate triangular mandible; uncinate

process bluntly triangular. Peristomial thickening moderate: a short, blunt

umbo develops on the cystid of the avicularium, and a second umbo on the

opposite lateral edge of the secondary aperture, with a small asymmetrically

developed notch between; in later ontogenetic stages, one or two short knobs

may be developed on the distal rim of the peristome. Frontal avicularia

typically limited to one on each zooid, on the proximal! half of the frontal wall;

rostrum triangular, directed proximally. Ovicell rather elongate, with an oval

area of uncovered entooecium frontally, and a large frontal labellum; obscured

by a thickened ooecial cover.

Etymology

Beatus (L.)—happy, an allusion to the broad sinus.

Remarks

Despite the progressive thickening of the frontal calcification in later

ontogeny, the marginal pores of R. beatulum remain distinct. Further, the

suboral avicularium is rarely completely hidden, and the peristomial rim shows

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

only a limited development of umbones. The frontal avicularium is remarkably

constant in position, shape and orientation in all the present specimens, and

assists in distinguishing this species from others in the South African fauna.

Measurements (means of 20 values) in mm

Lz Iz

0,45 0,35

Rhynchozoon incallidum sp. nov.

Figs 27D-F, 29A-B

Material

Holotype: SAM—A26450, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Other material: stations SM 163, SM 163/164, SM 180, SM 184.

Description

Colony encrusting, forming small unilaminar patches. Zooids oval to

hexagonal, frontal wall smooth and vitreous with few indistinct marginal pores.

Primary orifice as wide as long: anter semicircular, with finely denticulate rim,

poster forming a short quadrate sinus; up to four slender distal oral spines

present. Suboral avicularium on a voluminous cystid, developed apically as a

blunt umbo; rostrum triangular, perpendicular to frontal plane of zooid and

facing laterally, uncinate process poorly developed, not prominent. A second

umbo typically developed on opposite lateral border of orifice, a third may be

present distal to the suboral avicularium; these develop independently of the

spine bases. Frontal avicularia not found. Ovicell spherical, flattened frontally,

with an irregular area of uncovered entooecium and a short, broad labellum.

Ooecial cover developing early in ontogeny, frequently with one or more low

umbones, but ovicell remaining conspicuous. Zooid boundaries become

indistinct in later ontogenetic stages, additional umbones may be developed on

the frontal wall, but the primary orifice is not deeply immersed and generally

remains visible in frontal view.

Etymology

Incallidus (L.)—simple, referring to the relative simplicity of the zooidal

morphology.

Remarks

The shape of the primary orifice, the poor development of the uncinate

process and the relatively simple zooid morphology serve to distinguish R.

incallidum from other species of Rhynchozoon.

Measurements (means of 20 values) in mm

IL Iz

0,45 OZ

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 115

Ow?

SENS

SAS

Sos

Fig. 28. Rhynchozoon beatulum sp. nov. A. Zooids at a growing edge.

x 46,5. B. Details of a primary orifice. x 240. C. Later zooids, showing

the characteristic frontal avicularium. x 61.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

RhynchoZoon oscitans sp. nov.

Fig. 27G—H

Material

Holotype: SAM—A26451, station SM 185, 33°39,3’S 27°11,6’E, 90 m.

Other material: station SM 180.

Description

Colony encrusting, unilaminar. Zooids oval to hexagonal, convex,

distinct, separated by well-marked sutures. Frontal calcification thick and

smooth, with a few inconspicuous marginal pores. Primary orifice longer than

wide, lepralioid; proximal edge straight, distolateral rim denticulate above

massive rectangular, basally deflected condyles. Four or five oral spines

present, the distalmost pair persisting as short stumps in later ontogeny.

Peristome low; suboral avicularium situated medioproximally on outer peri-

stome rim, at oblique angle to frontal plane, mandible small, semi-elliptical.

Uncinate process large, quadrate, very conspicuous. Frontal avicularia and

ovicells not found.

Etymology

Oscitans (L.)—yawning, referring to the conspicuous primary orifice.

Remarks

Only two small specimens of this species were found, and neither

represents a complete astogenetic or ontogenetic sequence. However, the

shape of the orifice, and the massive uncinate process are very distinctive and

sufficient to distinguish R. oscitans from all known species of Rhynchozoon.

Measurements (means of 20 values) in mm

Lz Iz

0,53 0,37

Rhynchozoon stomachosum sp. nov.

Fig. 27I-K, 29C

Material

Holotype: SAM—A26452, station SM 234, 32°15’S 29°09,1’E, 500-520 m.

Description

Colony encrusting, multilaminar. Zooids at growing edge large, oval and

convex; frontal calcification granular, becoming rather nodular in later

ontogenetic stages, marginal pores few and small but distinct. Primary orifice |

D-shaped, wider than long; proximal border almost straight, distolateral rim

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES iLY/

Fig. 29. A-B. Rhynchozoon incallidum sp. nov. A. A group of zooids from a dead colony.

x 100. B. Detail to show primary orifice and spine bases. x 300. C. Rhynchozoon

stomachosum sp. nov. X 41.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

finely denticulate, apparently without condyles. No oral spines. Suboral

avicularium obliquely transverse to orifice, at a slight angle to frontal plane;

rostrum triangular, with hooked tip. Uncinate process large and conspicuous.

Peristome morphology variable; typically developing a columnar, clavate umbo

proximomedially, with a knobbed granular surface, adjacent to a shallow

notch; two, rarely more, short processes may also be present on the lateral

borders of the peristome. Alternatively, two shorter, more slender, equisized

umbones may develop, flanking a more prominent peristomial notch. Frontal

avicularia of two sizes: on peripheral zooids situated laterally, close to suture,

and directed proximolaterally, the plane of the palate typically at a right angle

to the frontal plane; in later ontogeny a larger avicularium may develop on the

frontal wall along the midline of the zooid, directed proximally with the plane

of the palate normal to the frontal plane. In both types the rostrum is elongate,

slightly expanded distally to give a slender spatulate shape, straight or gently

curved laterally; cross-bar entire, palate with a narrow, elongate central fissure.

Ovicell longer than wide, flattened frontally, with a distinct labellum; frontal

ectooecium finely tuberculate.

Etymology

Stomachosus (L.)—itritable, an allusion to the aggressive aspect of the

peristomial complex.

Remarks

The material comprised a single colony, 20 mm in diameter, with an

undulating, unilaminar growing edge and a thickened multilaminar central

region. The calcification is thick, particularly that of the basal walls; the basal

surface of the colony was apparently mostly unattached to the substratum, and

is encrusted with epizooites.

Measurements (means of 20 values) in mm

Lz Iz

OM 0,54

Rhynchozoon ptarmicum sp. nov.

Fig. 30

Material

Holotype: SAM-—A26453, station SM 179, 33°30,3’S 27°22,1’E, 80 m.

Other material: stations SM 185, SM 239, SM 250.

Description

Colony encrusting, multilaminar, forming extensive nodular sheets.

Zooids convex, large; frontal calcification thick and vitreous, becoming

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 30. Rhynchozoon ptarmicum sp. nov. A. Young zooids, showing the form of

the primary orifice. x 67. B. Later zooids, showing proliferation of avicularia and

orifices immersed in thickened calcification. x 50.

119

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

distinctly nodular in later ontogeny, marginal pores small. Primary orifice

slightly longer than wide; anter orbicular, with finely denticulate rim, poster

shallow, transversely elliptical; condyles large and conspicuous, triangular,

basally deflected. Four to six distal oral spines present in early ontogeny,

obscured by the development of a low thickened peristome, bearing a number

of short conical umbones. Suboral avicularium small, not enveloped by the

peristome and always distinct, rostrum at acute angle to frontal plane, directed

obliquely laterally, mandible short, broadly subtriangular; sporadically devel-

oped and frequently absent. No uncinate process. Additional avicularia often

abundant, of two types: small, adventitious, with a rounded cystid and short

semicircular or subtriangular mandible, developed along lateral borders of

zooid and encroaching on to frontal wall, often numerous, up to ten per zooid;

large, ?vicarious, with a swollen cystid supporting an elongate, triangular,

hooked rostrum (0,35 mm), with complete cross-bar and massive, quadrate,

bifid columnella. A slightly smaller version of the latter type (?adventitious)

may replace the suboral avicularium in some zooids. Ovicell elongate oval,

frontal surface granular, striated, labellum distinct; ooecial cover developed in

later ontogeny, often tuberculate.

Etymology

Ptarmos (Gr.)—a sneeze, an allusion to the open aspect of the orifice.

Remarks

Live colonies were collected from all four stations, the largest (SM 179)

was a massive, nodular, multilaminar growth measuring 70 mm by 35 mm. The

young colonies from SM 185 had a pale blue-grey coloration. The ?vicarious

avicularia resemble those of Strophiella tuberigera Jullien & Calvet (1903: 66,

pl. 9 (fig. 1)), reported from the Gulf of Gascony, from 135 m. S. tuberigera

has a ‘beaded’ orifice and is probably referable to Rhynchozoon; it differs

completely from R. ptarmicum in the shape of the primary orifice.

Measurements (means of 20 values) in mm

Lz Iz

0,73 Or52

Brodiella Uttley & Bullivant, 1972

Brodiella Uttley & Bullivant, 1972: 35.

Brodiella longispinata (Busk, 1884)

Fig. 31A-B

Schizoporella longispinata Busk, 1884: 163, pl. 17 (figs 2A, C).

Brodiella longispinata: Uttley & Bullivant, 1972: 36, fig. 26.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES IVA

Fig. 31. A-B. Brodiella longispinata (Busk). A. Three young zooids. B. Portion of a colony showing

ovicells and different types of avicularia. C—D. Brodiella ignota sp. nov. C. A typical zooid. D. Three

zooids showing variation in avicularia type. E-F. Turbicellepora conica (Busk). E. Portion of a colony

showing active frontal budding. F. Vicarious avicularia, and adventitious avicularia clustered on the

peristome rim of a brooding zooid. G-—H. Turbicellepora valligera sp. nov. G. Young zooids, showing

typical development of the peristome. H. Zooids from an older part of the colony, with numerous small

avicularia.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

Stations SM 185, SM 239.

Description

Colony encrusting, multilaminar. Zooids oval to hexagonal, heavily

calcified, separated by distinct grooves at first, boundaries later obscured by

secondary calcification. Primary orifice cleithridiate: anter orbicular, with a

denticulate rim, separated from the short rounded poster by prominent

condyles; operculum well chitinized, dark brown, with a distinct marginal

sclerite and a longitudinal median groove. Six distal oral spines, jointed basally,

of variable length, often greatly enlarged. Proximal side of orifice enclosed by a

low, thickened, peristomial rim, extending between the bases of the most

proximal pair of spines. Frontal wall thick, vitreous, with a characteristic

surface of rounded nodules imparting a rippled effect; marginal pores

infrequent and not readily visible. Avicularia adventitious, lateral to orifice,

single or paired, cystid low, tumid, rostrum elliptical, pivotal bar stout with a

thickened columnella, mandible semi-elliptical, directed laterally or obliquely

distally. Either or both avicularia may be replaced by an enlarged type with an

elongate, acute triangular mandible directed distally; alternatively or addition-

ally this enlarged avicularium may occur on the frontal wall, when the mandible

is directed proximally. Ovicell prominent, almost perpendicular to frontal plane

of zooid, oval, flattened frontally, with a short quadrate labellum above the

aperture and an uncovered frontal area of entooecium.

Remarks

The incidence of the different types of avicularia varies, and the larger

type may be more common than the small in some specimens. Both types are

seen in Busk’s original specimen (BMNH _ 1887.12.9.653), which in other

respects as well differs in no way from the Meiring Naude material.

Distribution

Brodiella longispinata was described by Busk (1884) from the Straits of

Magellan and has been reported subsequently from New Zealand and the

Chatham Islands (Uttley & Bullivant 1972). A synonymy of recent and fossil

New Zealand records is given by these authors.

Brodiella ignota sp. nov.

Fig. 31C-D

Material

Holotype: SAM—A26454, station SM 250, 31°59,3’S 29°22,5’E, 150-200 m.

Description

Colony encrusting, multilaminar. Zooids oval to hexagonal, large and

broad, convex, separated by distinct grooves. Primary orifice comprising an

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 123

orbicular anter, with finely denticulate rim, and a short, rounded poster below

blunt condyles. Six or seven slender, distal oral spines; proximally, a thin

peristomial rim encloses the orifice between the proximalmost pair of spines,

particularly developed in heavily calcified older zooids. Frontal wall thick,

vitreous and glistening, surface texture of densely packed rounded nodules;

marginal pores few in number, small and inconspicuous, generally visible only in

newly budded zooids. Avicularia paired, situated lateral to distal half of orifice;

cystid small, tumid, rostrum elliptical, directed laterally or medially, mandible

semicircular. Either or both avicularia frequently replaced by an enlarged type

with an elongate spatulate mandible directed distomedially, close to distal border

of orifice, reminiscent of Hippaliosina; palate with a small foramen almost

occluded by a thick columella. Rarely, one or both types of avicularia may occur

on the frontal wall. Small basal pore chambers present. Ovicells not found.

Etymology

Ignotus (L.)—unknown.

Remarks

The dimorphic avicularia, arising close to the primary orifice, the

denticulate anter, the peristomial rim and the characteristically thick, nodular

and vitreous frontal wall are all features of Brodiella, exemplified in the type

species B. longispinata (above). B. ignota differs from that species in the

proportions of the orifice, the shape and orientation of the enlarged avicularia

and in possessing more slender oral spines. D. P. Gordon (1979 in litt.) has

pointed out that the northern hemisphere species Lepralia armata Hincks

(Hayward & Ryland 1979: 206, as Buffonellaria) also belongs to Brodiella.

Measurements (means of 20 values) in mm

Lz Iz

0,83 0,61

Family Celleporidae Busk, 1852

Celleporidae Busk, 1852: 85. Hayward & Ryland, 1979: 274.

Turbicellepora Ryland, 1963

Turbicellepora Ryland, 1963: 34. Hayward & Ryland, 1979: 284.

Turbicellepora conica (Busk, 1884)

Fig. 31E—F

Cellepora conica Busk, 1884: 203, pl. 28 (fig. 10), pl. 36 (fig. 1).

Material

Stations SM 163, SM 163/164, SM 164, SM 179, SM 180, SM 184, SM 185.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Colony forming small nodules on erect substrata, pisiform, or branching in

one or more planes to give a stellate appearance. Zooids closely packed,

individual boundaries scarcely discernible. Primary orifice longer than broad;

anter suborbicular, poster deep, V-shaped, constituting one-third of total

orifice length. Peristome thin, tubular, incorporating proximolaterally a short

cylindrical avicularium; mandible semi-elliptical or subtriangular, acute to

frontal plane, directed obliquely laterally. Ovicell prominent, thinly calcified,

frontal surface closely punctured by numerous small round pores. Vicarious

avicularia of two types: elongate, broadly elliptical or slightly spatulate, with

thin cross-bar and large palatal foramen, small (0,2 mm long) and generally

infrequent; outnumbered by small vicarious avicularia, identical to suboral

type, with semi-elliptical mandible.

Remarks

The peristome tends to encircle the orifice completely, extending on to the

lateral regions of the ovicell. On older zooids the small avicularia tend to

multiply and typically become clustered around the rim of the deeply immersed

peristome, four or five commonly occur together. However, this very

characteristic feature seems to be present only in colonies in which growth by

frontal budding has slowed, in young colonies the continued budding of zooids

results in a surface composed of numerous cylindrical peristomes. The

multiplication of these small avicularia is thus apparently an ontogenetic, and

possibly gerontic, effect.

Turbicellepora conica is known only from Simon’s Bay, South Africa

(BMNH 1899.7.1.482, 483, 484, 486).

Measurements (means of 20 values) in mm

Lor lor

0,14 Oetat

Turbicellepora valligera sp. nov.

Fig. 31G—H

Material

Holotype: SAM-—A26455, station SM 163, 33°04,6’S 28°06,6’E, 90 m.

Other material: station SM 239.

Description

Colony developing from an encrusting base, forming a slender, erect

cylindrical growth. Zooids distinct, strongly convex and thickly calcified; frontal

wall fine-grained and smooth, with three or four very large marginal pores.

Primary orifice longer than broad, with a deep V-shaped poster comprising

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES WS

about one-third of the total length; hooded distally by a thin, erect peristomial

rim, the free edge of which is often peaked medially or produced into two or

more processes. Paired lateral oral avicularia; slender, cylindrical, with small

semicircular mandibles, linked distally by the peristome. Proximal to orifice a

conspicuous median umbo develops, often produced into an erect spike.

Vicarious avicularia of two types: broadly spatulate, cross-bar slender with a

thick median columella, palate with a small foramen; small, oval, with a

semicircular mandible, often abundant.

Etymology

Vallum (L.)—a palisade, referring to the enveloping peristome.

Remarks

This species is superficially similar to Osthimosia bicornis (Busk) from

which it differs in the position of the oral avicularia, which are proximolateral

in O. bicornis. Further, the ovicell of O. bicornis has the single central pore

characteristic of the genus; only a single partly immersed ovicell was found on

the material of T. valligera, but it was typical of Turbicellepora, being closely

punctured by small pores. The numerous small vicarious avicularia are also

seen in 7. pustulata (Busk), but this species is characterized by a single lateral

oral avicularium, and the absence of the columella on the cross-bar of the

spatulate avicularium.

Measurements (means of 20 values) in mm

Lor lor

0,15 Om

Celleporaria Lamouroux, 1821

Celleporaria Lamouroux, 1821: 43. Harmer, 1957: 663.

Celleporaria tridenticulata (Busk, 1881)

Fig. 19H

Cellepora tridenticulata Busk, 1881: 343, pl. 26 (fig. 9).

Celleporaria tridenticulata: Harmer, 1957: 670, pl. 42 (figs 510).

Material

Stations SM 162, SM 163, SM 163/164, SM 164.

Description

Colony encrusting, multilaminar, forming small nodules. Zooids oval,

convex, separated by distinct grooves; boundaries distinct in peripheral zooids,

later obscured. Primary orifice D-shaped, the straight proximal border with

three or four short conspicuous denticles, each of which may be bifid

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

terminally. Peristome scarcely developed, forming at the most a low rim around

the distal and lateral borders of the orifice; three or four short distal oral spines

present. Adventitious avicularium median suboral, cystid frequently columnar;

mandible short, semi-elliptical, typically directed transversely. Vicarious avicu-

laria sparsely developed, mandible narrowly spatulate, or subtriangular. Ovicell

prominent, oval, overarching much of the primary orifice; thinly calcified, with

a wide triangular frontal orifice.

Remarks

Celleporaria tridenticulata has been widely reported in the Indo-West-

Pacific region, from the Great Barrier Reef and New Guinea to Ceylon and

Mauritius. This is the first record of its occurrence in South African waters.

Celleporaria capensis (O’Donoghue & De Watteville, 1935)

Fig. 151-J

Holoporella capensis O’Donoghue & De Watteville, 1935: 203, pl. 5 (figs 9-10), pl. 6 (fig. 15).

Material

Stations SM 163, SM 163/164, SM 180, SM 185.

Description

Colony encrusting, developing multilaminar sheets and nodules. Zooids

oval, convex, thinly calcified; frontal calcification finely granular, translucent,

with a few inconspicuous marginal pores. Orifice large, subterminal, wider

than long, almost semicircular, proximal border slightly concave; two short

and thick, widely spaced, distal oral spines present in young zooids.

Peristome, when developed, forming simply a thin raised proximal lip,

occasionally overarching the orifice in older zooids; rarely, peaked medially to

form a short columnar umbo. Adventitious avicularia proximolateral to

orifice, mandible elongate, semi-elliptical, directed distally; one or two present

in newly budded zooids, later proliferating. Vicarious avicularia sporadically

developed, mandible elongate, semi-elliptical or slightly spatulate. No com-

plete ovicells found.

Remarks

C. capensis is known only from South Africa. Described originally from

Still Bay, southern Cape (O’Donoghue & De Watteville 1935), it was

subsequently reported from St. James and Oudekraal, south-western Cape

(O’Donoghue & De Watteville 1937), and from Port Elizabeth (O’Donoghue

& De Watteville 1944).

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 27

Family Vittaticellidae Harmer, 1957

Vittaticellidae Harmer, 1957: 765. Wass & Yoo, 1975: 286.

Vittaticella sp.

Fig. 11D

Material

Station SM 185.

Description

Zooids slender, clavate; small, less than 0,4 mm long, including proximal

node, daughter zooid of doubleton scarcely smaller than the mother zooid.

Primary orifice D-shaped, proximal border gently concave, a distinct lip

proximally. Frontal wall smooth, with scattered, minute, pores; vittae elongate,

narrow, extending from a point well proximal to the orifice, to close to the

proximal end of the zooid, each with a well-marked series of pores on each

side. Scapular chambers developed as prominent paired avicularia, with short

semicircular mandibles; usually symmetrical, except in daughter zooids in which

the inner, axial, avicularium is not developed. Suprascapular and infrascapular

chambers very small, closely united with avicularium and not conspicuous.

Ovicell not observed.

Remarks

The material was scanty, comprising part of a single small colony, and

insufficient for a more detailed morphological study. From its general features

this species seems to belong to the same group of species as the Indo-West-

Pacific V. tenella Harmer, V. venusta (MacGillivray), and V. praetenuis

(MacGillivray) (Harmer 1957: 776; Wass & Yoo 1975: 295), and could not be

readily identified with any species known from the South African region.

However, the vittaticellid fauna of the western Indian Ocean is poorly known.

Family Mamilloporidae Canu & Bassler, 1927

Mamilloporidae Canu & Bassler, 1927: 9. Harmer, 1957: 887.

Anoteropora Canu & Bassler, 1927

Anoteropora Canu & Bassler, 1927: 10. Harmer, 1957: 888.

Anoteropora latirostris Silén, 1947

Anoteropora latirostris Silén, 1947: 58, Pl. 5 (figs 25-27), figs 49-50. Hayward & Cook,

1979: 103.

Material

Stations SM 163/164, SM 185.

Description

Colony lunulitiform, large (up to 16 mm diameter), attached to sandy

sediments by numerous basal rhizoids with deep vertical walls; frontal shield

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

small, with an almost central orifice. Both autozooids and brooding zooids with

a large lateral avicularium, with curved, acute mandible.

Remarks

Eight complete colonies and fragments of several others were collected.

Two of the five colonies from station SM 185 were alive at the time of

collection and have intact basal rhizoids.

Distribution

Indo-West-Pacific.

ORDER CYCLOSTOMATA

Fragments of numerous species of Cyclostomata were present in the

bottom sediments of most of the stations reported upon here, and were

especially abundant in those from SM 163/164 and SM 129. Many of the

specimens were of dead and worn colonies, evidently transported, including

species of Hornera, Tubulipora, and ?Tervia. The free tubular parts of

peristomes and the brooding gonozooids are absent or damaged in most

specimens rendering identification difficult. Although at least 20 distinct colony

growth forms may be recognized, only 16 taxa have been identified, often

doubtfully, to genus or species level. More than 40 species of cyclostomata have

been reported from the South African region by Busk (1867, 1875, 1886),

O’Donoghue (1924), O’Donoghue & De Watteville (1935, 1944) and Brood

(1976b). Of these only 9 species have been found in the present Meiring Naude

collections, although 1 additional form, Crisia aff. holdsworthii, was listed by

Hayward & Cook (1979: 116). One notable absentee is Tennysonia stellata

Busk (1867: 242, pl. 36 (figs 10-11)), which was first reported from South

Africa, and has large, robust colonies. Another South African species, —

Disporella buski Harmer (1915: 161, pl. 12 (figs 4-5)), is known to have a wide

distribution but is also absent from the present collections.

Generally, nominal species, at least, have very wide distributions and

some species have been identified, both here and by Brood (1976b), with forms

reported from the Philippines by Canu & Bassler (1929), from Indonesia by

Harmer (1915), and from Australia by MacGillivray (1885). Examination of the

type suites of some species, for example those described by Busk (1875, 1886)

in the British Museum Catalogue and the Challenger Report respectively, has

revealed, however, a wide range of variation within populations of nominal

species. Some of these variants may even prove to be specifically distinct. It is

obvious that revision of the South African cyclostome fauna will require

detailed analysis of well-preserved, preferably living, colonies. Harmelin

(1976b) has demonstrated that within the Mediterranean Tubuliporina many

colony forms, and even zooid morphologies, are radically modified by

environmental influences, and that convergence of character states can occur

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Wy)

Fig. 32. A. Idmidronea contorta (Busk). X 16. B. Mecynoecia clavaeformis (Busk). X 17.

C_-D. Idmidronea crassimargo (Canu & Bassler). C. Two fragments of a colony, including

a gonoecium. X 26,3. D. Detail to show ooeciostome. xX 62.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

among systematically unrelated species. Until such a detailed study may be

made, it is pointless to attempt to revise specific and generic concepts, or to

assign very worn fragments to previously described species. In assigning the

present material to particular species it is not implied that the specimens have

been compared with type material, unless so stated; rather, it is considered that

the Meiring Naude fragments closely resemble the specimens illustrated by the

authors listed in the synonymies.

Family Diaperoeciidae Canu, 1918

Diaperoeciidae Canu, 1918: 329.

Diaperoecia Canu, 1918

Diaperoecia Canu, 1918: 329. Harmelin, 1976b: 78.

Diaperoecia sp.

Material

Station SM 163.

Remarks

A single colony, lacking gonozooids, was found encrusting a worn, erect

cheilostome bryozoan.

Family Entalophoridae Reuss, 1869

Entalophoridae Reuss, 1869: 285.

Mecynoecia Canu, 1918

Mecynoecia Canu, 1918: 326. Harmelin, 1976b: 160.

The characters of the genera Pustulopora and Mecynoecia have been

discussed by Brood (1976a, 1976b) and Harmelin (1976b).

Mecynoecia clavaeformis (Busk, 1875)

Fig. 32B

Pustulopora clavaeformis Busk, 1875: 22, pl. 14 (figs 1-4).

Material

Stations SM 131, SM 163/164.

Description

Colonies club-shaped, up to 20 mm high, arising from a small, encrusting

base. Zooids in irregular closely spaced whorls of three to six.

Remarks

Although somewhat worn, the fragments agree exactly with the material

from Algoa Bay (BMNH 1875.5.29.34, 1899.7.1.530) described by Busk (1875).

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 131

Busk’s specimens were also fragmentary and provide little information as to the

later growth form of the colony and the systematic relationships of the species.

Mecynoecia delicatula (Busk, 1875)

Pustulopora delicatula Busk, 1875: 21, pl. 6B (fig. 3). Brood, 1976b: 290, fig. 13E-F.

Mecynoecia delicatula: Harmelin, 1976b: 160, pl. 27 (figs 1-8).

Material

Stations SM 129, SM 163/164, SM 185.

Description

Colonies erect, occasionally branched, with long zooids opening irregu-

larly on all sides. Free peristomes long and curved. Gonozooids often at

bifurcations, simple.

Remarks

Most specimens were slightly worn, and lacked the long peristomes.

Brood (19766) noted that this species was common from 50 to 700 m in east

African waters.

Mecynoecia australis (Busk, 1852)

Pustulopora australis Busk, 1852: 350; 1875: 21, pl. 17A. Brood, 1976b: 291, fig. 13A.

Pustulopora proboscidea: Busk, 1886: 19, pl. 4 (fig. 2).

Material

Stations SM 129, SM 151, SM 163/164, SM 180, SM 185.

Description

Colonies similar to those of M. delicatula but with larger zooids and

coarser calcification.

Remarks

Brood (1976b) reported M. australis as rare in east African waters, with a

bathymetric range of 75 to 700 m.

Distribution

Indo-Pacific.

Family Diastoporidae Busk, 1859

Diastoporidae Busk, 1859: 91, 113. Harmelin, 1976b: 119.

Plagioecia Canu, 1918

Plagioecia Canu, 1918: 327. Harmelin, 1976b: 128.

Plagioecia patina (Lamarck, 1816)

Tubulipora patina Lamarck, 1816: 163.

Plagioecia patina: Harmelin, 1976b: 129, pl. 8 (figs 5-9), pl. 18 (figs 4-9), pl. 19 (figs 1-5).

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

Station SM 163/164.

Description

Colonies forming flattened discs, often with an extended peripheral lamina

and small subcolonies. Early budding fan-shaped, later zooids in repent,

connate, single, radial rows. Gonozooids peripheral, transversely extended,

ooeciostomes central.

Remarks

Two colonies were found, one of which was alive when collected.

Distribution

Temperate Atlantic, Mediterranean, and eastern Pacific.

Liripora MacGillivray, 1887

Liripora MacGillivray, 1887: 182.

Desmeplagioecia Canu & Bassler, 1920: 718. Brood, 1976b: 284.

MacGillivray (1887) introduced Liripora for two Recent Australian

species, L. lineata (see below) and L. fasciculata, both of which had been

originally assigned to Diastopora in an earlier paper (MacGillivray 1885). L.

fasciculata, together with several fossil species included in the genus by

MacGillivray (1895), is probably generically distinct from L. lineata. However,

as the first species described in 1885 and the first listed in 1887, L. lineata may

be regarded as the type species of Liripora, and Canu & Bassler (1920) seem to

have introduced Desmeplagioecia, for D. lineata, unnecessarily. Canu

(1908: 310) used the name Liripora incorrectly for flabellate species of

Tubulipora.

Liripora is obviously closely related to Plagioecia, being distinguished

principally by its multiple radial rows of connate zooids. The genus is also very

similar to the fossil Actinopora d’Orbigny (see Brood 1976b: 284).

Liripora lineata (MacGillivray, 1885)

Diastopora lineata MacGillivray, 1885: 96, pl. 3.

Liripora lineata: MacGillivray, 1887: 182.

Berenicea lineata: Harmer, 1915: 116, pl. 11 (figs 6-7).

Desmeplagioecia lineata: Canu & Bassler, 1920: 718, fig. 234. Brood, 1976b: 284.

Material

Stations SM 163/164, SM 185.

Description

Colony encrusting, discoid, with a basal lamina; forming subcolonies by

frontal extension. Primary zooids repent, budding distally and laterally. Later

zooids radially disposed; peristomes connate, raised terminally, becoming

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 133

multizooidal. Areas between zooid rows calcified but not reticulate. Gono-

zooids peripheral and subperipheral, with distal ooeciostomes.

Remarks

Three young colonies, and a single group of four subcolonies were found;

all were alive when collected. One further colony, on the concave side of a

lamellibranch shell, had three gonozooids, and was dead. The subcolonies are

formed by frontal extension of one or more peripheral zooids of the mother

colony, which forms a new basal lamina. Some of the Australian colonies

examined (BMNH 1897.5.1.1377, 1963.2.12.19.) are more robust than those

from South Africa, but others are almost exactly the same in all characters.

A similar species, Diastopora reticulata, described from the Antarctic by

Borg (1944: 68, pl. 4 (figs 3-8)), has reticulate interzooidal areas and a tubular

ooeciostome.

Distribution

Indo-Pacific, including Japan and Australia.

Family Tubuliporidae Johnston, 1838

Tubuliporidae Johnston, 1838: 247. Harmelin, 1976b: 165.

Idmidronea Canu & Bassler, 1920

Idmidronea Canu & Bassler, 1920: 784. Harmelin, 1976b: 181. Buge, 1979: 232.

Harmelin (1976) discussed the character of Idmidronea and related

genera, and noted the difficulties in distinguishing species, due to environmen-

tally induced variation between populations of single species. Six different

kinds of colony belonging to Jdmidronea were distinguished in the Meiring

Naude collections. Although these have been identified with known species, it

should be noted that several of these were originally described from one or two

fragments only, which may have represented parts of a single, variable species.

Idmidronea contorta (Busk, 1875)

Fig. 32A

Idmonea contorta Busk, 1875: 12, pl. 8. O’Donoghue, 1924: 24.

Tennysonia contorta: Canu & Bassler, 1922: 52, pl. 11 (figs 11-14); as Lobosoecia semiclausa,

in error, on p. 145.

Material

Stations SM 129, SM 131, SM 163/164, SM 185.

Description

Branches short, inflated, curved; flat basally. Zooids connate, in

alternating groups of six. Gonozooids occurring at bifurcations.

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The fragmentary specimens are closely similar to those from Algoa Bay

described by Busk (BMNH _ 1875.5.29.18). The gonozooid has not been

described before, but the specimens are worn and the ooeciostome is not

obvious. J. contorta was recorded by O’Donoghue (1924) from several South

African localities, from depths of 55-169 m. Harmelin (1976b) noted that

Mediterranean records of this species are referable to Tubulipora notomale

Busk (1875), which has much larger zooids.

Idmidronea crassimargo (Canu & Bassler, 1929)

Fig. 32C-D

Idmonea crassimargo Canu & Bassler, 1929: 545, pl. 85 (figs 2-3).

Idmidronea crassimargo: Brood, 1976b: 290, fig. SI-J, M.

Material

Stations SM 163/164, SM 185.

Description

Colonies with basal side of branches curved, zooids in alternating, connate

groups of three. Gonozooid short, inflated, frequently placed at a bifurcation;

ooeciostome small, rounded with a short tube, central or lateral.

Remarks

Originally described from the Philippine Islands, from 320 m, this species

was also recorded from several east African localities by Brood (1976b)

between 60-150 m. Several of the Meiring Naude specimens were alive when

collected; the gonozooids have not been figured before.

Idmidronea cf. parvula (Canu & Bassler, 1929)

Fig. 33A

Idmonea parvula Canu & Bassler, 1929: 546, pl. 85 (fig. 1).

Material

Stations SM 163/164, SM 185.

Description

Branches with flat basal side. Zooids in alternating connate groups of two

or three.

Remarks

Several fragments of this small species were found, two of which had

incomplete zigzag gonozooids. The zooids are distinctly smaller than those of J.

crassimargo, and larger than those of J. cf. biporata (below). The gonozooids

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 135

Fig. 33. A. Idmidronea c.f. parvula (Canu & Bassler). x24. B-D. Hornera erugata sp. nov.

B. Frontal view of a branch, including a gonoecium. X 26,7. C. Detail to show ooeciostome. x 50.

D. Basal view of colony and gonoecium. x 17,8.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

are not inflated like those of J. biporata, but it is possible that all of these small

forms, including J. cf. antarctica (below) are either very closely related

systematically, or represent ecologically varying populations of the same

species.

Idmidronea cf. biporata Brood, 1976

Idmidronea biporata Brood, 1976b: 290, fig. SF—-H.

Material

Stations SM 163/164, SM 185.

Description

Zooids very small, in alternating pairs, gonozooids inflated.

Remarks

Only three fragments were found. the zooids are minute, even smaller

than those figured by Brood (1976b).

Idmidronea cf. antarctica Borg, 1944

Idmidronea antarctica Borg, 1944: 84, pl. 6, (fig. 4), pl. 7 (figs 1-2).

Material

Stations SM 129, SM 131, SM 163/164.

Description

Branches flat basally; zooids in closely spaced, alternating, connate groups

of five. Gonozooids forming a zigzag ridge frontally.

Remarks

Only four minute fragments were found. The zooids are smaller than

those figured by Borg (1944), but the gonozooid closely resembles that of J.

antarctica, which also has five zooids in each group.

Idmidronea cf. atlantica (Forbes in Johnston, 1847)

Idmonea atlantica Forbes in Johnston, 1847: 278.

Idmidronea atlantica: Harmelin, 1976b: 182, pl. 32 (figs 1-11). Hayward & Cook, 1979: 116.

Buge, 1979: 232, pl. 7 (fig. 4).

Material

Stations SM 129, SM 163/164.

Remarks

Numerous worn fragments were found, none of which had complete

gonozooids. They are provisionally assigned to this widely distributed and very

variable species.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 37)

Family Crisiidae Johnston, 1847

Cristidae Johnston, 1847: 282.

Crisia Lamouroux, 1812

Crisia Lamouroux, 1812: 183. Ryland, 1967: 272.

Crisia elongata Milne Edwards, 1838

Crisia elongata Milne Edwards, 1838: 203, 235, pl. 7 (fig. 2). Busk, 1875: 5, pl. 4 (figs 5-6).

Harmer, 1915: 103, pl. 8 (figs 13-17). Brood, 1976b: 282, fig. 4F, J, N.

Material

Stations SM 123, SM 163/164.

Description

Colonies with elongated, almost straight internodes, consisting of ten to

twenty alternating zooids. Peristomes curved forward, short. Basis rami short,

wedged in between two zooids, branches usually occurring high in an

internode. Gonozooid dilated distally, ooeciostome slit-like, without a raised

rim. Joints black.

Remarks

The material comprised numerous well-developed living colonies with

extensive kenozooidal rooting systems. The joints are dark yellow early in

ontogeny, rapidly becoming black. No gonozooids were present. The colonies

are slightly less robust than Busk’s colonies from Algoa Bay (BMNH

1875.5.29.5), but are closely similar to Brood’s (1976b) figures of east African

material. C. elongata differs from C. transversata Brood (1976b: 282, fig. 4E,

G-I), which also has black joints, in having shorter zooids, longer and

straighter internodes, less prominent peristomes and a longer gonozooid. C.

holdsworthii Busk, which was reported from the earlier Meiring Naude

collections, has much longer, more delicate zooids, and relatively colourless

joints.

Distribution

Indo-Pacific, including Japan and Australia.

Family Lichenoporidae Smitt, 1866

Lichenoporidae Smitt, 1866: 404, 474.

Lichenopora Defrance, 1823

Lichenopora Defrance, 1823: 257.

Lichenopora novae-zealandiae (Busk, 1875)

Discoporella novae-zealandiae Busk, 1875: 32, pl. 30 (fig. 2).

Lichenopora novae-zealandiae: Harmer, 1915: 155, pl. 12 (figs 6-11). Brood, 1976b: 299,

fig. 17H-I.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

Stations SM 131, SM 163/164, SM 184, SM 185.

Description

Colonies discoidal, raised centrally, with almost vertical sides. Zooids in

radial, connate rows, the ends of the peristomes not free. Gonozooids central,

surrounded by extrazooidal calcification; ooeciostomes large, rounded.

Remarks

Sixteen colonies were found, five of which were alive when collected. L.

novae-zealandiae differs from Disporella buski Harmer, originally described (as

D. ciliata Busk, 1875) from the Cape of Good Hope, in its regularly radial,

connate zooid series, which are not free terminally (see Brood 1976b: 299,

fig. 17D-G; Buge 1979: 242, pl. 9 (fig. 1)).

Distribution

Indo-Pacific, including Japan, Australia and New Zealand.

Family Crisinidae d’Orbigny, 1853

Crisinidae d’Orbigny, 1853: 902. Borg, 1941: 35.

Crisina d’Orbigny, 1850

Crisina d’Orbigny, 1850: 265; 1853: 912. Borg, 1941: 2.

Crisina radians (Lamarck, 1816)

Retepora radians Lamarck, 1816: 183.

Crisina watersi Borg, 1941: 16, pl. 2 (figs 3-4), pls 3-4.

Crisina radians: Brood, 1976b: 297, fig. 1SH-J.

Material

Station SM 131.

Description

Colonies profusely and regularly branched in one plane; basal surface of

branches flat, composed of kenozooids. Zooids in alternating connate groups of

three to four.

Remarks

Two bifurcating fragments were found, one of which bore a gonozooid.

Brood (1976b) considered that the east African material described as C. watersi

by Borg (1941) was within the range of variability displayed by C. radians.

Distribution

Crisina radians is widely distributed in the Indo-West-Pacific region, from

east Africa to Australasia, and also in New Zealand waters.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 139

Family Horneridae Smitt, 1866

Horneridae Smitt, 1866: 404, 465. Gregory, 1899: 360. Borg, 1926: 385.

Hornera Lamouroux, 1821

Hornera Lamouroux, 1821: 41. Harmer, 1915: 147. Borg, 1926: 204, 385.

Hornera erugata sp. nov.

Fig. 33B—D

Hornera sp. Cook, 1968: 238.

Material

Holotype: BMNH 1842.11.30.45, Cape of Good Hope.

Paratypes: SAM—A26456, as above, Cape of Good ves

Other material: station SM 163.

Description

Colony erect, branching; branches curved in more than one plane. Zooids

in laterally contiguous series of three to seven; peristomes long, often bifurcate

in lateral zooids. Secondary branches formed by fascicles of three to four

zooids. Basal extrazooidal calcification smooth, without obvious pores. Gono-

zooid basal, surface smooth; ooeciostome large, slit-like, lateral, closely

opposed to the side of the branch, without a raised rim.

Etymology

Erugatus (L.)—smooth, referring to the basal calcification.

Remarks

Of the numerous worn fragments of Hornera present in the Meiring

Naude bottom samples (p. 142), only three, relatively unworn, had the smooth

basal calcification of H. erugata. The basal calcification of other species of

Hornera is invariably corrugated, with large pores, pits and intervening ridges.

The lack of corrugations on the gonozooid of this species is also unusual, as is

the absence of a rim to the ooeciostome.

A somewhat similar species, which has ridged, porous basal calcification

and a reticulate gonozooid with a raised, lateral ooeciostome, was described

from the Antarctic as H. smitti by Borg (1944: 199, pl. 15 (figs 2-8)).

Pseudidmonea gracilis Androsova (1965: 80, fig. 18), another Antarctic species,

has a gonozooid somewhat like that of H. erugata, but a more delicate colony,

with alternating series of three to five connate zooids.

Measurements (range of 10 values) in mm

Lz Lg lg Loo.st.

0,40,5 2,5—3,0 2,0-2,5 0,4-0,55

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

DISCUSSION

General review of collection

The particular importance of the material described in the preceding

account derives from the fact that it was collected from a range of depths for

the most part far shallower than those from which the first series (1975-6) of

Meiring Naude samples was taken. The bryozoan collections from the first two

cruises originated from stations further to the north and east than those of

subsequent cruises, and could be characterized largely as typical continental

slope assemblages (Hayward & Cook 1979). The present collections are from a

number of stations established off-shore between Durban and East London, the

majority of which were located on the continental shelf of this region. Thus, 9

of the samples reported upon here were collected from depths of less than

200 m, 6 range between 500 m and 700 m, and only 3, 1 of which (SM 103)

represents a residue from the 1976 cruise, were obtained from depths in excess

of 700 m. The greatest depth sampled for bryozoans in the later survey was

station SM 151, at 900 m. By contrast, the greatest depth represented in the

first series of samples (Hayward & Cook 1979) was 1300 m, with the majority

of the material originating from between 600m and 1000m. It is not

surprising, then, that the bryozoan faunas revealed in each case should differ

markedly.

The first report described 51 species, 48 of which were cheilostomata; the

number of anascan species was only a little smaller than the number of

ascophora and included several which were considered to be particularly

adapted to life on the fine, unconsolidated sediments of the continental slope,

such as the small colonies of Setosellina, Heliodoma and Inversiscaphos. There

were also representatives of cellularine genera, for example Columnella,

Notoplites, and Bugulella, frequently associated with slope faunas. The

ascophora, similarly, included a number of highly specialized species in the

genera Anoteropora, Batopora and Lacrimula. The present collection offers an

immediate contrast in both the total number of taxa recorded, 130, and in the

increased proportion of ascophorans. There were almost twice as many

ascophorans (Table 2) as anascans (Table 1), and these were predominantly

encrusting forms, together with a significant proportion with erect, rigid

colonies, most notably the species of Adeonella. The anascans included a range

of encrusting species, and a number of cellularines, such as Menipea, typically

associated with shallow water faunas. A further difference is seen in the

substantial number (sixteen) of Cyclostomata identified in these samples; with

two exceptions (Table 3) live specimens of cyclostome species were recorded

only at stations with depths of less than 100 m.

Only twenty-two of the species described here had been reported in the

first Meiring Naude collections. For the most part these comprised species

which occurred only at stations deeper than 500 m (e.g. Tessaradoma spp.), or

which were represented in the first collections by specimens from the shallowest

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 141

stations only (e.g. Flustramorpha angusta, Reteporella dinotorhynchus), or

which appear to have very broad bathymetric distribution patterns (Setosellina

roulei). Thus, the two collections complement each other most usefully, and

together provide a representative survey of the shelf and slope faunas of

eastern South Africa. The results of the Meiring Naude cruises, together with

the accounts published in the century following the Challenger investigations,

provide a sound base line for future research on the bryozoan fauna of this

region.

Including the new species described here, more than 280 nominal forms of

Bryozoa have been reported from South Africa. Descriptions of these may be

found in Busk (1852, 1854, 1884), Marcus (1922), Harmer (1926, 1934, 1957),

the papers of O’Donoghue and O’Donoghue & De Watteville (q.v.), and a

very few other sources. A precise enumeration is not yet possible, largely

through outstanding problems of synonymy. Many South African forms were

originally assigned the names of European species and, while it has been

possible to review the systematic status of some of these, others will perhaps

never be recognized. O’Donoghue (1924), for example, noted that both

Cellaria fistulosa (Linnaeus) and ‘Lepralia’ foliacea (Ellis & Solander) had been

reported from South Africa, yet in view of the known geographical distribution

of these two species (Ryland & Hayward 1977; Hayward & Ryland 1979) this

seems improbable, and there is no way of ascertaining to which species these

names were applied. Despite such problems, some useful synonymizing has

been achieved by previous authors. Caberea boryi (Audouin), listed by Busk

(1852) from Algoa Bay, may now be recognized as C. darwinii Busk

(O’Donoghue 1924), and all records of Steginoporella magnilabris (Busk) may

be assigned to S. buskii Harmer (O’Donoghue 1957). Several names introduced

for South African bryozoans have proved to be junior synonyms of previously

described species; for example Bicellariella capensis O’Donoghue is B.

chuakensis Waters (O’Donoghue 1957), and Beania paucispinosa O’Donoghue

& De Watteville is B. vanhoffeni Kluge (O’ Donoghue 1957).

Research on the Meiring Naude samples has allowed some opportunity for

revision and reassessment of previously known species. For example, Hasen-

bank’s (1932) record of Beania erecta Waters is here redescribed as B. rediviva

sp. nov. However, although it is clear that much similar revisionary work

remains to be done, only 51 of the species described in this report have been

described before from South African waters; with the exception of B. rediviva,

the rest are either new species or newly reported for the region. For the slope

fauna (Hayward & Cook 1979) 44 out of a total of 51 species belonged to these

latter two categories. Thus, the bryozoan fauna of both the shelf and slope of

eastern South Africa has a far greater diversity than was hitherto suspected and

it is probable, from evidence discussed below, that its complete diversity is as

yet uncharted. The fauna now known has a taxonomic diversity comparable to

those of other, more widely studied, areas of continental shelf seas. For

example, Cook (1983b) has listed 222 species from the shelf waters of west

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Africa, and the cheilostome fauna of the British sea area (Ryland & Hayward

1977; Hayward & Ryland 1979) totals 186 species. The Meiring Naude

investigations revealed a very high proportion of formerly undescribed

cheilostomata which, together with the considerable numbers of apparently

rare species also collected, suggests that the eastern South African fauna

includes a substantial endemic element. It is likely that further surveys, with a

coverage as intense as that accomplished by Millard (1975, 1978) for the

hydroids, encompassing the whole of southern Africa will greatly enhance the

presently known diversity of bryozoan species.

Analysis of bottom deposits

Two immediate impressions are gained from initial examination of the

abundant sediment samples collected from a number of the Meiring Naude

stations. Firstly, there is an almost complete absence of the minute ‘sand fauna’

species discussed in the previous report (Hayward & Cook 1979); secondly, a

high proportion of the sediment at some stations consists of bryozoan skeletai

remains.

The reasons for the virtual absence of such genera as Batopora, Lacrimula

and Heliodoma, which were so abundant in the samples from stations

SM 16-SM 109 (see Hayward & Cook 1979) are not obvious, and do not

appear to be related directly to depth. These specialized forms were found in

the first series of samples from depths of 376 to 1300 m. Samples from similar

depths were included in the present collections, yet H. implicata was frequent

only in the residue of station SM 103, and none of the ascophoran genera, with

the exception of Anoteropora, were found. The range of sediment particle size

seemed essentially very similar in both series of samples, and although, as

stated above, the stations of the 1977-9 surveys were situated further south

than those of the 1975-6 cruises, other environmental parameters, such as

bottom temperature, appear to have the same range with depth in both

collections (Louw 1977, 1980).

An analysis of sediment types for ten stations for which samples were

available is given in Table 4. The data comprise the approximate proportions

of different constituents in a single sample for each station, graded in three

coarse size fractions. The number of bryozoan species identified is given for

each station including the number present as dead material only. There is

considerable variation in sediment size between stations; for example, for

SM 184 40 per cent of the sample was retained by the 5mm sieve, while

nothing passed the 1 mm sieve, whereas for SM 185 48 per cent of the sample

passed the 1 mm sieve and nothing was retained by the 5 mm sieve. Inspection

of the table reveals no apparent correlation between bryozoan diversity and

sediment size.

The proportion of biogenic sediments in each size fraction also varies

widely from station to station and again suggests no correlation with bryozoan

diversity. Shell fragments constituted from 10 to 50 per cent of each sample,

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 143

often in the two smaller size fractions; these were largely finely comminuted

molluscan shell, although remains of echinoderm test were frequent, and the

coarsest fragments often included a significant proportion of decapod shell. The

most interesting feature of this table is the significant contribution to the

sediment made by bryozoan skeletal remains at several stations, viz. SM 129,

SM 131, SM 151, SM 163/164, SM 179. At SM 179, in particular, bryozoan

remains comprised 50 per cent of the sample. Most of the fragments were of

erect genera, such as the cheilostome$ Flustramorpha, Adeonella and Sertella,

and the cyclostomes Mecynoecia, Idmidronea and Hornera. Three subsamples

from station SM 163/164, each of 1 cm? volume when dried, were studied in

greater detail; an average of 400 bryozoan fragments was found per sample, of

which 25 per cent were cyclostomata and 75 per cent cheilostomata. The total

sediment sample for this station was estimated to contain 18000 bryozoan

fragments; 56 species were recorded for this station, including 24 that were

represented by dead fragments only. The high species diversity and abundance

of bryozoans in this region is thus reflected in the constitution of the bottom

deposits; similar results were found by Wass et al. (1970) in a study of the

Southern Australian shelf and slope. However, Table 4 also emphasizes the

problems encountered in attempting to describe patterns of geographical and

bathymetric distribution, and the need to differentiate carefully between living

and dead, or attached and loose, specimens in benthic samples. A number of

the species described above were represented by dead material only, and their

real distribution remains unknown. The proportion of species represented by

dead specimens only increases with depth; the stations in Table 4 with a

high content of bryozoan remains in the sediment range in depth from 80 m

(SM 179) to 850m (SM 129). It is likely that each includes a different

proportion of transported material while none probably represents a true

thanatocoenosis. The proportion of very worn material varied from station to

station; for example, 30 per cent of the skeletal material from SM 163/164

was too abraded to be identified, and 70 per cent of that at SM 179 was

similarly unidentifiable. While some specimens could not even be readily

assigned to a family grouping, others could be recognized as species, or even

genera, which were not otherwise recorded in these collections, but which

could not be reliably characterized. For example, abraded internodes of a

species of Margaretta were frequent in several of the sediment samples. The

last three stations in Table 4 (SM 180, SM 184, SM 185) each had a high

diversity of bryozoan species, including a significant proportion of dead

material, yet bryozoan skeletal remains were negligible in, or absent from,

the sediments from these stations. In these three cases the living material

included several cellularine species and a substantial number of large erect

species (Adeonella, Gigantopora) encrusted with numerous other cheilos-

tomata. Although the removal of large dead fragments from these samples

prior to analysis undoubtedly biases the results, the lack of finer skeletal

remains and the predominance of erect branching species probably reflect

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

local hydrographic conditions which are not conducive to the accumulation

of bryozoan deposits.

Colonies of Dactylostega prima and Chaperia familiaris, particularly from

stations SM 162 and SM 180, were often frequent on the tests of large

specimens of the rhizopod foraminiferan Schizammina pinnata (Pearcey). This

foraminiferan is known only from the south-eastern coast of South Africa

(between 57 and 228m), where it is frequently, although sporadically,

abundant. The family Schizamminidae was introduced by Né@rvang (1961) for

TABLE 4

Analysis of sediment components (%) from ten Meiring Naude stations.

A=<1mm fraction; B=1-5 mm fraction; C=>5 mm fraction; T = terrigenous

material; S=shell; FS =foraminiferans with sand accreted tests; FC = foraminiferans

with calcareous tests; BR = bryozoan fragments.

Bryozoan spp.

Station Depth m r S FS FC BR Total Dead

SM 103 680 A 20 0 50 0 0 7 6

B 5 10 10 0 2

Cc 3 0 0 0 0

SM 123 690 A 10 0 0 10 0 2 0

B 0 10 0 20 0

C 10 10 10 20 0

SM 129 850 A DD 5 2 3 0 7 7

B 5 10 10 0 10

C 10 5 5 0 10

SM 131 780 A 10 20 1 1 5 22 21

B 0 25 1 0 25

Cc D 5 0 0 5

SM 151 900 A 0 5 5 0 ») 8 8

B 0 20 10 0 15)

C 0 10 10 0 20

SM 163/164 90 A 15 0 0 5 0 67 24

B 0 20 0 5 30

‘C 0 10 0 0 115)

SM 179 80 A 5 5 0 0 10 24 4

B 5 5 0 0 10

C 15 IS 0 0 30

SM 180 80 A 25 20 5 0 0 Di, Dh

B 20 20 5) 5 0

Cc 0 0 0 0 0

SM 184 86 A 0 0 0 0 0 23 12

B WS) 30 5 0 0

€ 15 20 5 0 0

SM 185 90 A 20 25 3 0 D 5 iS)

B 20 25 3 0 a,

C 0 0 0 0 0

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 145

two genera of large Foraminifera, Schizammina and Jullienella, some species of

which may grow to 70 mm in height or diameter, and which have dark or

reddish-brown, rigid tests composed of agglutinated sand grains. S. pinnata

(N¢rvang, 1961: 192, pl. 7 (figs 9-17)) is robust, elongated, and has regularly

alternating, short, sympodial branches, which form a zigzag pattern. Another

South African species, Jullienella pearceyi Ngrvang (1961: 198, pl. 9 (figs 1-11,

14)), is smaller, more inflated and almost polygonal in shape. In west African

waters the large fimbriated plates of J. foetida Schlumberger (N@rvang,

1961: 195, pl. 8 (figs 1-13)) form the major substratum in some areas for a

diverse fauna of bryozoan species (Cook 1968, 1983b). The mode of life of the

foraminiferans is not known, but they are frequently covered on both surfaces

by encrusting bryozoan colonies. Worn and fragmented specimens often form a

significant proportion of bottom sediments (Table 4).

Geographical distributions

In Table 5 known geographical ranges are given for seventy-three

previously described species. Fifty-one of these have been reported before from

South Africa. The twenty-two species here recorded for the first time include a

number which appear to have generally broad geographical distribution

patterns, such as Parasmittina tropica, Escharina pesanseris, Trypostega

venusta, Celleporaria tridenticulata and Crisina radians. Several others, such as

Cribrilaria innominata and Cleidochasma porcellanum, are known to be

complex variable entities in which apparently very broad distributions may be

simply artefacts of systematic uncertainty. However, at least half of these

twenty-two species are characterized by less broad distributions suggestive of

more interesting patterns. Hippoporidra senegambiensis has an otherwise rather

limited distribution on the west African shelf; Cupuladria multispinata,

Setosellina roulei, Heliodoma implicata, and perhaps Cribrilaria venusta (p. 43)

appear to show similarly narrow distributions, but in total these species

constitute a minority in the eastern South African fauna. Caberea darwinii and

Brodiella longispinata seem to be more widely distributed, cold-temperate

Southern Hemisphere species, probably at the extreme northern limit of their

range in South African waters. This pattern is seen in another species,

Amphiblestrum inermis, long known from South Africa. However, the

arrangement of Table 5 emphasizes the accepted faunal similarity of eastern

South Africa with the Indo-West-Pacific realm and, like the previously reported

species, these new records are predominantly of species with distributions

extending through part or whole of this realm. Escharina waiparaensis,

formerly known only from New Zealand, and Hippomenella avicularis,

described from a few stations in the western Pacific, represent particularly

significant range extensions; the distribution of Hippoporella spinigera is

established through the inclusion in its synonymy of Mucronella serratilabris

O’Donoghue (1924). Crassimarginatella marginalis seems to have a narrow

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 5

Geographical distributions of previously described species.

Malay

Atlantic West South East Indian Archi- West East New Sub- Antarctic

Africa Africa Africa Ocean pelago Pacific Pacific Zealand antarctic

Carbasea mediocris

Cupuladria multispinata x

Discoporella umbellata x

Setosellina roulei x

Heliodoma implicata x

Amphiblestrum inermis

Crassimarginatella marginalis x

Foveolaria imbricata

Chaperia multifida ?

Chaperia capensis

Chaperia stephensoni

Notocoryne cervicornis

Steginoporella buskii x x

Cellaria tectiformis

Cellaria punctata

Cellaria paradoxa

Caberea darwinii x x x x

Eupaxia quadrata

Menipea crispa

Menipea triseriata

Menipea ornata

Menipea marionensis

Bugulella australis

Beania magellanica x x

Bugula dentata x

Cribrilaria innominata x

Cribrilaria venusta x x

Figularia philomela

Escharoides contorta x

Pachycleithonia mutabilis x x x

Tropidozoum burrowsi x x

Parasmittina tropica x x x x x

Porella capensis x

Arthropoma cecilii x x x x x x x

Arthropoma circinatum x

Escharina pesanseris x x x x x x

Escharina waiparaensis x

Calyptotheca nivea x

Calyptotheca porelliformis x

Stomachetosella balani x

Cleidochasma porcellanum xX x x x x x

Cleidochasma protrusum x x x x x

Cleidochasma cribritheca x

Hippoporidra senegambiensis x

Hippoporella spinigera x x x

Hippomenella avicularis x

Flustramorpha flabellaris

Flustramorpha marginata

Flustramorpha angusta

Trypostega venusta x x x x x x x x

Gigantopora polymorpha

Adeonella majuscula

Adeonella cracens

Tessaradoma bispiramina

Tessaradoma circella

Sertella lata

Schizoretepora tessellata

Reteporella dinotorhynchus

Brodiella longispinata x x |

Turbicellepora conica

Celleporaria tridenticulata x x x x x

Celleporaria capensis

Anoteropora latirostris

Mecynoecia clavaeformis x

Mecynoecia delicatula x

Mecynoecia australis x

Plagioecia patina x x x

Liripora lineata x x

Idmidronea contorta x

Idmidronea crassimargo x

Crisia elongata x x

Lichenopora novae-zealandiae x x x

Crisina radians x

x eS % HK OS OK OK OK XK oS OS OK OK OK OK

xX xX KK KK XK XK

x X XK X

x X &

xX XX XK X XK XK X

x X

xX X X X

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 147

distribution within the Indian Ocean and is of interest in being, apparently, the

first Mauritian species to be reported from South Africa.

A notable feature of this essentially shallow shelf fauna is that of the

fifty-one species reported before from this region, twenty-nine have not been

recorded from anywhere else. If at least a similar proportion of the forty-four

new species described prove to be limited to eastern South Africa, then the

endemic component of this fauna will be considerable. Millard (1978) showed

that the endemic component of the South African hydroid fauna achieves its

highest levels in the region between Cape Agulhas and Durban, and that while

the eastern and southern regions of the South African shelf have a hydroid

fauna with a substantial proportion of Indo-West-Pacific species, the character

of the southern fauna, centred on Agulhas Bank seems sufficiently distinctive to

confirm Briggs’s (1974) contention that a separate faunal province may be

recognized in this region. Bryozoan species diversity is frequently related to the

availability of substratum and might be expected to be high on the Agulhas

Bank. The most southerly of the stations represented in the present collections

(SM 184, SM 185) were located at the northern extremity of the Bank; by

analogy with Millard’s (1978) results it seems probable that the South African

bryozoan fauna may prove to be even richer than the present report suggests,

and even more distinctive than the hydroid fauna.

The most exciting aspect of the geographical distribution patterns of the

South African bryozoan fauna is the increasing evidence of strong similarities

with eastern Australian and northern New Zealand faunas, and indications that

this link may be of considerable antiquity. Three examples are given in Table 4:

Bugulella australis and Schizoretepora tessellata have been reported elsewhere

only from South Australia; Escharina waiparaensis was originally described

from Miocene (Brown 1952) and Pliocene (Brown 1954) deposits of New

Zealand, and subsequently live specimens were reported (Powell 1967) from

Three Kings Islands, northern New Zealand. The occurrence of this species in

South African waters is thus of the greatest interest. The new species described

here also include several with equally extraordinary time-space distributions.

For example, Dimorphocella moderna appears to be the only Recent

representative of a genus well known from Australian Tertiary deposits. The

large live colony was collected from the same station that produced the

abundant, also live, material of Aspidostoma livida. Few Recent species of

Aspidostoma are known, and most of these are small encrusting forms with a

habit quite unlike the massive foliose colonies of A. livida. However,

Aspidostoma also seems to be an ancient genus; Brown (1952) described a

number of species with erect colony forms from New Zealand Tertiary

deposits, and other fossil species are known from Australia (Maplestone 1902,

1911). Macropora is another interesting case; the genus is widespread through

the Tertiary deposits of Australia and New Zealand, and one species with a

long fossil history survives today in New Zealand and has also been reported

from the Philippines. Few nominal Recent forms have been described; these

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

require re-examination and it is possible that M. africana may be only the

second Recent form of this ancient genus.

A number of authors have published valuable integrated studies of

Tertiary to Recent bryozoan faunas (for example, Lagaaij & Cook 1973; Cook

& Lagaaij 1976; David & Pouyet 1978) that demonstrate the importance of

such research to bryozoan phylogeny and the contribution it may make to the

broader field of marine zoogeography. Bryozoans are particularly useful

subjects for zoogeographical research; with very short larval lifespans, and

consequent poor dispersal ability, with high species diversity in most shelf seas,

and in many Tertiary deposits, and with perhaps rapid rates of phylogenetic

change through time, they offer great potential for detailed analysis. As Lagaaij

& Cook (1973) reiterate, wide distributions observed in Recent seas are the

legacy of past expansions and contractions. The evident richness of the eastern

South African fauna and its apparently high level of endemism, the numerous

systematic affinities with the fauna of the south-west Pacific (see, for example,

p. 47), and the number of ancient genera and species found in the present

collections demand further investigation. Whether these features of the South

African fauna reflect relict elements, or are truly autochthonous, can only be

decided by further study of Recent populations, in comparison with Tertiary

fossil deposits and Quaternary sediments.

SUMMARY

A total of 130 species of Bryozoa have been identified from samples

collected by the R.V. Meiring Naude between 1977 and 1979. The sampling

stations ranged in depth from 80 m to 900 m, with the majority in depths of less

than 100 m. Bryozoan diversity was highest in the samples from the shallowest

stations; 44 new species are described, including a high proportion of encrusting

cheilostomata. The fauna described is considered to be representative of a

typical shallow shelf-sea assemblage, in marked contrast to the deeper slope

fauna reported upon in the first publication on the Meiring Naude Bryozoa

(Hayward & Cook 1979). Together with the earlier contributions on the South

African Bryozoa, referred to in the text, these two reports now provide a sound

introduction to the bryozoan fauna of this region. Although some taxonomic

problems remain and the question of broader systematic relationships is further

complicated in some cases by these results, the particular character of the

eastern South African fauna is more clearly defined and shown to include a

substantial and distinctive endemic element. Further surveys on the Agulhas

Bank may be expected to yield results that will further enhance this

distinctiveness. Bryozoan skeletal remains were important components of

biogenic carbonate sands in some of the areas sampled, and the presence of

abraded fragments of unrecognized species in these sands shows that the

taxonomic diversity of this fauna is still to be fully recorded.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 149

The discovery of species and genera with extensive time-space distribu-

tions, and in particular the apparent faunal similarity between this region and

Tertiary fossil deposits of Australia and New Zealand, suggest interesting and

fruitful possibilities for further research.

ACKNOWLEDGEMENTS

We are grateful to Dr N. A. H. Millard and Dr P. A. Hulley (South

African Museum) for the opportunity to work on the Meiring Naude

collections, and to Mr P. J. Chimonides (British Museum, Natural History) for

the Scanning Electron Microscopy. We should also like to thank Drs K. Brood

(Naturhistoriska Riksmuseet, Stockholm) and D. P. Gordon (New Zealand

Oceanographic Institute, Wellington) for their helpful comments and criticisms.

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ABBREVIATIONS

Lz length of zooid Iz width of zooid

Lop length of opesia lop width of opesia

Lor length of orifice lor width of orifice

Lov length of ovicell lov width of ovicell

Lg length of gonozooid lg width of gonozooid

Loo.st. length of ooeciostome eho} taza width of brooding zooid

Ie ibieZ: length of brooding zooid ] br.or. width of brooding orifice

Lbr.or. length of brooding orifice Im. width of mandible

Lav length of avicularium

Lad.av. length of adventitious avicularium

Lint.av. length of interzooidal avicularium

Lm length of mandible

BMNH British Museum (Natural History)

SAM — South African Museum

ANNALS OF THE SOUTH AFRICAN MUSEUM

APPENDIX 1

MEIRING NAUDE STATIONS THAT PRODUCED BRYOZOA

Station Co-ordinates

Ss Ss)

SM 103 Dirge 3244

SM 123 30°33 ,4’ 30°48,6’

SM 129 S05845 S0eoileve

SM 131 30°43 2’ 30°40,8’

SM 151 30°14’ S27 207

SM 162 3255 28 ile

SM 163 33°04,6’ 28°06,6’

SM 164 33°04,6’ 28°06,6’

SM 179 SES.) 3 Di A

SM 180 33°29 ,4’ DE De

SM 184 33°39 4’ rahe 4

SM 185 33°39 ,3’ Dg AEG:

SM 232 32°14,9’ 29°10,4’

SM 233 SRS JH 29°09 ,8’

SM 234 SAIS 29°09,1’

SM 239 SP aIARSe 29°00,8’

SM 250 BP S)3)" 29 DIS

APPENDIX 2

Depth, m Date

680 24.5.76

690 105972

850 bMS) 57/7/

780 OMS) 7/7/

900 17-3

630 2d

90 26.5.78

80 29°5218

80 295518

86 BIBS Vis:

90 3578

560-620 25.6.79

540-580 25.6.79

500-520 29:69

90 25.6.79

150-200 2 O9

INDEX TO GENERA AND SPECIES

Correct names are given in italic; synonyms in roman.

abdita 93,97, 99, 100

alia 100

confusanea 92

conspicua 93

coralliformis 88, 92

cracens 88, 90,95

cultrata 88

decipiens 90

distincta 95

expansa 88, 97

falcicula 88

gibba 88, 97, 99, 100

infirmata 95

jellyae 88

ligulata 88

majuscula 88, 90

meandrina 88

pectinata 88, 99

ponticula 86, 88

Pygmaea 88

regularis 87

triton 47

abdita, Adeonella 93,97, 99, 100

Acamarchis dentata 42

acanthina, Chaperia 19

Actinopora 132

Adeona 47,51

Adeonella 51, 87, 140, 143

Adeonellopsis 47

meandrina 88

africana, Cribrilaria 45

africana, Macropora 33, 148

africana, Pachycleithonia 53, 55

alia, Adeonella 100

ambigua, Emballotheca 73

Amphiblestrum 11,16

capense 22

imbricatum 16

inermis 12, 145

pontifex 13

triangulare 12

angusta, Flustramorpha_ 84, 85, 141

Anoteropora 127, 140, 142

latirostris 127

antarctica, Idmidronea 136

Arachnopusia 27, 29

corniculata 29

arcuata, Watersipora 55

armata, Brodiella 103, 123

Arthropoma_ 69

cecilii 69

circinatum 69

Aspidostoma 36, 147

giganteum 38

livida 36,51, 147

asymmetrica, Mucropetraliella 67

(atlantica, Idmidronea overleaf)

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES eyf/

atlantica, Idmidronea 136

australis, Bugulella 40, 147

australis, Chaperia 19

australis, Mecynoecia 131

avicularis, Hippomenella_ 81, 145

balani, Stomachetosella 74

Batopora 140, 142

Beania 41

erecta 42, 141

magellanica 42

paucispinosa 141

rediviva 42, 141

vanhoffeni 141

beatulum, Rhynchozoon 113

Berenicea lineata 132

Bicellariella

capensis 141

chuakensis 141

bicornis, Osthimosia 125

bifaciata, Cellaria 36

biporata, Idmidronea 134, 136

bispiramina, Tessaradoma 102

boryi, Caberea 141

Brodiella 103, 104, 120, 123

armata 103, 123

ignota 103, 122

longispinata 103, 120, 145

Buffonellaria armata 123

Bugula 42

dentata 42

Bugulella 40, 140

australis 40, 147

problematica 40

burrowsi, Tropidozoum 55

buski, Disporella 138

buskii, Steginoporella 32, 141

Caberea 38

boryi 141

darwinii 38, 141, 145

calcarata, Smittoidea 60

Callopora 11

Calyptotheca 72,74

nivea 72

porelliformis 73

capensis, Bicellariella 141

capensis, Celleporaria 126

capensis, Chaperia 19, 22, 23

capensis, Porella 64

Carbasea 6

mediocris 6

cecilii, Arthropoma 69

Cellaria 34

bifaciata 36

fistulosa 141

gracilis 34

Johnsoni 35

paradoxa 35

Ppunctata 34

tectiformis 34

cellariiforme, Tropidozoum 55

Cellepora conica 123

senegambiensis 78

tridenticulata 125, 145

Celleporaria 125

capensis 126

tridenticulata 125

Cellularia crispa 39

infantae 39

ornata 39

quadrata 38

centralis, Macropora 33

cervicornis, Notocoryne 26

Chaperia 19, 25

acanthina 19

acanthina var. australis 22

acanthina var. polygonia 23

australis 19

capensism 19522773

familiaris 23, 144

multifida 20

stephensoni 23

chuakensis, Bicellariella 141

ciliata, Microporella 82

circella, Tessaradoma 102

circinatum, Arthropoma 69

circumspecta, Smittoidea 59

clavaeformis, Mecynoecia 130

Cleidochasma_ 75, 104

contractum 104

cribritheca 76

perspicua 76, 104

porcellanum 75, 78, 145

protrusum 75

Columnella 140

confusanea, Adeonella 92

conica, Turbicellepora 123

conspicua, Adeonella 93

contorta, Escharoides 46

contorta, Idmidronea 133

contractum, Cleidochasma 104

Copidozoum 13

transversum 14

coralliformis, Adeonella 88, 92

corniculata, Arachnopusia 29

cracens, Adeonella 88, 90, 95

crassicollis, Stomachetosella 74

Crassimarginatella 14

marginalis 14, 145

crassimargo, Idmidronea_ 134

cribraria, Porina 87

Cribrilaria 43

africana 45

innominata 43, 145

venusta 43, 145

Cribrilina philomela 46

cribritheca, Cleidochasma 76

Crisia 137

elongata 137

holdsworthii 128, 137

transversata 137

(Crisina overleaf)

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crisina 138

radians 138, 145

watersi 138

crispa, Menipea 39

cultrata, Adeonella 88

Cupuladria 6

multispinata 6, 145

owen 8

Cupularia multispinata 6

cyclops var. tessellata, Odontionella 29

Dactylostega 26, 144

nigrans 29

prima 27

spissimuralis 29

tubigera 29

darwinii, Caberea 38, 141, 145

decipiens, Adeonella 90

delicatula, Mecynoecia 131

dentata, Bugula 42

Desmeplagioecia lineata 132

Diachoris magellanica 42

Diaperoecia 130

Diastopora

lineata 132

reticulata 133

Dimorphocella 13, 47, 62

moderna 48, 147

portmarina 47

pyriformis 47, 48

triton 47,51

dinotorhynchus, Reteporella 108, 141

Discoporella 8

novae zealandiae 137

umbellata 8

discors, Escharella 66

Disporella

buski 138

ciliata 138

distincta, Adeonella 95

documentum, Rhynchozoon 110

elongata, Crisia 137

Emballotheca 73

ambigua 73

nivea 72

errata, Smittoidea 59

erugata, Hornera 139

Eschara contorta 46

flabellaris 84

Escharella 66

discors 66

Escharina 71

pesanseris 71, 145

waiparaensis 71, 145, 147

Escharoides 46

contorta 46

spinigera 76

Eupaxia 38

incarnata 38

quadrata 38

Exechonella 52

expansa, Adeonella 88, 97

falcicula, Adeonella 88

familiaris, Chaperia 23, 144

fasciculata, Liripora 132

Fenestrulina 85

indigena 85

ferruginea, Smittina 58

Figularia 45

philomela 46

fistulosa, Cellaria 141

flabellaris, Flustramorpha 84

flosculum, Iodictyum 109

Flustra acanthina 19

cecilii 69

marginata 84

spinosa 20

Flustramorpha_ 84, 143

angusta 84, 85, 141

flabellaris 84

marginata 84

foraminosa, Gigantopora 86

Foveolaria 13, 16, 29

imbricata 16

Galeopsis mutabilis 53

Gemellipora cribritheca 76

protrusa 75

Gephyrophora polymorpha 86

gibba, Adeonella 88, 97, 99, 100

giganteum, Aspidostoma 38

Gigantopora 86, 143

foraminosa 86

mutabilis 53

polymorpha _ 86, 87, 88

gracilis, Pseudidmonea_ 139

grandis, Macropora 33

Heliodoma 10, 140, 142, 145

implicata 11, 142

Hincksina nigrans 29

Hippaliosina 123

Hippomenella 81

avicularis 81, 145

spatulata 81

Hippoporella 79, 104

hippopus 104

labiata 80, 104

multidentata 80

spinigera 79, 145

Hippoporidra 78

picardi 79

senegambiensis 78, 145

hippopus, Hippoporella 104

Hippothoa pes anseris 71

holdsworthii, Crisia 128, 137

(Holoporella capensis overleaf)

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 159.

Holoporella capensis 126

Hornera 128, 139, 143

erugata 139

smitti 139

Idmidronea_ 133, 143

antarctica 136

atlantica 136

biporata 134, 136

contorta 133

crassimargo 134

parvula 134

Idmonea atlantica 136

contorta 133

crassimargo 134

parvula 134

ignota, Brodiella 103, 120

imbricata, Foveolaria 16

implicata, Heliodoma 11, 142, 145

incallidum, Rhynchozoon 103, 114

indigena, Fenestrulina 85

inermis, Amphiblestrum 12, 145

infirmata, Adeonella 95

innominata, Cribrilaria 43, 145

Inversiscaphos 140

Iodictyum 109

flosculum 109

jellyae, Laminopora 88

Jullienella 145

foetida 145

pearceyi 145

labiata, Hippoporella 80, 104

Lacrimula 140, 142

Laminopora 51

jellyae 88

lata, Sertella 105

latirostris, Anoteropora 127

Lepralia armata 103, 123

circinata 69

foliacea 141

innominata 43

porcellana 75

triangularis 12

tuberculata var. avicularis 81

venusta 86

Lichenopora 137

novae-zealandiae 137

ligulata, Adeonella 88

lineata, Liripora 132

Liripora 132

lineata 132

fasciculata 132

livida, Aspidostoma 36, 51, 147

Lobosoecia semiclausa 133

longispinata, Brodiella 103, 120, 123, 145

Lunulites umbellata 8

Macropora 33, 147, 148

africana 33

centralis 33

grandis 33

magellanica, Beania 42

magnilabris, Steginoporella 33, 141

majuscula, Adeonella 88, 90

Margaretta 143

marginalis, Crassimarginatella 14, 145

marginata, Flustramorpha 84

marionensis, Menipea 40

meandrina, Adeonellopsis 88

Mecynoecia 130, 143

australis 131

clavaeformis 130

delicatula 131

mediocris, Carbasea 6

Membranipora galeata var. inermis 22

galeata var. multifida 20

imbricata 16

inermis 12

marginalis 14

spinosa 19

tubigera 29

Membraniporidra spissimuralis 29

Menipea 39, 140

flabellum 39

crispa 39

marionensis 40

ornata 39

triseriata 39

Micropora 31

normani 32

similis 31, 69

Microporella 82

ciliata 82

flabellaris 84

moderna, Dimorphocella 48, 147

mooraboolensis, Mucronella 80

Mucronella contorta 46

mooraboolensis 80

serratilabris 79, 145

Mucropetraliella 67

asymmetrica 67

watersi 67, 69

multidentata, Hippoporella 80

multifida, Chaperia 20

multispinata, Cupuladria 6, 145

mutabilis, Pachycleithonia 53

nigra, Pachycleithonia 53

nigrans, Dactylostega 29

nivea, Calyptotheca 72

normani, Micropora 32

Notocoryne 26

cervicornis 26

Notoplites 140

novae-zealandiae, Lichenopora 137

novella, Parasmittina 63

(Odontionella cyclops overleaf)

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Odontionella cyclops var. tessellata 29

ornata, Menipea 39

oscitans, Rhynchozoon 103, 116

Osthimosia bicornis 125

owenii, Cupuladria 8

Pachycleithonia 52

africana 53,55

mutabilis 53

nigra 53

paradoxa, Cellaria 35

Parasmittina 62

novella 63

serrula 63, 64

tropica 62, 145

parvula, Idmidronea 134

patina, Plagioecia 131

perforata, Schizoporella 74

perspicua, Cleidochasma 76, 104

pesanseris, Escharina 71, 145

Petralia vultur var. armata 67

peyroti, Discoporella umbellata 10

Dhilomela, Figularia 46

picardi, Hippoporidra 79

Plagioecia 131, 132

patina 131

polymorpha, Gigantopora 86, 87, 88

pontifex, Amphiblestrum 13

porcellanum, Cleidochasma_ 75, 78, 145

Porella 64

capensis 64

porelliformis, Calyptotheca 73

Porina cribraria 87

portmarina, Dimorphocella 47

praetenuis, Vittaticella 127

prima, Dactylostega 27, 144

problematica, Bugulella 40

protrusum, Cleidochasma 75

Pseudidmonea gracilis 139

ptarmicum, Rhynchozoon 103, 118

Puellina venusta 43

pulchra, Schizoporella 104

punctata, Cellaria 34

pustulata, Turbicellepora 124, 125

Pustulopora

australis 13

clavaeformis 130

delicatula 131

proboscidea 131

pygmaea, Adeonella 88

pyriformis, Dimorphocella 47, 48

quadrata, Eupaxia 38

radians, Crisina 138, 145

rediviva, Beania 42,141

regularis, Adeonella 87

Retepora lata 105

radians 138

tessellata 106

Reteporella 103, 108

dinotorhynchus 108, 141

Rhynchopora spinifera 80

Rhynchozoon 102, 103, 104, 110

beatulum 113

documentum 110

incallidum 103, 114

oscitans 103, 116

ptarmicum 103, 118

stomachosum 103, 116

roulei, Setosellina 10, 141, 145

Salicornaria punctata 34

Schizammina 145

pinnata 25, 144

Schizomavella 70

Schizoporella balani 74

cecilit 69

longispinata 103, 120

nivea 72

perforata 74

porelliformis 73

pulchra 104

submersa 48

tenuis 72

Schizoretepora 106

tessellata 103, 106, 147

Schizotheca 102, 103

senegambiensis, Hippoporidra 78, 145

serrula, Parasmittina 63, 64

Sertella 103, 105, 143

lata 105

verecunda 105

Setosellina 10, 140, 145

roulei 10, 141

similis, Micropora 31, 69

sitella, Smittina 56

smitti, Hornera 139

Smittia tropica 62

Smittina 56

ferruginea 58

sitella 56

tropica 62

Smittoidea 58

circumspecta 59

calcarata 60

errata 59

hexagonalis 59

spinifera, Rhynchopora 80

spinigera, Hippoporella 79, 145

spissimuralis, Dactylostega 29

Steginoporella 32

buskii 32, 141

magnilabris 33, 141

stellata, Tennysonia 128

stephensoni, Chaperia 23

Stomachetosella 74

balani 74

crassicollis 74

(stomachosum, Rhynchozoon overleaf)

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 161

stomachosum, Rhynchozoon 103, 116

Strophiella tuberigera 120

submersa, Schizoporella 48

tectiformis, Cellaria 34

tenella, Vittaticella 127

Tennysonia contorta 133

stellata 128

Tervia 128

Tessaradoma 102, 140

bispiramina 102

circella 102

tessellata, Schizoretepora 103, 106, 147

transversata, Crisia 137

transversum, Copidozoum 14

tridenticulata, Celleporaria 125, 145

Triphyllozoon 103

triseriata, Menipea 39

triton, Dimorphocella 47,51

Tropidozoum 55

burrowsi 55

cellariiforme 55

tropica, Parasmittina 62, 145

Trypostega 86

venusta 86, 145

tuberigera, Strophiella 120

tubigera, Dactylostega 29

Tubulipora 128

notomale 134

patina 131

Turbicellepora conica 123

pustulata 125

valligera 124

umbellata, Discoporella 8

valligera, Turbicellepora 124

vanhoffeni, Beania 141

venusta, Cribrilaria 43, 145

venusta, Trypostega 86, 145

venusta, Vittaticella 127

verecunda, Sertella 105

Vittaticella 127

praetenius 127

tenella 127

venusta 127

waiparaensis, Escharina 71, 145, 147

watersi, Mucropetraliella 67, 69

Watersipora arcuata 55

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

(particularly Articles 22 and 51).

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

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

nov., syn. nov., etc.

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

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

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

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

scientific names by which the species previously has been designated are listed in chronological

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

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, 1861: 87.

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

Note punctuation in the above example:

comma separates author’s name and year

“semicolon separates more than one reference by the same author

full stop separates references by different authors

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

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

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

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

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

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

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

7. SPECIAL HOUSE RULES

Capital initial letters

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

e.g. “... the Figure depicting C. namacolus ...’; ‘. .. in C. namacolus (Fig. 10)...’

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

by initials or full names

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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

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

book or article, such as

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

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

to initial capital letter, provided the same generic name is used consecutively.

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

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.

P. J. HAYWARD

P. L. COGK

THE SOUTH AFRICAN MUSEUM’S

MEIRING NAUDE CRUISES

PART 13

BRYOZOA II

a :

S67X T 2 JANUARY 1983 ISSN 0303-2515

‘i AMT NAS \

( MAR 22.963

NH .

S_LIBR ARIES. oe

ANNALS

| F THE OUTH AFRICAN

° : MUSEUM

CAPE ‘TOWN

INSTRUCTIONS TO AUTHORS

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

2. LAYOUT should be as follows:

(a) Centred masthead to consist of

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

Author’s(s’) name(s)

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

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

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

(d) Introduction

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

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous

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

with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of

legends for illustrations should be typed separately, their positions indicated in the text. All

pages should be numbered consecutively.

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

capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered

italics. Further subdivisions should be avoided, as also enumeration (never roman numerals)

of headings and abbreviations.

Footnotes should be avoided unless they are short and essential.

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

up should be left to editor and publisher.

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