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

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

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

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

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 103 °+#Band

December 1993 Desember

Part 5 Deel

9.8.3.9)

a C

LOI

NEW TAXA AND DISTRIBUTIONAL RECORDS

OF AZOOXANTHELLATE SCLERACTINIA

(CNIDARIA, ANTHOZOA) FROM

THE TROPICAL SOUTH-WEST INDIAN OCEAN,

WITH COMMENTS ON THEIR ZOOQGEOGRAPHY

AND ECOLOGY

S. D. CAIRNS

N. B. KELLER

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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Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 147 4

Printed in South Africa by In Suid-Afrika gedruk deur

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

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

D2246

NEW TAXA AND DISTRIBUTIONAL RECORDS OF

AZOOXANTHELLATE SCLERACTINIA (CNIDARIA, ANTHOZOA)

FROM THE TROPICAL SOUTH-WEST INDIAN OCEAN,

WITH COMMENTS ON THEIR ZOOGEOGRAPHY AND ECOLOGY

S. D. Cairns

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560

N. B. KELLER

Institute of Oceanology, Moscow, 117218, Russia

(With 13 figures and 2 tables)

[MS accepted 26 October 1992]

ABSTRACT

Seventy-seven species of azooxanthellate Scleractinia are reported from collections made by R.V.

Anton Bruun, Meiring Naude, and Vityaz from deep-water (to 1 720 m) in the tropical south-west

Indian Ocean. Thirty-six new distributional records are noted for this region, increasing the total

number of azooxanthellates from 64 to 100—one of the most diverse regions in the world oceans for

azooxanthellate Scleractinia. Among the 77 records, seven species (Caryophyllia elongata, Rhizosmilia

robusta, Sphenotrochus (S.) evexicostatus, S$. (S.) imbricaticostatus, Truncatoflabellum gardineri,

T. zuluense and T. multispinosum) and one subspecies (Flabellum (Ulocyathus) japonicum bythios) are

described as new, and four new combinations are proposed.

The distribution and bathymetric ranges of all 100 species are given and zoogeographic affinities

discussed. Twelve per cent of the 100 tropical south-west Indian Ocean species are cosmopolitan or

widespread in distribution and six per cent were uncategorized. Of the remaining 82 species, the

largest distributional pattern is that of Indo-West Pacific (43 %), followed by those species known only

from the Indian Ocean (36,5 %); however, it is believed that, because of the relatively homogeneous

nature of the deep-water fauna, there will be a tendency to find an increase of the Indo-West Pacific

component at the expense of the Indian Ocean ‘endemics’ as the deep-water corals become better

known. Ten species (12%) co-occur in tropical and southern temperate regions and seven species

(8,5%) have a shared distribution with the Atlantic Ocean, five of which are found in the western

Atlantic. Three significant bathymetric zones are defined, based on south-west Indian Ocean azooxan-

thellates: 0-300 m, 300-1 300 m, and 1 300-2 000 m. The potential for using skeletal morphology as

an indicator of environmental conditions (i.e. nutrient level) is discussed.

CONTENTS

PAGE

AN troductOn Peper: Crees es ee te ace 214

ZOOLCOLTAPDY He seek ei Leek ney ie 2A)

Bathymethicaistributionys eye renee eee acai 222

COLOR Vier ee sea k caterer rire epg apse eer Seagtcat nape NMe aha 223

Matenaliandimethodsicycrerrcrr tate tae Nope iene 226

SySLematiclaccoun tae Pere eee eta 228

NCKNOWLEd SEMENISH ae cee seein rere terete learners at 284

FRELELETICES ior eye tayo ne eee et ee ATE ET EN cere a neal, 285

PRY PVCTA I occ SWE ash arses occa store, Meaney alesse austere yotecensrisiake, sete needs 290

213

Ann. S. Afr Mus. 103 (5), 1993: 213-292, 13 figs, 2 tables, app.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

Our knowledge of the azooxanthellate scleractinian fauna of the tropical south-

west Indian Ocean is based on approximately 31 papers (Table 1), most of

which include only one or several incidentally collected species or constitute reports

on expeditions that briefly entered this region (e.g. Valdivia, John Murray Expedi-

tion, Percy Sladen Trust Expedition). The newly reported specimens presented

herein are also based on expeditionary collecting, namely by R.V. Anton Bruun,

Vityaz and the Meiring Naude; however, this paper also includes a re-analysis of

previously reported specimens and a compilation of all previous records for the

region (Table 2). This paper should not be considered as a faunistic revision, but

rather as an annotated checklist of those species for which additional specimens were

collected.

Sixty-four azooxanthellate Scleractinia have been reported previously from

the tropical south-west Indian Ocean (Tables 1, 2). An additional 36 new records

(Table 2) for the region are reported herein, resulting in a total of 100 azooxanthellate

species for this region. Of the 36 new south-west Indian Ocean records, 11 species are

also new records for the Indian Ocean, seven are described as new species, one as a

new subspecies, and four new combinations are proposed. Additional records of 77

of the 100 species (77% of the fauna) are presented in the ‘Systematic Account’.

Examination of an unpublished collection (Zibrowius in prep.) from the same region

TABLE 1

Annotated, chronological list of publications on azooxanthellate Scleractinia from the tropical south-west

Indian Ocean (* denotes significant papers).

Year Author Remarks

18485 Milne Edwards & Haime Two species of Tubastraea from off Seychelles.

1876 Duncan Culicia natalensis from off Natal, South Africa.

1902 Gardiner Two species of Flabellum from South Africa.

*1904 Gardiner Fifteen azooxanthellate species from off South Africa.

1904 Von Marenzeller Six deep-water species from Valdivia stations 243-247 off Kenya

and Tanzania and other records to north in the Indian Ocean.

1926 Van der Horst Eight shallow-water, mostly dendrophylliid species collected on

Percy Sladen Trust Expedition of 1905 off Zanzibar,

Seychelles, Mauritius, and Saya de Malha.

*1927 Van der Horst Eight shallow-water species, primarily dendrophylliids from off

South Africa.

1931 Van der Horst Seven azooxanthellate species from off Seychelles, Mauritius,

and Saya de Malha.

*1938 Gardiner & Waugh Seventeen deep-water flabellids and caryophylliids collected on

the John Murray Expedition (1933-1934) stations 102-133

off Kenya, Tanzania, and Seychelles.

TABLE | (cont.)

Year

*1939

1958

1964

1974a

1974b

1974

1975

1976

1978

1979

1980

1981

*1981

1982

1984

1985

1989a

1989b

Author

Gardiner & Waugh

Macnae & Kalk

Pichon

Zibrowius

Pichon

Zibrowius, Southward &

Day

Keller

Pichon

Rosen

Cairns

Best, Faure & Pichon

Zibrowius

Keller

Boshoff

Zibrowius

Schuhmacher

Zibrowius & Grygier

Zibrowius

Grygier

Cairns

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 215

Remarks

Eleven deep-water dendrophylliids and oculinids collected from

the John Murray Expedition (1933-1934) stations 102-133

off Kenya, Tanzania, and Seychelles.

Uncritical listing of eleven shallow-water azooxanthellates from

Inhaca, Mozambique.

List of four shallow-water azooxanthellates from Tuléar,

Madagascar.

Record of Deltocyathus north-west of Madagascar.

Javania insignis from off north-west Madagascar.

Sphenotrochus sp. from reefs off Madagascar.

Polychaete symbionts from three species of Flabellum collected

off Natal and Mozambique.

Fungiacyathus pseudostephanus from off Seychelles and mid-

Indian Ridge.

Uncritical list of eleven shallow-water azooxanthellates from

Tuléar, Madagascar.

List of four shallow-water azooxanthellates from off Aldabra.

Labyrinthocyathus delicatus from off Natal.

List of six shallow-water azooxanthellates off Seychelles.

Lophelia pertusa from north-west of Madagascar.

Caryophyllia scobinosa from off Tanzania.

Forty-two azooxanthellates reported from off South Africa.

Two azooxanthellates from off Natal and Madagascar, both

hosting crab galls.

Discussion of two species of Tubastraea, including records

from off Comores and Madagascar.

Nine species of azooxanthellates, all hosting ascothoracid para-

sites, off Natal, Mozambique, Kenya, Comores, and

Seychelles.

Additional records of Balanophyllia stimpsonii from off

Mozambique and Natal, South Africa.

Two shallow-water azooxanthellates hosting ascothoracid

parasites, from off Kenya, Reunion, and Comores Islands.

Several records of Flabellum pavoninum off Kenya and Durban,

South Africa.

Eight species of azooxanthellates discussed incidental to review

of Philippine fauna, from off Natal, Mozambique, Tanzania,

Kenya, and Reunion.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

seen at the Station Marine d’Endoume, Marseille (specimens from Marion Dufresne,

Meiring Naude, Galathea, Cruise BENTHEDI on Le Suroit, Plante collection)

revealed approximately 25 additional azooxanthellate species from this region, which

would result in a fauna of about 125 species—one of the most diverse azooxanthellate

scleractinian faunas in the world oceans.

Historical resumé. Milne Edwards & Haime (1848b) would appear to have

reported the first azooxanthellate scleractinians from the tropical south-west Indian

Ocean, Coenopsammia ehrenbergiana (= Tubastraea coccinea) and Coenopsammia

viridis (= Tubastraea micrantha), from the ‘Seychelles’, two common, shallow-water

species. Another shallow-water species, but less commonly collected, is Culicia natal-

ensis, reported by Duncan (1876) from off Natal, South Africa. All of the remaining

records of azooxanthellate corals from this region were made in the twentieth century

and are summarized in Table 1; only the five most significant contributions are dis-

cussed in greater detail. Most of the other references listed in Table 1 are uncritical

checklists or concentrate on other geographic areas or topics that only incidentally

mention Scleractinia from the south-west Indian Ocean.

The first significant paper on south-west Indian Ocean azooxanthellates was that

of Gardiner (1904), who examined over 2 000 specimens collected off South Africa

and reported 15 species from this region, including five new species. Although some

of his identifications subsequently have been changed, this paper remains the founda-

tion for serious work on azooxanthellate corals from this region. As a counterpart to

Gardiner’s (1904) contribution, which dealt only with the caryophyllids and flabellids,

Van der Horst (1927) reported eight species of dendrophylliids from the same region,

based on the same collection sources.

In another pair of papers, Gardiner & Waugh published the results of the Sclerac-

tinia collected off Kenya, Tanzania and the Seychelles from the John Murray

Expedition. Their first paper (Gardiner & Waugh 1938), like Gardiner’s (1904), was

restricted to the caryophylliids and flabellids; their second (Gardiner & Waugh 1939),

like Van der Horst’s (1927), reported the dendrophyllids and other minor families.

Whereas Gardiner and Van der Horst reported primarily shallow-water azooxan-

thellates, the Gardiner & Waugh papers reported deeper-water species, their two

contributions discussing 28 species collected from H.E.M.S. Mabihiss — stations

102—133 within the south-west Indian Ocean (Sewell 1935).

Boshoff’s (1981) annotated checklist of South African Scleractinia included

42 azooxanthellate species from the south-west Indian Ocean. Unfortunately, none of

his specimens were illustrated, his localities are confused and obscure, and many of his

identifications are incorrect (Zibrowius & Gili 1990). We are forced to agree with

Zibrowius & Gili that Boshoff’s contribution is misleading, and we look forward to

Zibrowius’ revision of the deep-water corals from this region, which will include a

re-analysis of all the Boshoff specimens.

Other papers that include useful information on south-west Indian Ocean azoo-

xanthellates, but do not report specimens from this region, include: Wood-Mason &

Alcock (1891a, 18916); Alcock (1893, 1898, 1902c); Bourne (1905); Von Marenzeller

(1907a, 1907b); Van der Horst (1922); Gardiner (1929); Wells (1935); Scheer & Pillai

(1974, 1983); Pillai & Scheer (1976); Zibrowius (1980); Fricke & Schuhmacher (1983);

Zibrowius & Gili (1990); and Sheppard & Sheppard (1991).

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN Dale)

ZOOGEOGRAPHY

Of the 100 species of azooxanthellate Scleractinia known from the tropical south-

west Indian Ocean, 12 are cosmopolitan or widely distributed (Fig. 1; Table 2: distri-

bution pattern 6) and six are not categorized because they are not identified to the

species level or have a disjunct distribution (Table 2: pattern 7), e.g. Fungiacyathus

pseudostephanus. Neither of these categories contributes to an understanding of zoo-

geographic affinities.

Of the remaining 82 species, the largest component is the Indo-West Pacific

pattern (Table 2: pattern 3), shared by 35 species (43 %). Thirty species (36,5 %) are

known only from the Indian Ocean: 20 species (24,5 %, Table 2: pattern 1) thus far

Tropical

South-West

Indian Ocean

(20)

Indian Ocean

(10)

Indo-West Pacific

(35)

Fig. 1. Pie-diagram illustrating the seven zoogeographic patterns of the 100 species of south-west

Indian Ocean azooxanthellate Scleractinia. Numbers in inner circle label the seven zoogeographic

patterns discussed in the text and tabulated in Table 2. Numbers in parentheses are the numbers of

species having that pattern.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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222 ANNALS OF THE SOUTH AFRICAN MUSEUM

known only from the tropical south-west Indian Ocean, and the other 10 species

(12 %, Table 2: pattern 2) are more widely distributed in the Indian Ocean. Although

Briggs (1974) divided the tropical Indo-Pacific region into the western Indian Ocean

Province and the Indo-Polynesian Province, separated at the Persian Gulf, he also

stated that there is no obvious physical barrier between east Africa and the Malay

Peninsula and urged caution in interpreting endemic faunas for either province. We

agree. To assume that the 20 species now known only from the tropical south-west

Indian Ocean are endemic to that region would be misleading. The finding of 11 new

records for the Indian Ocean, nine of which were heretofore known only from the

western Pacific, reinforces the supposition that the south-west Indian Ocean fauna is

an outpost of an essentially homogeneous Indo-West Pacific fauna and, as the deep-

water corals are better understood, the Indo-West Pacific component will probably

increase at the expense of the ‘endemic’ and ‘exclusively’ Indian Ocean species.

Ten species (12 % of the 82 species) have a shared distribution with temperate

regions to the south: eight species being found also off temperate South Africa

(Table 2: pattern 4A) and two species occurring in the temperate/circum-Subantarctic

regions (Table 2: pattern 4B).

The final category (Table 2: pattern 5) is perhaps of greatest interest because of

its unexpected nature: seven (8,5 %) of the tropical south-west Indian Ocean azooxan-

thellates have a shared distribution in the Atlantic Ocean. Six of the species occur in

the eastern Atlantic and five occur in the western Atlantic.

As mentioned in the introduction, at least 125 species of azooxanthellate Sclerac-

tinia are believed to occur in the tropical south-west Indian Ocean, which represents

one of the highest diversities of this type of coral in the world oceans. For comparative

purposes, the number of azooxanthellates in some other well-studied regions, in

descending order of diversity, are: 140 species (estimate), New Caledonia region

(Zibrowius in prep.); 116, tropical western Atlantic (Cairns 1979); 110 (estimate),

Philippine region (Cairns 19896); 102, Japanese region (Yabe & Eguchi 1942; Eguchi

1968; Cairns in prep.); 85, north-east Atlantic (Zibrowius 1980); 54, Hawaiian Islands

(Cairns 1984); 48, South Australia—Victoria—Tasmania region (Cairns & Parker 1992);

37, entire Antarctic and Subantarctic region (Cairns 1982); 25, temperate north-east

Pacific (Cairns in prep.); and 14, temperate north-west Atlantic (Cairns 1981).

BATHYMETRIC DISTRIBUTION

The azooxanthellate corals of the south-west Indian Ocean can be divided into

three bathymetric zones: 0-300 m, 300-1 300 m and 1 300-2 000 m. These zones

were established by tabulating the number of species occurring in each 100 m interval

and then looking for depths at which major gains or losses of species occurred.

Sixty-eight species occur in the subtidal zone (0-300 m), slightly over half of

which (35) occur only in this zone. Species of the Dendrophylliidae (21 species) and

Caryophylliidae (22 species) predominate, as well as representatives of the Flabellidae

(eight species), Turbinoliidae (seven species), Micrabaciidae (three species), Guynii-

dae (two species), Pocilloporidae, Oculinidae, Fungiacyathidae, Rhizangiidae and

Anthemiphylliidae—the last five families each being represented by only one species.

The number of species in the subtidal zone reaches a maximum (49) in the

100-200 m interval, and then decreases abruptly to only 41 species in the 200-300 m

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 223

interval. Only 35 species occur at a depth range of 300-400 m, most of them also

known from shallower depths, five appearing for the first time, but another 35 species

do not cross this border from shallower water. Thus, the subtidal zone appears to be

well isolated and we consider 300 m to be its lower boundary, which is 100 m lower

than the generally accepted model of marine zonation suggested by Belyaev et al.

(1959).

In the upper bathyal zone (300-1 300 m) the number of species steadily decreases

from the lower boundary of the subtidal zone (41 species) to 1 300 m (only 12 spe-

cies). This lower boundary is also 100 m deeper than that based on bivalve species

investigated by Krylova (1989). We may thus conclude that the outer shelf margin and

upper continental slope (0-1 300 m) are the most favourable zones for azooxanthel-

late corals. The upper region of the slope to 700 m is inhabited by the same families

(with the exception of the Pocilloporidae) as occur in the subtidal zone. The families

Guyniidae, Rhizangiidae and Anthemiphylliidae do not occur deeper than 700 m.

Furthermore, six species of caryophylliids do not occur deeper than 700 m, and two

are endemic to the 500-700 m interval. The families Fungiacyathidae, Guyniidae and

Flabellidae each lose a species at depths below 700 m. Thus, it appears that the

300-700 m interval is populated by a transitional coral fauna, the lower boundary of

this transitional region (700 m) lying 300 m shallower than was suggested in the

general model of Belyaev et al. (1959) of 1 000 m.

The fauna of the lower bathyal zone (1 300-2 000 m) is the most isolated, consist-

ing of 12 species, 10 of which have a broad bathymetric range. Three of the 12 lower

bathyal species also range into the subtidal zone, seven occur as shallow as the upper

bathyal zone, and two occur in the lower interval of the upper bathyal. No species

makes a first appearance in this zone. Only three families are represented: Caryophyl-

liidae (six species), Flabellidae (four species) and Fungiacyathidae (two species). Only

six of the 12 species reach the depth of 2 000 m, the lower boundary of this zone. The

position of the lower boundary of the bathyal zone is shallower by 1 000 m than the

model of Belyaev et al. (1959), but this difference may be explainable due to a paucity

of deep-water collections in the south-west Indian Ocean.

All boundaries established for the subtidal and bathyal zones, based on the

azooxanthellate corals, coincide with those found for Brachiopoda and Bivalvia by

Zezina (1976) and Krylova (1989), respectively. This is not surprising, since all three

taxa belong to the same trophic feeding group: the sestonfeeders.

ECOLOGY

It is sometimes possible to predict the environment in which a deep-water coral

lives by an analysis of the morphology of the coral, which is essentially an immobile

organism with finely-tuned adaptations to variations in the environment. For this

purpose, it is better to use the solitary (non-colonial) species because the more

complex construction of colonial species may obscure the interpretation (Keller 1981).

Analysis of this kind may also serve as a model for palaeoecological reconstructions.

In the subtidal zone of the tropical south-west Indian Ocean, in conditions of high

water productivity (Bogorov et al. 1968; Koblenz-Mishke 1977), there are many kinds

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

of azooxanthellate Scleractinia of various sizes and shapes. The largest specimens and

most diverse taxa settle on hard substrates, i.e. muddy sand, pebbles, rubble, or rocky

bottoms covered with loose sediment. Species common to this environment are

Stephanophyllia complicata, S. fungulus, Dasmosmilia variegata, Rhizosmilia robusta

sp. nov., Javania insignis, Flabellum pavoninum, Truncatoflabellum multispinosum

sp. nov., 7. inconstans, and numerous dendrophylliids.

Another homogeneous coral association occurs on loose sandy bottoms. Most of

the species of this association are part of the interstitial fauna, e.g. Caryophyllia

rugosa, C. cornuformis, Sphenotrochus evexicostatus sp. nov., S. gilchristi, S. auran-

tiacus, §. imbricaticostatus sp. nov., Peponocyathus australiensis, Aulocyathus juven-

escens and Guynia annulata. These species belong to three families—Caryophylliidae,

Turbinoliidae and Guyniidae—but have many similar features: small, unattached

coralla with a GCD usually less than 10 mm; low number of septa, usually 48 or

fewer; dense skeletal elements; narrow, conical coralla; and smooth calicular margins.

Species of Truncatoflabellum dwell on the sand in the same locations, but differ in

having larger coralla (GCD often over 10 mm) and a trapezoid corallum shape result-

ing from asexual transverse division. However, their skeletal elements are dense and

their calicular margins are smooth.

There may be many reasons for a small-sized coral association. Thiel (1975)

believed that invertebrate groups that are faced with constant food restrictions consist,

on average, of small individuals. My investigations (Keller 1978, 1989) on the coral

associations of the abyssal region and mid-oceanic, bathyal submarine ridges, which

both have low nutrient levels, confirm this assertion. However, in addition to their

small size, corals in oligotrophic regions also adapt by having a deep fossa, a scalloped

calicular margin—the exsert septa forming tall thecal extensions, and a delicate,

fragile skeleton. These characteristics allow the coral to accomplish the function of

support in the most economical way, which is important in low nutrient level regions.

On the contrary, species forming the sandy-bottom associations have dense, robust

coralla and their fossae are shallow, with smooth calicular margins due to little or non-

exsert septa. This morphology is indicative of normal trophic conditions. Maps of the

primary production and zooplankton distribution (Bogorov et al. 1968; Koblenz-

Mishke 1977) confirm this hypothesis. We suppose that the small size of subtidal

corals is related to their location on a shifting sand substrate, not favourable to their

existence or normal development. The corals occurring on loose, mobile sand often

possess fewer septa than average, possibly as a result of neotenic development, which

is common in unfavourable conditions (Keller 1978).

As noted previously, species of three families (Fungiacyathidae, Micrabaciidae

and Caryophylliidae) are widely distributed in the upper bathyal and transitional

regions. These families have the greatest bathymetric ranges among the Scleractinia,

because each includes one genus that occurs at abyssal depths. Fungiacyathidae is rep-

resented by three species of Fungiacyathus: F. paliferus, F. stephanus and F. sibogae;

Caryophylliidae by two species of Deltocyathus: D. andamanicus and D. rotulus; and

Micrabaciidae by Letepsammia formosissima. As noted before (Keller 1978), there are

definite peculiarities in construction of scleractinian skeletons that make it possible for

them to adapt to unfavourable environmental conditions, in particular, life in low

nutrient levels associated with environments at great depths. This distinctive morpho-

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 225

logy consists of a flattened discoidal growth form, a widely open calice with a deep

fossa, highly exsert septa, and a light fragile skeleton. Corals with such a morphology

maximize the area for food capture and for respiration, and can live in conditions of

high nutrient levels as well as being pre-adapted to survive in low nutrient levels.

Species of Flabellum (Flabellidae) also occur in the upper bathyal and transitional

regions. In the nearshore areas and in other highly productive areas, such as over

seamounts (e.g. Saya de Matha Bank), most species of this genus have a serrate cali-

cular edge, a shallow fossa, a narrow elliptical calice with a smooth calicular margin,

and numerous septa. The widely distributed F. pavoninum is a good example. Lower

in the transitional region, but still in the upper bathyal zone where the productivity is

not very high (Bogorov et al. 1968), two different types of Flabellum construction

exist. Typical of the first type are F. messum and F. lowekeyesi, the latter being very

common in the south-west Indian Ocean at depths of 800-1 000 m. Its adaptive

characteristics are a deep fossa and large calice, a scalloped calicular edge, the septa

of which are quite exsert, and a rather fragile skeleton. The shallower-water

F. messum (430-835 m) has a similar morphology and both species are very similar to

F. marcus Keller, 1981, which occurs on the oligotrophic mid-oceanic Marcus-Necker

submarine ridge. The construction of these three species is well adapted to catching

and digesting food in oligotrophic waters, since it requires less calclum carbonate and

energy compared with a similarly-sized dense skeleton with a smooth calicular margin.

A second, deeper-water group of Flabellum species typifies a second kind of skeletal

construction, including the species F. apertum (863-1 400m) and F. japonicum

bythios subsp. nov. (1 095-1 720 m). These species, like the first type, have a deep

fossa, an open calice, and a fragile skeleton, but are distinguished from the first group

by having a flattened corallum, somewhat similar to that of Stephanocyathus.

Three species of Caryophyllia, two of which dwell in the upper bathyal and tran-

sitional zones, are also characterized by an open calice and highly exsert septa

producing a scalloped calicular margin: C. grandis (183-490 m), C. scobinosa

(535-960 m) and C. ambrosia (430-2 000 m). The morphology of C. ambrosia is most

similar to C. grandis, the former replacing the latter at greater depth. The main differ-

ence between the two species is that the number of septa of the shallower species,

C. grandis, is greater than that of the deeper C. ambrosia, which supports the corre-

lation suggested by Keller (1978) of a decreasing septal number with increase in

depth.

The upper bathyal species C. profunda (80-755 m) has a somewhat different

morphology: a shallow fossa with only a slightly scalloped calicular margin. However,

its shallow depth range is reflected in a relatively large number of septa (up to

96 septa and 24 pali). It is advantageous for this species to exist in unfavourable

trophic conditions, because its numerous septa (and corresponding mesenteries)

provide an increased area for food capture and absorption.

The four species of Stephanocyathus live in relatively deep water, two of them

occurring in the upper bathyal zone—S. explanans (183-614 m) and _ S. spiniger

(210-695 m), and two others in the lower bathyal—S. campaniformis (1 600-1 610 m)

in the south-west Indian Ocean and the better known S. nobilis (609-2 000 m).

All species in this genus are characterized by having a flattened corallum, an open

calice, and highly exsert septa, which produce a scalloped calicular margin. As

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

mentioned above, corals with such a morphology are best adapted for living in waters

having a poor nutrient level.

We conclude, therefore, that a certain kind of environment, i.e. low nutrient

level, leads to a certain type of skeletal morphology that occurs in parallel in different

families, in agreement with the law of homology series of variations (Vavilov 1967).

However, for an accurate reconstruction of environmental conditions, either in the

Recent or in the geologic past, it is necessary to analyse the entire coral assemblage at

a particular depth. If in a community there exist species with exsert septa and scal-

loped margins and also flattened forms, it is not significant. However, if most or all of

the species have highly exsert septa, a deep fossa, and a fragile skeleton, as is typical

of those species in the bathyal of the south-west Indian Ocean, we may conclude that

they lived in a region of low nutrient levels. This conclusion is in agreement with maps

of primary production and zooplankton distribution published by Koblenz-Mishke

(1977) and Bogorov et al. (1968).

MATERIAL AND METHODS

Material. The specimens that form the basis of this study derive primarily from

three sources (see Station List). Many were obtained from 38 stations of cruises 7 and

8 of the Indian Ocean International Expedition, the specimens collected from waters

of a wide range of depths (34-1 360 m) off Mozambique and Kenya using R.V. Anton

Bruun in 1964 (see Anonymous 1965). Secondly, deeper-water (to 1 720 m) specimens

were examined from the Madagascar and Mascarene plateaus, off south-eastern

Mozambique and off Madagascar from 33 stations of the Russian research vessel

Vityaz (cruise 17) in 1988-1989. Third, a collection of relatively shallow-water azoo-

xanthellates (primarily 50-100 m) was studied, comprising specimens from 52 stations

made by the R.V. Meiring Naude off Zululand, South Africa. The geographic and

bathymetric data for all of these stations are listed in the Station List. The R.V. Anton

Bruun, Meiring Naude, and some of the Vityaz specimens are deposited at the

National Museum of Natural History, Washington, D.C. (NMNH), but most of the

Vityaz specimens are deposited at the Institute of Oceanology, Moscow (IOM).

In addition to the newly reported material, previously reported specimens of his-

torical interest were examined by the first author from the following institutions: BM

(Gardiner 1904; Van der Horst 1926, 1931; Gardiner & Waugh 1938, 1939); ORI

(part of Boshoff 1981); ZMA (Alcock 1902a, 19026); and ZMB (Von Marenzeller

1904). Finally, a reference collection of Indian Ocean deep-water corals was examined

in 1991 by the first author at the Station Marine d’Endoume, Marseille, but is not

cited in this publication.

Methods. The geographic region considered in this report is the tropical south-

west Indian Ocean (Fig. 2). The south-western border between the tropical and tem-

perate regions has been variously defined from Algoa Bay to just south of Durban,

but we have chosen to follow Briggs (1974) in setting this boundary at the mouth of

the Kei River (28°22'E 32°41’S). The southern boundary of the region is somewhat

arbitrarily taken to be 40°S; the eastern boundary, 70°E; the northern boundary, 2°S

(border between Kenya and Somalia); and the western boundary, the coast of eastern

Africa and a N-S line from the mouth of the Kei River to 40°S. The region includes

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 227

0°

SEYCHELLES

TANZANIA ZANZIBAR IS.

= Q

\ S 9 ®) de Malha Bank 10

( bey bes. Seo es

3 MASCARENE

P of O

ys - MOZAMBIQUE

/ PLATEAU

MADAGASCAR

MAURITIUS Sune 20°

/ RODRIGUES

REUNION

South West

30°

Indian Ocean

MOZAMBIQUE MADAGASCAR

PLATEA

g Walters Shoal

PLATEAU

40°

30° 40° 50° 60° TOZE

Fig. 2. Map of the region covered in this checklist.

the following countries: north-eastern South Africa, Mozambique, Tanzania (includ-

ing Pemba and Zanzibar), Kenya and Madagascar; the following islands: Comores,

Seychelles, Mauritius, Reunion and Rodriques; and the following submarine features:

Madagascar and Mascarene plateaus, Walters Shoal and Saya de Malha Bank.

Species synonymies are complete for records within the Indian Ocean, although

additional citations are often given that provide more complete synonymies, descrip-

tions, and/or good illustrations. In the ‘New Records’ sections, the station number is

followed by the number of specimens examined, and finally the catalogue number (if

any) or museum of deposition. Station data are found in a separate Station List. Types

are deposited primarily at the NMNH, but some are also at the IOM and SAM (see

text). The new species were distinguished and described by Cairns and thus should be

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

cited as: Cairns in Cairns & Keller. The descriptive, historical and zoogeographic sec-

tions are also the work of the first author; the bathymetric and ecology sections are

the work of the second author.

Only those species new to the south-west Indian Ocean and taxa not definitively

identified to the species level are illustrated. The scanning electron microscopy was

done by the first author on a Cambridge Stereoscan 100.

The following abbreviations are used in the text:

Museums

BM British Museum (Natural History), London

IOM Institute of Oceanology, Moscow

NM Natal Museum, Pietermaritzburg

NMNH National Museum of Natural History, Washington, D.C.

ORI Oceanographic Research Institute, Durban

SAM South African Museum, Cape Town

USNM United States National Museum (now the NMNH, Smithsonian, Wash-

ington, D.C.)

ZMA Zoological Museum, Amsterdam

ZMB Zoologisches Museum, Berlin

Vessels

AB. R.V. Anton Bruun

JM John Murray Expedition (H.E.M.S. Mabihiss)

MN R.V. Meiring Naude

Vv R.V. Vityaz

Morphological terms

GCD Greater Calicular Diameter

GCD : LCD Ratio of Greater Calicular Diameter to Lesser Calicular Diameter

SEM Scanning Electron Microscopy

Sx, Cx, Px Septa, Costae, or Pali (respectively) of cycle designated by numerical

subscript

SYSTEMATIC ACCOUNT

A checklist of species recorded from the tropical south-west Indian Ocean and

their distribution is provided in Table 2.

Suborder ASTROCOENIINA

Family Pocilloporidae

Genus Madracis

Madracis sp. A

Fig. 3A-B

New records

V-2697, 1 branch, IOM; AB-372B, 1 colony, USNM 91499; AB-400C,

1 branch, USNM 91498.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 229

Remarks

A well-preserved branch tip from Walters Shoal (Fig. 3A—B) measures 14,5 mm

long and 2,2—2,5 mm in diameter, supporting about 20 corallites. Costae are highly

echinulate and faintly striate. Corallites are circular to slightly elongate, each with

10 primary septa and traces of 10 secondary septa. Very small paliform lobes occur on

inner edges of primary septa. Styliform columella massive.

Like most other genera containing shallow-water azooxanthellate species (e.g.

Culicia, Polycyathus, Oculina, Balanophyllia, Dendrophyllia and Tubastraea), many

species have been described (about 15 in the case of Madracis) but none of these

genera has been comprehensively revised. Until a worldwide revision is available for

this genus, we opt to leave these specimens unidentified to species.

At least three species of Madracis have been reported from the western Indian

Ocean: an unidentified species reported by Gardiner & Waugh (1939) from off

Pemba, Tanzania (73-165 m); Madracis sp. cf. M. decactis from the reefs of Tuléar,

Madagascar (Pichon 1978); and M. interjecta von Marenzeller, 1907b, from the Gulf

of Aqaba, Red Sea (see also Scheer & Pillai 1983; Fricke & Schuhmacher 1983) at

122-350 m.

Distribution

Off Mozambique; off Walters Shoal; 42-160 m.

Suborder FUNGIINA

Family Fungiacyathidae

Genus Fungiacyathus

Fungiacyathus (Bathyactis) sibogae (Alcock, 1902a)

Bathyactis sibogae Alcock, 1902a: 108; 1902c: 38 [part.—lectotype from Siboga—208].

Bathyactis symmetrica: von Marenzeller, 1904: 312-313, pl. 18 (fig. 25). Gardiner & Waugh, 1939:

231"

Bathyactis stabilis Gardiner & Waugh, 1939: 231-232, text-figs 1-2 [syn. nov.].

Fungiacyathus (Bathyactis) sibogae: Cairns, 1989b: 10-11, pl. 3 (figs d-k), pl. 4 (figs a—c) [synonymy].

New records

AB-365D, 3, USNM 91487; AB-369J, 3, USNM 91486; AB-370D, 1,

USNM 91489; AB-373H, 2, USNM 91488; AB-373J, 3, USNM 91485.

Remarks

The south-west Indian Ocean specimens collected by R.V. Anton Bruun are

indistinguishable from the lectotype of the species ard to those reported by Cairns

(1989b) from Indonesia. Gardiner & Waugh’s (1939) specimens of Bathyactis stabilis

and B. symmetrica (BM 1939.7.13.92 and 1939.7.13.93, respectively) and Von Maren-

zeller’s (1904) B. symmetrica from Valdivia—245 (ZMB 5066) were examined by SDC

and also found to be conspecific, the latter being 20,8 mm in calicular diame er.

Distribution

Indian Ocean: off Zululand, South Africa; off south-eastern Mozambique; off

Pemba, Tanzania (Von Marenzeller 1904); off Kenya (Gardiner & Waugh 1939); off

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

south-west Madagascar; Gulf of Oman (Gardiner & Waugh 1939); 463-1 948 m. Else-

where: Indonesia; 522—1 914 m (Cairns 1989b). Fungiacyathus sibogae is a new record

for the Indian Ocean in name only, being previously reported by Gardiner & Waugh

(1939) and Von Marenzeller (1904) as B. stabilis and B. symmetrica.

Fungiacyathus (Fungiacyathus) stephanus (Alcock, 1893)

Bathyactis stephanus Alcock, 1893: 149, pl. 5 (fig. 12, 12a).

Bathyactis symmetrica: Gardiner & Waugh, 1939: 230-231 [part.—John Murray-185].

Bathyactis stephana: Gardiner & Waugh, 1939: 232.

Fungiacyathus (F.) stephanus: Cairns, 1989b: 7-9, pl. 1 (figs a-k), pl. 2 (figs a-n) [synonymy and

distribution].

New records

AB-370D, 1, USNM 91500; AB-373H, 4, USNM 91501, 1, SAM-—H4574;

AB-374D, 1, USNM 81534; AB-399C, 13, USNM 80859.

Distribution

Indian Ocean: off Natal (Gardiner & Waugh 1939) and Zululand, South Africa;

off Mozambique; Gulf of Aden (Gardiner & Waugh 1939); Bay of Bengal (Alcock

1893); 880-2 000 m. Elsewhere: Indonesia; Philippines; 245-1 977 m (see Cairns

19895).

Fungiacyathus (Fungiacyathus) paliferus (Alcock, 1902a)

Bathyactis palifera Alcock, 1902a: 108; 1902c: 38, pl. 5 (fig. 34, 34a).

Fungiacyathus sp. Zibrowius & Grygier, 1985: 119-120 (figs 6-7).

?Fungiacyathus sp. Zibrowius & Grygier, 1985: 120 (figs 8-9).

Fungiacyathus (F.) paliferus: Cairns, 1989b: 9-10, pl. 2 (figs c-i), pl. 3 (figs a-c) [synonymy and

distribution].

New record

V-2722, 1, IOM.

Distribution

Indian Ocean: ?off Natal, South Africa (Zibrowius & Grygier 1985); Reunion

(Zibrowius & Grygier 1985); Walters Shoal; 99-720 m. Elsewhere: Philippines; Indo-

nesia; Great Australian Bight; 75-522 m (Cairns 1989b).

Family Micrabaciidae

Genus Letepsammia

Letepsammia formosissima (Moseley, 1876)

Fig. 3D

Stephanophyllia formosissima Moseley, 1876: 561-562; 1881: 201-204, pl. 4 (fig. 11), pl. 13 (figs 6-7),

pl. 16 (figs 8-9). Van der Horst, 1927: 7. Gardiner & Waugh, 1939: 234. Boshoff, 1981: 24.

Zibrowius & Grygier, 1985: 120.

Letepsammia formosissima: Owens, 1986: 487. Cairns, 1989b: 15-18, pl. 6 (fig. j), pl. 7 (figs g-i),

pl. 8 (figs a—d) [synonymy and distribution].

Stephanophyllia: Williams, 1986, upper left colour photo.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 231

New records

V-2608, 1, IOM, 1, USNM 91506; V-2626, 33, IOM, 6, USNM 91507;

AB-358A, 2, USNM 91508; AB-370G, 10, USNM 75642; AB-370H, 1, USNM

75641; AB-372G, 2, USNM 75643; AB-390R, 1, USNM 91509; AB-390S, 30,

USNM 75640; Vema-14 SAT6, 6, USNM 75644; MN-ZL3, 1, USNM 91505.

Remarks

An undescribed species of Letepsammia from the south-west Indian Ocean was

mentioned first by Squires (1967: 505), and later by Owens (1986: 487) and Cairns

(1989b: 16). This ‘species’ differs from typical L. formosissima in having a denser and

more robust corallum, coarser septal teeth, and a papillose columella. Because these

differences are slight and may be interpreted as intraspecific variation, a new species is

not described at this time.

Distribution

Indian Ocean: off Nieca River mouth, South Africa (Zibrowius & Grygier 1985);

off Durban, South Africa (Van der Horst 1927; Boshoff 1981); off Zululand; off

Mozambique; off Pemba, Tanzania (Gardiner & Waugh 1939); off north-west Mada-

gascar; 320-780 m. Elsewhere: Japan; Philippines; Hawaiian Islands; New Zealand;

Australia; Tasmania; 97-828 m (Cairns 1989b).

Genus Stephanophyllia

Stephanophyllia fungulus Alcock, 19026

Stephanophyllia fungulus Alcock, 1902b: 122-123; 1902c: 40, pl. 5 (fig. 35a—b). Gardiner & Waugh,

1939: 234. Pillai & Scheer, 1976: 14. Cairns, 1989b: 21-23, pl. 10 (figs a-k), pl. 11 (figs a—b) [syn-

onymy and distribution].

Non Stephanophyllia fungulus Alcock. Boshoff, 1981: 24.

New records

AB-373B, 1, USNM 91510; MN-—ZCC1, 1, USNM 91511.

Remarks

The specimen from MN-ZCC1 is very similar to the specimen illustrated by

Cairns (19895, pl. 10h) from off Natal.

Distribution

Indian Ocean: off Natal, South Africa (Cairns 19896); off south-eastern Mozam-

bique; Chagos Archipelago (Gardiner & Waugh 1939); Maldive Islands (Pillai &

Scheer 1976); 98-236 m. Elsewhere: western Pacific Ocean; 15-635 m (Cairns 1989b).

Stephanophyllia complicata Moseley, 1876

Stephanophyllia complicata Moseley, 1876: 558; 1881: 198-201, pl. 4 (fig. 12), pl. 13 (figs 3-5). Van

der Horst, 1926: 51; 1931: 11. Gardiner & Waugh, 1939: 234. Pillai & Scheer, 1976: 14. Cairns,

1989b: 21, pl. 12 (figs a—b).

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

New record

V—2804, 1, IOM.

Remarks

Three of the specimens reported by Van der Horst (1926, 1931) from Saya de

Malha Bank are deposited at the USNM (81875).

Distribution

Indian Ocean: Saya de Malha Bank (Van der Horst 1926, 1931); Chagos Archi-

pelago (Gardiner & Waugh 1939); Maldive Islands (Pillai & Scheer 1976); 229-236 m.

Elsewhere: Indonesia; 236 m (Moseley 1881).

Suborder FAVIINA

Family Rhizangiidae

Genus Culicia

Culicia sp. cf. C. natalensis (Duncan, 1876)

Fig. 3G

Cylicia tenella var. natalensis Duncan, 1876: 439-440, pl. 40 (fig. 3).

?Culicia tenella: Gardiner & Waugh, 1939: 230 (John Murray—123). Boshoff, 1981: 24.

New record

AB-421A, 2 colonies, USNM 91515.

Diagnosis

Larger colony (Fig. 3G) roughly spherical, about 4 cm in diameter, consisting of

50 closely spaced corallites. Corallites elongate and tubular, with circular calices up to

6,5 mm in diameter. Epitheca smooth and thin. Septal symmetry irregular: 10-12

primary septa, an equal number of secondary septa, and occasionally rudimentary

tertiary septa in some sectors. Primary septa large, each having 4 or 5 prominent

teeth, and joined to columella. Columella rudimentary and papillose.

Remarks

Specimens of Culicia have been reported at least four times from the south-west

Indian Ocean: C. excavata (Milne Edwards & Haime, 1849) from the Cape of Good

Hope; C. tenella var. natalensis (Duncan, 1876) from off Natal; C. tenella by Gardiner

& Waugh (1939) from off Pemba, Tanzania; and C. tenella by Boshoff (1981) from off

Mozambique and the Natal regions. In calicular size, septal number and arrangement,

and distribution (i.e. tropical south-west Indian Ocean), the R.V. Anton Bruun speci-

mens resemble C. natalensis (Duncan, 1876) more so than C. excavata, the latter

having smaller corallites, fewer septa, and appearing to be restricted to temperate

South Africa. The type of C. natalensis could not be found for comparison. Culicia

natalensis is very similar to C. hoffmeisteri Squires, 1966, a species known only from

South Australia at 0-29 m (Cairns & Parker 1992). When more specimens of the

south-west Indian Ocean species are collected, detailed comparisons should be made

to this Australian species.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 233

Distribution

Off Natal, South Africa (Duncan 1876; Boshoff 1981); ?off Pemba, Tanzania

(Gardiner & Waugh 1939); off Kenya; 34 m.

Family Oculinidae

Genus Madrepora

Madrepora oculata Linnaeus, 1758

Madrepora oculata Linnaeus, 1758: 798. Zibrowius, 1974a: 762-766, pl. 2 (figs 3-5); 1980: 36-40,

pl. 13 (figs A-P) [synonymy]. Cairns, 1979: 39-42, pl. 3 (fig. 2), pl. 4 (fig. 5), pl. 5 (figs 1-3)

[synonymy].

Lophohelia investigatoris Alcock, 1898: 24—25.

Amphihelia (Diplohelia) moresbyi Alcock, 1898: 25-26 [fide Zibrowius, 1974a].

Amphihelia oculata: von Marenzeller, 1904: 308-310, pl. 14 (fig. 1).

Madrepora kauaiensis: Gardiner & Waugh, 1939: 227.

Non Madrepora oculata Linnaeus. Boshoff, 1981: 27 [= a dendrophylliid, H. Zibrowius, pers.

comm. ].

Lophelia exigua: Boshoff, 1981: 37.

New records

V-2699, 7 branches, IOM; AB-—365D, 2 branches, USNM 77210.

Distribution

Indian Ocean: Off Pemba, Tanzania (Gardiner & Waugh 1939); off south-west

Madagascar; Madagascar Plateau; off Somalia (Von Marenzeller 1904); Red Sea (Von

Marenzeller 1904); Arabian Sea (Alcock 1898); Laccadive Islands (Alcock 1898);

St Paul and Amsterdam Islands (Zibrowius 1974a); 732-1 270 m. Elsewhere: cosmo-

politan; 80—1 500 m (Cairns 1979).

Family Anthemiphylliidae

Genus Anthemiphyllia

Anthemiphyllia dentata (Alcock, 1902a)

Fig. 3E

? Discotrochus investigatoris Alcock, 1893: 142, pl. 5 (fig. 5, Sa).

Discotrochus dentatus Alcock, 1902a: 104; 1902c: 27, pl. 4 (fig. 26). Gardiner & Waugh, 1938: 194.

Pillai & Scheer, 1976: 16.

Anthemiphyllia dentata: Cairns & Parker, 1992: 16-17, pl. 4 (figs e-f) [synonymy].

New record

V—2804, 1, IOM.

Distribution

Indian Ocean: Saya de Malha Bank; Arabian Sea (Alcock 1893; Gardiner &

Waugh 1938); Maldive Islands (Gardiner & Waugh 1938; Pillai & Scheer 1976);

193-494 m. Elsewhere: off Japan; Indonesia; South Australia; 75-522 m (see Cairns

& Parker 1992).

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Suborder CARYOPHYLLIINA

Family Caryophylliidae

Genus Caryophyllia

Caryophyllia ambrosia ambrosia Alcock, 1898

Fig. 3H

Caryophyllia communis: Wood-Mason & Alcock, 1891a: 6. Alcock, 1898: 11-12.

Caryophyllia ambrosia Alcock, 1898: 12, pl. 1 (fig. 1, 1a). Zibrowius, 1980: 63-65, pl. 25 (figs A-K)

[synonymy].

Caryophyllia clavus: van der Horst, 1931: 3 [part.—specimens from Laccadive Sea]. Gardiner &

Waugh, 1938: 176.

Caryophyllia ambrosia ambrosia: Cairns, 1979: 59.

New records

V-2668, 12, IOM; V-—2671, 1, IOM; V—2816, 4, IOM. Reference specimens:

Gardiner & Waugh’s (1938) Caryophyllia clavus: JM-119, 3, BM 1950.1.9.159-183;

JM-122, 2, BM 1950.1.9.75-80; JM-185, 3, BM 1950.1.9.134-158.

Distribution

Indian Ocean: off Pemba and Zanzibar, Tanzania (Gardiner & Waugh 1938);

Saya de Malha Bank; Madagascar Plateau; Gulf of Aden (Gardiner & Waugh 1938);

Maldive Islands (Van der Horst 1931); Laccadive Islands (Alcock 1898); 430-2 000 m.

Elsewhere: western and eastern Atlantic; 1 600-2 670 m (Zibrowius 1980).

Caryophyllia grandis Gardiner & Waugh, 1938

Caryophyllia clavus: von Marenzeller, 1904: 281 [part.—Valdivia—186, pl. 16 (fig. 9, 91)].

Caryophyllia grandis Gardiner & Waugh, 1938: 177, pl. 1 (fig. 2). Pillai & Scheer, 1976: 16. Zibrowius

& Gili, 1990: 32.

New records

V-2631, 12, IOM, 1, USNM 91521; 12 miles (19 km) north of Durban,

183-220 m, 4, USNM 62497. Reference specimens: 4 syntypes of C. grandis from

JM-145, BM 1950.1.9.211-225.

Remarks

Caryophyllia grandis is similar to C. ambrosia, both having unattached coralla of

approximately the same size. To reiterate and add to Gardiner & Waugh’s (1938) dis-

tinctions, C. grandis has a brownish theca; more crowded septa (and more septa than

C. ambrosia at a corresponding size, sometimes up to a full fifth cycle); narrower pali;

less exsert septa; and a shallower depth range (183-595 m vs 430-2 670 m). Compari-

sons to another unattached, cornute species, C. valdiviae, known only from the

Walvis Ridge and off north-west Africa (882-1 230 m), are made by its authors Zibro-

wius & Gili (1990).

Distribution

Off Natal, South Africa (Zibrowius & Gili 1990); off south-eastern Mozambique;

Maldive Islands (Gardiner & Waugh 1938); ?off western Sumatra (Von Marenzeller

1904); 183-595 m.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 235

Caryophyllia scobinosa Alcock, 1902a

Caryophyllia scobinosa Alcock, 1902a: 90; 1902c: 8, pl. 1 (fig. 2, 2a). Gardiner & Waugh, 1938:

177-178. Keller, 1976: 17-18.

Caryophyllia clavus: von Marenzeller, 1904: 281 [part.—Valdivia—246, pl. 16 (fig. 9c—g, but not 9h)].

New records

V-2650, 1, IOM; V—2699, 2, IOM; AB-365D, 11, USNM 91519.

Distribution

Indian Ocean: off Pemba and Dar es Salaam, Tanzania (Von Marenzeller 1904;

Gardiner & Waugh 1938; Keller 1976); south-west of Madagascar; off Walters Shoal,

Madagascar Plateau; 535-960 m. Elsewhere: Celebes and Sulu Seas; 786-805 m.

Caryophyllia sp. cf. C. cornuformis Pourtalés, 1868

Fig. 3C, F

?Caryophyllia cornuformis: Gardiner & Waugh, 1938: 179, text-fig. 2.

New records

V-2626, 1, IOM; AB-370G, 32, USNM 91525, 1, SAM—H4585; AB-370H, 1,

USNM 91526.

Remarks

Both Cairns (1979) and Zibrowius (1980) stated that Gardiner & Waugh’s (1938)

south-west Indian Ocean specimens of C. cornuformis were not that species, but did

not state which species the John Murray specimens are. Comparisons of our new

records to typical western Atlantic C. cornuformis show them to be very similar in

size, shape, and septal and palar arrangement. The south-west Indian Ocean speci-

mens differ only in having a thinner, sometimes ridged, non-porcellaneous theca, and

some specimens have an intact base. It is suggested that two similar species may be

involved: one very similar, if not identical, to C. cornuformis, which always has a

broken base and irregular septal symmetry, and another as yet unnamed species (illus-

trated by Gardiner & Waugh 1938) that has an intact base and hexameral (three

cycles) symmetry.

Distribution

Indian Ocean: off south-western Mozambique; ?off Pemba and Zanzibar, Tanza-

nia (Gardiner & Waugh 1938); ?Gulf of Aden (Gardiner & Waugh 1938); 91-347 m.

Distribution of C. cornuformis: western Atlantic from Brazil to 63°N at 37-931 m

(Cairns 1979) and eastern Atlantic in area bounded by Celtic Sea, Azores, and

Morocco at 1 300-2 200 m (Zibrowius 1980).

Caryophyllia profunda Moseley, 1881

Caryophyllia profunda Moseley, 1881: 138-139, pl. 1 (fig. 6) [part.—not specimen from Cape Verde

Islands]. Von Marenzeller, 1904: 298. Zibrowius, 1974a: 751-755, pl. 1 (figs 1-10) [synonymy].

Cairns, 1982: 17-19, pl. 5 (figs 1-5) [synonymy]. Zibrowius & Gili, 1990: 25-26, pl. 4 (figs L-R).

Caryophyllia cyathus: von Marenzeller, 1904: 295, pl. 16 (fig. 6).

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

New records

V-2686, 12, IOM, 1, USNM 91527; V-2722, 3, IOM; V-2731, 11, IOM;

V-2733, 9, IOM.

Distribution

Indian Ocean: Agulhas Bank and Cape Agulhas (Von Marenzeller 1904); Mada-

gascar Plateau; St Paul and Amsterdam Islands (Zibrowius 1974a); 80-755 m.

Elsewhere: circum-Subantarctic; 35-1 116 m (Cairns 1982).

Caryophyllia rugosa Moseley, 1881

Figs 3

Caryophyllia rugosa Moseley, 1881: 141-143, pl. 1 (fig. 8). Cairns, 1984: 11-13, pl. 2 (figs A-B), pl. 4

(fig. I) [synonymy].

New records

AB-371E, 6, USNM 77212; AB-371F, 5, USNM 77213; Manihine 381-1, 2,

USNM 91528; MN-ZD4, 2, USNM 91529; MN—ZQ8a, 1, USNM; JM-157 (attached

to a colony of Balanophyllia diffusa), 1, BM 1950.1.6.35.

Distribution

Indian Ocean: off Zululand; off south-eastern Mozambique; off Mombasa,

Kenya; off Maldive Islands; 95-250 m. Elsewhere: Hawaiian Islands; Philippines;

Ceram Sea; Japan; Bikini; 71-230 m (Cairns 1984).

Caryophyllia elongata sp. nov.

Fig. 4A-B

Records

Holotype: V—2716, 1, IOM.

Description

Corallum attached, subcylindrical, straight, and elongate: 25,6 mm in height,

9,3 x 7,9 mm in calicular diameter, and 5,7 mm in pedicel diameter. Costae poorly

developed, only slightly ridged Ci and C2 present near calice. Theca otherwise porcel-

laneous, covered by low, rounded granules. Corallum primarily white, but theca light

brown near calice.

Septa hexamerally arranged in four complete cycles (48 septa) according to the

formula: $i>S2>S.:>Ss3. $i moderately exsert (2-2,5 mm), with slightly sinuous inner

edges that reach about three-quarters distance to columella. S2 slightly less exsert and

equally sinuous, extending about two-thirds distance to columella. S3 least exsert and

smallest septa, extending only about half distance to columella. Ss twice as exsert and

slightly wider than Ss, both cycles having sinuous inner edges. Septal granules promi-

nent, usually rectangular in profile, sometimes extending as short ridges paralleling

inner septal edge. Twelve P3 form a deeply recessed palar crown, each palus about

1,2 mm wide, highly sinuous, and separated from its respective Ss; by a deep and

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN DSi]

narrow notch. Each couple of P3; within a system are slightly closer to each other than

to those of adjacent systems, giving the impression of paired pali. Fossa quite deep,

containing the palar crown and even deeper columella, consisting of two narrow fas-

cicular elements.

Remarks

Among the 56 Recent species of Caryophyllia listed by Cairns (1991), 18 have

attached coralla with septa hexamerally arranged in four cycles, and in only five of

these species are the S; larger than the Ss, the set of characters found in Caryophyllia

elongata sp. nov.: C. polygona Pourtalés, 1878; C. calveri Duncan, 1873; C. alberti

Zibrowius, 1980; C. atlantica (Duncan, 1873); and C. panda Alcock, 1902a. Caryo-

phyllia elongata is distinguished from these species by having Si: larger than S2 (S: and

S2 are equal in size in most species), a very deep fossa, and deeply recessed, ‘paired’

pali.

Etymology

The species name elongata (from the Latin elongatus, prolonged) is an allusion to

the elongate corallum of this species.

Distribution

Known only from the type locality of 33°17’S 44°55’E (Madagascar Plateau, off

Walters Shoal), 630-680 m.

Subgenus Caryophyllia (Premocyathus)

Caryophyllia (Premocyathus) zanzibarensis Zou, 1984 [comb. nov.].

Caryophyllia compressa Gardiner & Waugh, 1938: 180, pl. 2 (fig. 4) [junior secondary homonym of

Caryophyllia (Premocyathus) compressus Yabe & Eguchi, 1942].

?Trochocyathus pileus: Gardiner & Waugh, 1938: 187.

Caryophyllia zanzibarensis Zou, 1984: 52, 53 [replacement name for C. compressa Gardiner &

Waugh, 1938].

New record

Manthine 381-3, 4, USNM 91542.

Remarks

This species is transferred to the subgenus Premocyathus based on its highly com-

pressed corallum and carinate thecal edges. Other Recent species in this subgenus

include: C. (P.) compressus Yabe & Eguchi, 1942; C. (P.) spinacarens (Moseley,

1881), comb. nov.; C. (P.) burchae Cairns, 1984; and ?C. (P.) dentiformis (Alcock,

1902b). Caryophyllia (Premocyathus) zanzibarensis is most similar to C. (P.) spina-

carens, differing primarily in having fewer septa and in being more compressed.

Distribution

Off north-eastern Tanzania, Maziwi Island (Gardiner & Waugh 1938);

238-302 m.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. A-B. Madracis sp. A, V-2697, IOM, branch and calicular views. AX 10, B X 20.

C, F. Caryophyllia sp. cf. C. cornuformis, V-2626, IOM, lateral and calicular views. C x 5,8, F x 6,7.

D. Letepsammia formosissima, V-2608, USNM 91506, calice. 1,3. E. Anthemiphyllia dentata,

V-2804, IOM, calice. x 1,4. G. Culicia sp. cf. C. natalensis, AB—421A, USNM 91515, colony. x 0,9.

H. Caryophyllia ambrosia ambrosia, V-2668, IOM, calice. x 1,3. I. Caryophyllia rugosa, AB-371E,

USNM 77212, calice. x 9,4.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 239

Fig. 4. A-B. Caryophyllia elongata sp. nov., V-2716 (holotype), lateral and stereo calicular views.

Ax1,6, Bx3,9. C-D. Trochocyathus sp. A, V-2662, IOM, lateral and calicular views. C x 1,7,

D x 4,5. E, H. Trochocyathus sp. cf. T. rawsonii, V-2733, USNM 91568, lateral and calicular views.

E x 3,6, Hx 3,3. F-G. Conotrochus brunneus, AB-365D, USNM 91556, lateral and calicular views.

EG3 55 Geas29:

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Trochocyathus

Trochocyathus rhombocolumna Alcock, 1902c

Trochocyathus rhombocolumna Alcock, 1902c: 16, pl. 2 (fig. 12).

Paracyathus gardineri: Gardiner & Waugh, 1938: 183-184 (part. —JM-157, pl. 3 (fig. 5)).

New records

AB-371E, 5, USNM 91563; AB-371F, 2, USNM 91564. Reference material:

Gardiner & Waugh’s (1938) P. gardineri: JM-157, 4, BM 1950.1.9.724-826; JM-106,

1, BM 1950.1.9.718.

Remarks

Specimens herein reported from off Mozambique were compared to those iden-

tified by Gardiner & Waugh (1938) as Paracyathus gardineri from the Maldive Islands

(JM-157) and found to be conspecific; however, the John Murray specimen from off

Tanzania (JM-—106) was too damaged to identify. Several specimens from the Maldive

Islands are extremely similar to the figured holotype of T. rhombocolumna. A pecu-

liarity of this species is that not only are the Ss slightly wider that the S3, but those S,

adjacent to S: are wider than those adjacent to S2 (about equal in width as an Sz).

As noted by Cairns (1984), the Indian Ocean specimens reported by Gardiner &

Waugh (1938) as P. gardineri Vaughan, 1907, are not conspecific with that Hawaiian

species, the former differing in having a basal attachment, more crowded septa, and

transverse costae.

Distribution

Off south-western Mozambique; Maldive Islands (Gardiner & Waugh 1938);

110-229 m. Elsewhere: Sulu Archipelago (Alcock 1902c); 522 m.

Trochocyathus sp. A

Fig. 4C-—D

Records

V-2662, 1, IOM; V-2803, 1, USNM 91567; AB-371F, 2, USNM 91565;

MN-ZDD3, 1, USNM 91566.

Diagnosis

Corallum ceratoid and firmly attached either basally or laterally by epithecal

bands. Coralla up to 10,3 x 9,1 mm in calicular diameter and 15 mm in height. Costae

granular and well developed; transverse sculpturing not present. Corallum light

brown, especially near calice. Septa hexamerally arranged in four cycles according to

the formula: Si and S2>S;3 and Ss. All septa have straight, vertical inner edges, except

the S3, which are sinuous. Pali occur before the highest three cycles, the P2 and P3

about the same size, the P; about twice that width. All pali extend to columella.

Columella papillose, consisting of 5-12 elements in an elliptical field.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 241

Remarks

Trochocyathus sp. A is similar to T. rhombocolumna in having robust palar and

columellar elements and having 48 septa, but differs in having: P3 recessed from the

columella, forming chevrons with the P2; longitudinal (not transverse) costae; less

crowded septa; brown theca; and equally wide Si: and S:.

Distribution

Off Zululand, South Africa; off south-eastern Mozambique; south-west of Mada-

gascar; Saya de Malha Bank; 74-315 m.

Trochocyathus rawsonii sensu Gardiner, 1904

Non Trochocyathus rawsonii Pourtalés, 1874: 35.

Trochocyathus rawsonii: Gardiner, 1904: 100-103, pl. 1 (fig. 2a—b), pl. 2 (figs A—-K).

New record

MN-SM7232, 3, USNM 77220. Reference specimen: 7. rawsonii of Gardiner

(1904; pl. 1 (fig. 2)), 1, BM 1950.1.10.112.

Remarks

As noted by Cairns (1979) and Zibrowius & Gili (1990), the South African speci-

mens reported by Gardiner (1904) as T. rawsonii are incorrectly identified and

probably represent an undescribed species. The South African specimens differ in

having elongate, often curved coralla attached by a slender pedicel; better developed

costae; and a paucity of Ss. But, as stated above, until this genus is better known, we

prefer not to introduce a new name for this species at this time, but await Zibrowius’

anticipated faunistic revision, which will include many more specimens of this taxon.

Distribution

Off South Africa from Cape of Good Hope to Buffalo River (Gardiner 1904);

560-620 m (depths of Gardiner’s specimens unknown).

Trochocyathus sp. cf. T. rawsonii Pourtales, 1874

Fig. 4E, H

?Trochocyathus rawsonii Pourtalés, 1874: 35, pl. 6 (figs 7-10). Cairns, 1979: 77-79, pl. 13 (figs 5-7),

pl. 14 (figs 1-6) [synonymy].

Non Trochocyathus rawsonii Pourtalés. Gardiner, 1904: 100-103.

New records

V-2608, 4, IOM; V—2731, 1, IOM; V—2733, 1, USNM 91568.

Remarks

Although Gardiner’s (1904) records of T. rawsonii are thought to be misiden-

tified, three specimens collected off Madagascar and the Madagascar Plateau are

indistinguishable from small specimens of the attached, trochoid form of T. rawsonii

described and figured by Cairns (1979: 77, pl. 13 (fig. 6)). The largest of the three

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

specimens (V—2608) is 11,6 mm in calicular diameter and 13,4 mm in height, having

48 septa arranged in four complete cycles.

Distribution

Indian Ocean: off north-western Madagascar and off Walters Shoal, Madagascar

Plateau; 750-780 m. Distribution of T. rawsonii: Georgia to Brazil; 82-622 m (Cairns

1979).

Genus Stephanocyathus

Stephanocyathus (Odontocyathus) nobilis (Moseley, 1873)

Fig. SD-E

Ceratotrochus nobilis Moseley, 1873: 402, text-fig. 3.

Stephanotrochus nobilis: Moseley, 1881: 155, pl. 3 (fig. 3a—b).

Stephanotrochus nitens Alcock in Wood-Mason & Alcock, 189la: 7-8. Alcock, 1898: 18-19, pl. 2

(fig. 6, 6a); 1902d, text-fig. 92.

Stephanotrochus oldhami Alcock, 1894: 187-188; 1898: 19-20.

Stephanocyathus nobilis: Gardiner & Waugh, 1938: 189-192, pl. 6 (figs 13, 15). Pillai & Scheer, 1976:

16. Zibrowius, 1980: 101-103, pl. 51 (figs A-K) [synonymy].

Non Stephanocyathus nobilis: Boshoff, 1981: 39-40 (= S. explanans). Zou, 1988: 74-75

(= S. weberianus).

Stephanocyathus (Odontocyathus) nobilis: Cairns, 1979: 110-111, pl. 20 (figs 7, 10).

New records

V-2629, 6, IOM, 1, USNM 91543; V-2653, 3, IOM; V-2814, 2, IOM;

AB-399B, 1, USNM 91544; AB-399C, 28, USNM 91545, 2, SAM-—H4576.

Remarks

Although Zibrowius (1980) expressed reservation about the authenticity of Indian

Ocean records of S. nobilis (type locality, Azores), comparison of Indian Ocean speci-

mens with those from the eastern Atlantic, including the holotype, convince us that

they are the same species, a conclusion also reached by Zou (1988); however, we do

not include western Pacific specimens as S. nobilis. Although not examined (types

presumably deposited in the Indian Museum, Calcutta), we concur with Gardiner &

Waugh (1938) that S. nitens Alcock, 1891a (in Wood-Mason & Alcock), and S. old-

hami Alcock, 1894, are undoubtedly junior synonyms of S. nobilis, but agree with

Zibrowius (1980) that S$. weberianus Alcock, 1902a, and S. campaniformis von Maren-

zeller, 1904, are distinct species.

Distribution

Indian Ocean: off south-eastern Mozambique; off Zanzibar, Tanzania (Gardiner

& Waugh 1938); off Mombasa, Kenya (Gardiner & Waugh 1938); off south-western

Madagascar; off Saya de Malha Bank; Gulf of Aden (Gardiner & Waugh 1938);

Arabian Sea off India (Wood-Mason & Alcock 1891a; Alcock 1894); Maldive Islands

(Gardiner & Waugh 1938; Pillai & Scheer 1976); 609-2 000 m. Elsewhere: off

England; Azores; and Gulf of Guinea (Zibrowius 1980); off Brazil (Cairns 1979);

763-2 200 m.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 243

Stephanocyathus (Odontocyathus) campaniformis (von Marenzeller, 1904)

Fig. 5A—B

Stephanotrochus campaniformis von Marenzeller, 1904: 302-304, pl. 18 (fig. 20, 20a).

Stephanocyathus campaniformis: Zibrowius & Gili, 1990: 32-35, pl. 4 (figs A-K), pl. 5 (figs E—-J)

[synonymy].

New record

V—2674, 1, IOM.

Remarks

Stephanocyathus campaniformis is very similar to S. nobilis, the major differences

being its smaller adult size (distance from centre of base to edge break

= 7,5-11,5 mm vs 10-15 mm for adult S. nobilis), and its characteristic bell shape,

usually with a much higher H: W ratio than S. nobilis (i.e. 0,65-1,3 mm _ vs

0,42-0,61 mm for S. nobilis). The south-west Indian Ocean specimen, 29,5 mm in

calicular diameter and 19,2 mm in height (H : W = 0,65), was compared to topotypic

specimens of S$. campaniformis reported by Zibrowius & Gili (1990) from the Walvis

Ridge (USNM 86873-86876), the Indian Ocean specimen being very similar to a

specimen figured by Zibrowius & Gili (1990, pl. 4 (figs F—G)).

Distribution

Indian Ocean: south-west Indian Ridge south of Madagascar Plateau;

1 600-1 610 m. Elsewhere: Walvis Ridge, off Namibia; 882-1 230 m (Zibrowius &

Gili 1990).

Stephanocyathus (Acinocyathus) spiniger (von Marenzeller, 1888)

Stephanotrochus spiniger von Marenzeller, 1888: 20-21.

Odontocyathus spiniger: Eguchi, 1968: C39—C40, pl. C20 (figs 12-14), pl. C23 (figs 1-2) [synonymy].

Stephanocyathus spiniger: Boshoff, 1981: 39.

Stephanocyathus (Acinocyathus) spiniger: Wells, 1984: 209, figs 2.10-13 [synonymy]. Cairns & Parker,

1992: 26-27, pl. 7 (figs g-i) [synonymy].

New records

V-2635, 4, IOM; AB-365D, 14, USNM 77215, 1, SAM-—H4595.

Distribution

Indian Ocean: off Durban, South Africa (Boshoff 1981); off south-eastern

Mozambique; off south-western and northern Madagascar; 210-695 m. Elsewhere:

Japan; Philippines; Indonesia; Great Australian Bight; 120-560 m (Cairns & Parker

1992).

Stephanocyathus (Acinocyathus) explanans (von Marenzeller, 1904)

Stephanotrochus explanans von Marenzeller, 1904: 304-307, pl. 18 (fig. 19a—b).

Stephanocyathus explanans: Gardiner & Waugh, 1938: 192.

Stephanocyathus nobilis: Boshoff, 1981: 39.

New records

19 km north of Durban, South Africa, 183-220 m, 4, USNM 62500.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Although having the same size range, S. explanans differs from S. spiniger in

having shorter, thinner costal spines; having much less exsert septa (Si = S2, whereas

Si of S. spiniger are much larger than S2); and in lacking corallum pigmentation.

Distribution

Off Durban, South Africa (Boshoff 1981); off Pemba and Zanzibar, Tanzania

(Von Marenzeller 1904); west of Sumatra (Von Marenzeller 1904); 183-614 m.

Genus Labyrinthocyathus

Labyrinthocyathus delicatus (von Marenzeller, 1904)

Ceratotrochus delicatus von Marenzeller, 1904: 302, pl. 18 (fig. 18).

Cyathoceras cornu: Gardiner, 1904: 121-122.

Labyrinthocyathus sp. Cairns, 1979: 70, pl. 11 (figs 10-11).

Labyrinthocyathus delicatus: Zibrowius & Gili, 1990: 44.

New records

V-2637, 2, IOM; AB-357E, 1, USNM 77219; MN-SM162, 1, USNM 91546.

Distribution

Known only from the Indian Ocean off South Africa, from Cape Town to Durban

(Von Marenzeller 1904; Gardiner 1904; Cairns 1979), and off south-eastern Mozam-

bique; 155-1 000 m.

Genus Deltocyathus

Deltocyathus andamanicus Alcock, 1898

Fig. 5F

Deltocyathus andamanicus Alcock, 1898: 16-17, pl. 1 (fig. 5, Sa). Gardiner & Waugh, 1938: 196. Pillai

& Scheer, 1976: 16.

Non Deltocyathus sp. cf. D. andamanicus: Cairns, 1984: 15, pl. 3 (figs A-B).

New record

Manihine 381-63, 1, USNM 91548.

Remarks

At least eight species of Deltocyathus have been reported from the Indian Ocean:

D. andamanicus Alcock, 1898; D. rotulus (Alcock, 1898); D. murrayi Gardiner &

Waugh, 1938; D. varians Gardiner & Waugh, 1938; D. sarsi (Gardiner & Waugh,

1938); D. nascornatus (Gardiner & Waugh, 1938); Deltocyathus sp. sensu von Maren-

zeller, 1904; and D. italicus sensu Zibrowius, 1974a. The species D. minutus Gardiner

& Waugh, 1938, and D. lens Alcock, 1902, pertain to the genus Feponocyathus (see

Cairns 19896: 30). The specimen reported herein, 15,5 mm in calicular diameter, cor-

responds to the original description and illustrations of Alcock’s D. andamanicus.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 245

Distribution

Off Zanzibar, Tanzania (Gardiner & Waugh 1938); Gulf of Aden (Gardiner &

Waugh 1938); Maldive Islands (Gardiner & Waugh 1938; Pillai & Scheer 1976);

Andaman Sea (Alcock 1898); 240-1 463 m.

Deltocyathus rotulus (Alcock, 1898)

Fig. 51

Trochocyathus rotulus Alcock, 1898: 16, pl. 2 (fig. 1, 1a).

Deltocyathus fragilis Alcock, 1902a: 99-100; 1902c: 21, pl. 2 (fig. 15, 15a).

Deltocyathus rotulus: van der Horst, 1931: 6. Gardiner & Waugh, 1938: 196.

New records

AB-365C, 9, USNM 91549; AB-389C, 1, USNM 91550. Reference material:

Gardiner & Waugh’s (1938) D. rotulus from JM-119, 2, BM 1950.1.9.1159-1162.

Remarks

We concur with Gardiner & Waugh’s (1938) evaluation that D. fragilis is a junior

synonym of D. rotulus, even though the type specimens of each species have a differ-

ent number of septa: 96 and 72, respectively. Of the approximately 15 valid Recent

species in the genus, only one other, D. magnificus Moseley, 1876, has five cycles of

septa. Deltocyathus rotulus is distinguished from D. magnificus by having a scalloped

calicular edge and in lacking the V-shaped deltoid septal fusions characteristic of the

genus.

Distribution

Indian Ocean: off Durban, South Africa; off south-eastern Mozambique; off Zan-

zibar, Tanzania (Gardiner & Waugh 1938); Gulf of Aden (Gardiner & Waugh 1938);

Maldive Islands (Alcock 1898; Gardiner & Waugh 1938); off Sri Lanka (Van der

Horst 1931); 510-1 986 m. Elsewhere: Flores Sea; 794 m (Alcock 1902a, 1902c).

Deltocyathus sp. A

Fig. 5SG—H

?Deltocyathus italicus: Zibrowius, 1974a: 757.

Deltocyathus lens: Gardiner & Waugh, 1938: 198.

Records

V-2662, 9, IOM; MN-ZV5, 1, USNM 91551.

Remarks

This species is similar to the Miocene D. italicus (Michelotti, 1838) and Recent

Atlantic specimens described by Cairns (1979) as Deltocyathus sp. cf. D. italicus [cited

by Zibrowius (1980) and Zibrowius & Gili (1990) as D. conicus Zibrowius, 1980].

Deltocyathus italicus and the south-west Indian Ocean specimens are similar in size

and shape, having a highly conical corallum with a pointed base; however, Delto-

cyathus sp. A differs in having granular, rounded costae (not dentate, ridged costae as

in D. italicus); relatively small P3 not fused to their corresponding S3 (in D. italicus, P3

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

are quite large and solidly fuse to their S3); and a fossa moderate in depth (the central

calice of D. italicus is usually elevated, having no fossa). Although not described or

illustrated, it is likely that Zibrowius’ (1974a) ‘conical species’ from north-west of

Madagascar is the same. No other Indian Ocean species of Deltocyathus is known to

have a conical corallum.

Distribution

Known only from the Indian Ocean off Durban, South Africa; off Zanzibar (Gar-

diner & Waugh 1938); ?off south-western and north-western Madagascar (Zibrowius

1974a); 207-315 m.

Genus Desmophyllum

Desmophyllum cristagalli Milne Edwards & Haime, 1848a

Desmophyllum cristagalli Milne Edwards & Haime, 1848a: 253, pl. 7 (fig. 10, 10a). Zibrowius, 1974a:

758-761, pl. 3 (figs 1-10); 1980: 117-121, pl. 61 (figs A-O). Cairns, 1979: 117-119, pl. 21

(figs 7-8), pl. 22 (fig. 8) [synonymy]. Zibrowius & Gili, 1990: 35-36.

Desmophyllum capense Gardiner, 1904: 96-97.

?Desmophyllum sp. Gardiner & Waugh, 1938: 176.

Non Desmophyllum cristagalli Milne Edwards & Haime. Boshoff, 1981: 37.

New records

V-2699, 1 IOM; V—2716, 6, IOM; V—2722, 1, IOM.

Distribution

Indian Ocean: off Cape of Good Hope (Gardiner 1904); off Pemba, Tanzania

(Gardiner & Waugh 1938); off Walters Shoal, Madagascar Plateau; Gulf of Aden

(Gardiner & Waugh 1938); Maldive Islands (Gardiner & Waugh 1938); 81-2 000 m.

Elsewhere: cosmopolitan; 60-2 460 m (Cairns 1979).

Genus Conotrochus

Conotrochus brunneus (Moseley, 1881)

Fig. 4F-G

Pleurocyathus brunneus Moseley, 1881: 159-160, pl. 2 (fig. la—c).

Phloeocyathus hospes: Alcock, 19026: 116-117.

Ceratotrochus (Phloeocyathus) hospes: Alcock, 1902c: 12, pl. 2 (fig. 8, 8a).

Conotrochus brunneus: Gardiner & Waugh, 1938: 175-176, pl. 5 (figs 11-12). Zibrowius, 1980: 79.

New record

AB-365D, 2, USNM 91556.

Distribution

Indian Ocean: off south-west Madagascar; Maldive Islands (Gardiner & Waugh

1938); 475-695 m. Elsewhere: Indonesia (Moseley 1881; Alcock 1902b, 1902c);

110-1 089 m.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 247

Genus Aulocyathus

Aulocyathus recidivus (Dennant, 1906)

Fig. 5C

Ceratotrochus recidivus Dennant, 1906: 159-160, pl. 6 (figs la—b, 2a—c). Zibrowius, 1980: 107.

Aulocyathus recidivus: Cairns, 1982: 25-26, pl. 7 (figs 7-9), pl. 8 (fig. 1). Cairns & Parker, 1992:

22-24, pl. 6 (figs d, h) [synonymy].

New record

Unspecified R.V. Anton Bruun station ‘off Madagascar’, depth unknown, 1,

USNM 91555.

Remarks

Aulocyathus recidivus is distinguished from A. juvenescens by its much larger and

stouter corallum and by having a greater number of septa. Three additional species

are known in this genus from the eastern Atlantic and western Pacific (see Zibrowius

1980).

Distribution

Indian Ocean: ‘off Madagascar’; depth unknown. Elsewhere: off South Australia

and Tasmania (Dennant 1906; Cairns & Parker 1992); Macquarie Ridge (Cairns

1982); 128-1 000 m.

Aulocyathus juvenescens von Marenzeller, 1904

Aulocyathus juvenescens von Marenzeller, 1904: 301-302, pl. 18 (fig. 17). Zibrowius, 1980: 107.

New record

Manihine 381-3, 2, USNM 91554. Reference material: syntypes of A. juvenescens

from Valdivia—243 and 245 (ZMB 5064, 7032).

Remarks

The Manihine specimens, collected only several kilometres from the type locality

(Valdivia station 245), are the only specimens to have been reported subsequent to its

description in 1904.

Distribution

Known only from the Indian Ocean off Pemba and Zanzibar, Tanzania (Von

Marenzeller 1904); 302-463 m.

Genus Dasmosmilia

Dasmosmilia variegata (Pourtalés, 1871)

Fig. 6C

Parasmilia variegata Pourtalés, 1871: 21, pl. 1 (fig. 13).

Non Dasmosmilia variegata: Gardiner & Waugh, 1938: 172-173.

Dasmosmilia variegata: Cairns, 1979: 134-135, pl. 25 (figs 4-7, 10), pl. 26 (fig. 1) [synonymy]. Zibro-

wius, 1980: 71-72, pl. 30 (figs A-K) [synonymy].

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. A-B. Stephanocyathus campaniformis, V—2673, IOM, calicular and basal views. Both X 1,7.

C. Aulocyathus recidivus, Anton Bruun station ‘off Madagascar’, USNM 91555, lateral view. x 2,9.

D-E. Stephanocyathus nobilis, AB-399C, USNM 91545, calicular and lateral views. Both x 1,5.

F. Deltocyathus andamanicus, Manihine 381-63, USNM 91548, calice. x 2,2. G—H. Deltocyathus

sp. A, MN-ZV5, USNM 91551, calicular and lateral views. Both 3,8. I. Deltocyathus rotulus,

AB-389C, USNM 91550, calice. x 1,5.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 249

New record

V—2644, 1, IOM.

Remarks

The figured specimen, measuring 10,1 x 8,3 mm in calicular diameter and

14,8 mm in height, was compared to specimens from both the western and eastern

Atlantic and found to be indistinguishable in most characters, except that the Indian

Ocean specimen is firmly attached to a substrate (a cylindrical bryozoan), not asex-

ually regenerated from a parent corallum fragment, as is typical for most other known

specimens.

Distribution

Indian Ocean: off south-western Madagascar; 330-335 m. Elsewhere: off Florida,

Venezuela, Brazil, Cape Verde Islands, and the Azores; 110-600 m (Cairns 1979;

Zibrowius 1980).

Genus Asterosmilia

Asterosmilia marchadi (Chevalier, 1966)

Fig. 6A—B

Ceratotrochus johnsoni: Gardiner, 1904: 118-119, pl. 1 (fig. Sa—c), pl. 2 (fig. M). Gardiner & Waugh,

1938: 188.

Dasmosmilia marchadi Chevalier, 1966: 944—949, pl. 5 (figs 3-4).

Asterosmilia marchadi: Cairns, 1979: 140-142, pl. 26 (figs 7, 9-10). Zibrowius, 1980: 141-142, pl. 74

(figs A-K) [synonymy].

New records

V-—2634, 1, IOM; AB-372J, 2, USNM 91557; AB-372L, 26, USNM 91559, 1,

SAM-—H4594; AB-373B, 1, USNM 91560; MN-—ZD8, 1, USNM 91561; MN-ZH27,

1, USNM 91562. Reference specimens: C. johnsoni of Gardiner & Waugh (1938):

JM-106, 2, BM 1950.1.9.1065-1069; JM107, 1, BM 1950.1.9.1284; C. johnsoni of

Gardiner (1904) from Cape Natal, 1, BM 1950.1.10.118.

Remarks

Zibrowius (1980: 141, 142) noted the extreme resemblance of the Indian Ocean

specimens reported by Gardiner (1904) and Gardiner & Waugh (1938) to Atlantic

A. marchadi, but did not commit to that identification. Comparisons of the south-west

Indian Ocean specimens reported herein and Gardiner (1904) and Gardiner &

Waugh’s (1938) specimens, with typical eastern and western Atlantic specimens of

A. marchadi, show no significant differences. Therefore, A. marchadi is considered to

have a disjunct distribution in both the Atlantic and western Indian oceans.

Distribution

Indian Ocean: South Africa off Bisho, Cape Natal (Gardiner 1904), and Zulu-

land; off south-eastern Mozambique; off Pemba, Tanzania (Gardiner & Waugh 1938);

off the Maldive Islands (Gardiner & Waugh 1938); 57-229 m. Elsewhere: eastern

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Atlantic from Spanish Sahara to Gabon (Zibrowius 1980); western Atlantic from off

Florida and the northern coast of South America (Cairns 1979); 79-229 m.

Genus Solenosmilia

Solenosmilia variabilis Duncan, 1873

Fig. 6D

Solenosmilia variabilis Duncan, 1873: 328, pl. 42 (figs 11-18). Von Marenzeller, 1904: 310-311, pl. 15

(fig. 4, 4a). Zibrowius, 1974a: 768-769; 1980: 143-145, pl. 75 (figs A—N) [synonymy]. Cairns,

1979: 136-138, pl. 26 (figs 2-4) [synonymy]. Scheer & Pillai, 1983: 160. Cairns & Parker, 1992:

29-30, pl. 8 (figs d-e).

Solenosmilia Jeffreyi Alcock, 1898: 27-28, pl. 3 (fig. 3, 3a—b).

Non Solenosmilia variabilis Duncan. Gardiner & Waugh, 1939: 229-230.

New records

MN-SM129, 1 branch, USNM 77211; MN-SM162, 2 colonies, USNM 91690;

MN-SM226, branch fragments, USNM 91691.

Distribution

Indian Ocean: off South Africa from Agulhas Bank (Von Marenzeller 1904) to

off Durban; off Somalia (Von Marenzeller 1904); Laccadive Sea (Alcock 1898);

366-1 079 m. Elsewhere: amphi-Atlantic; South Australia; circum-Subantarctic;

220-2 165 m (Cairns & Parker 1992).

Genus Goniocorella

Goniocorella dumosa (Alcock, 1902c)

Fig. 6E

Pourtalosmilia dumosa Alcock, 1902c: 36-37, pl. 5 (fig. 33, 33a).

Goniocorella dumosa: Cairns, 1982: 31-34, pl. 9 (figs 7-9), pl. 10 (figs 1-2) [synonymy].

New record

MN-SM174, 4 branches, USNM 77221.

Distribution

Indian Ocean: off Bisho, South Africa; 760 m. Elsewhere: off Japan, Banda Sea,

and New Zealand region; 100-638 m (Cairns 1982).

Genus Rhizosmilia

Rhizosmilia robusta sp. nov.

Fig. 6F—I

Records

Holotype: AB—373B, 1, USNM 91681. Paratypes: AB-371F, 1, USNM 91682;

AB-408D, 2 corallites, USNM 91683; MN-ZB23, 1, SAM—H4572; MN-ZC10, 1,

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN Dok

USNM 91684; MN-ZD5, 1, USNM 91685; MN-ZDD3, 3, USNM 91689; MN-ZD7,

2, USNM 91686, 1, SAM-—H4573; MN-ZK20, 1, USNM 91687; MN-ZK21, 1,

USNM 91688.

Description

Corallites trochoid in shape, firmly attached through a massive pedicel and thin

expansive base. Largest corallite of holotypic colony 31,0 x 26,2 mm in calicular

diameter, 25,0 mm in height, and 16,8 mm in pedicel diameter. Lower pedicel and

base reinforced with concentric rings of hollow chambers formed by adding exothecal

dissepiments over raised costae (Fig. 6G), as is characteristic of the genus (Cairns

1978). Calice elliptical in outline, even in small specimens. Costae equal (0,4-0,5 mm

wide) and quite low, separated by very narrow (0,10—-0,12 mm), shallow intercostal

striae. Costae covered with low, rounded granules. Corallum white.

Septa arranged in five cycles according to the formula: S:i>S2>S3>Si>Ss, the

fourth cycle complete at a GCD of 8—9 mm and the fifth cycle complete at a GCD of

19-21 mm; Se not observed even in largest calice of 31 mm diameter. Si moderately

exsert (up to 2,7 mm above calicular edge), their inner edges vertical and straight,

extending to the columella. Septa of higher cycles progressively less exsert and

smaller, except for those Ss adjacent to Si, which are more exsert than their adjacent

S:. Inner edges of S2 also straight; inner edges of Ss and Su slightly sinuous; Ss

rudimentary, with irregularly shaped inner edges. Septal faces relatively smooth,

bearing very low and sparsely placed granules. Septa well spaced, each separated from

one another by approximately twice the septal thickness. Small paliform lobes

present deep in fossa before septa of penultimate cycle (Pa, if Ss present; P3, if only S4

present in a half-system) and in such a manner of insertion as described by Cairns

(1978) for R. gerdae. Paliform lobes sometimes dissected into three or four thin,

elongate ribbons, similar in shape to columellar elements, but occurring higher in

fossa.

Fossa deep, containing a trabecular columella. Vescicular endothecal dissepi-

ments present, giving corallum a low density.

Remarks

Although the holotype is a phaceloid colony of four corallites, all paratypes are

represented as individual corallites, either broken from a larger colony or not yet

having formed a colony.

Only two other species of Rhizosmilia are known: R. maculata (Pourtalés, 1874)

and R. gerdae Cairns, 1978, both species known only from relatively shallow water

(3-287 m) in the western Atlantic. Rhizosmilia robusta sp. nov. is most similar to

R. maculata, especially in corallum size and shape and septal, palar, and costal mor-

phology, but differs in having an entirely white corallum (that of R. maculata is

speckled brown), and in having fewer septa at a corresponding calicular diameter.

Rhizosmilia robusta attains its fifth cycle at a GCD of 19-21 mm, whereas R. macu-

lata attains its at a GCD of only 11 mm (Cairns 1977) and often has additional Se in

larger corallites.

A probable fourth species of Rhizosmilia was reported as Caryophyllia gigas by

Van der Horst (1931) from off Mauritius— Rhizosmilia gigas comb. nov. Because only

Dy, ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. A-B. Asterosmilia marchadi, V-—2634, IOM, lateral and calicular views. A X 1,5, B x 2,1.

C. Dasmosmilia variegata, V—2644, IOM, calice. x 4,7. D. Solenosmilia variabilis, MN-—SM162,

USNM 91690, partial colony. x 0,7. E. Goniocorella dumosa, MN-SM174, USNM 77221. x 1,6.

F-I. Rhizosmilia robusta sp. nov. F, I. AB—373B (holotype), lateral and calicular views. F x 0,9,

Ix1,5. G. MN-ZD7, USNM 91686, paratype illustrating exothecal roots. x 2,5. H.MN-ZDS5,

USNM 91685, calice of a paratype. X 2,2.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 253

the holotype is known, which has a broken base, the colonial nature of the species is

unknown, but the arrangement of paliform lobes (P4), endothecal dissepiments, and

epithecal rootlets at the broken pedicel, suggest a placement in Rhizosmilia. The holo-

type is deposited at the BM (1939.7.20.851).

Etymology

The species name robusta (from the Latin robustus, hard, strong, like oak) is an

allusion to the thick pedicels of this species, which contribute to a robust colony.

Distribution

Off Zululand, South Africa; off south-eastern Mozambique; Mozambique

Channel off Madagascar; 66-150 m. Type locality: 26°00'S 33°05’E (off south-eastern

Mozambique), 135 m.

Family Turbinoliidae

Genus Tropidocyathus

Tropidocyathus lessoni (Michelin, 1842)

Fig. 7C

Flabellum lessoni Michelin, 1842: 119.

Tropidocyathus lessoni: Gardiner & Waugh, 1938: 194. Cairns, 1989b: 33-34, pl. 16 (figs d—l)

[synonymy].

New records

MN-ZAS1, 1, USNM 91692; MN-ZB24, 1, USNM 91693; MN-ZD4, 1,

USNM 91694; MN-—ZD6, 2, USNM 91695; MN-ZD7, 4, USNM 91696; MN-ZD8, 6,

USNM 91697, 1, SAM-—H4583; MN-ZDD3, 4 USNM 91706; MN-ZDD4, 7,

USNM 91707, 2, SAM-—H4588; MN-ZDD7, 1, USNM 91708; MN-ZH18, 1,

USNM 91698; MN-ZH19, 1, USNM 91699; MN-ZH23, 4, USNM 91700;

MN-ZH26, 3, USNM 91701; MN-ZK20, 1, USNM 91702; MN-ZK25, 1,

USNM 91703; MN-ZV20, 2, USNM 91704; MN-—ZV21, 3, USNM 91705.

Remarks

This is a commonly collected south-west Indian Ocean shallow-water azooxan-

thellate solitary coral, easily distinguished from all other species by its pale-orange

corallum and prominent thecal edge crests. Endopachys grayi also has edge crests but

always has a white corallum, the characteristic porous dendrophylliid theca, and a

Pourtalés Plan.

Distribution

Indian Ocean: South Africa from off Natal to Zululand; off south-eastern

Mozambique (Cairns 1989b); off Zanzibar, Tanzania (Gardiner & Waugh 1938); off

Kenya and north-eastern Somalia (Cairns 1989b); 62-155 m. Elsewhere: western

Pacific from East China Sea to Indonesia; 68—421 m (Cairns 1989b).

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Thrypticotrochus

Thrypticotrochus multilobatus Cairns, 1989b

Figa/El

Thrypticotrochus multilobatus Cairns, 1989b: 37, pl. 19 (figs b—g).

New records

AB-365D, 3, USNM 91709; AB-370G, 2, USNM 91710; AB-399A, 1, USNM

(lost).

Remarks

All specimens were asexually regenerated from parent fragments, the largest

corallum 4,4 mm in calicular diameter.

Distribution

Indian Ocean: off south-western Madagascar; off south-eastern Mozambique;

347-925 m. Elsewhere: South China Sea; Philippines; off south-eastern Australia

(Cairns 19896); 130-507 m.

Genus Sphenotrochus

KEY TO THE SOUTH-WEST INDIAN OCEAN SPECIES OF SPHENOTROCHUS

1A. Each costa composed of asingle; elongate ridge..>. 22.3: 0.5...) ae eee 7

1B. Each costa composed of two or more irregular rows of short, narrow ridges.......

a eer eee Le Mery AV ane ELCs ea ate Os APOE S. gilchristi Gardiner, 1904

2A. Corallum cuneiform but full (appears swollen); full fourth cycle (48) of costae

present; columella massive, rising well above upper septal edges ................

BRS aso UE Der rar eats Sind pes ree ie S. aurantiacus von Marenzeller, 1904

2B. Corallum cuneiform but compressed; only four pairs of Cs present in end half-

systems (total = 32); columella only as thick as an S: and not as exsert as an Si

3A. Costae rounded, equal in width, and do not overlap adjacent costae; intercostal

striae broad (35-55 % width of a costa); corallum triangular, faces diverging at a

constantianclemromibasei meres aoe oe oe ee S. evexicostatus sp. nov.

3B. Costae flattened, unequal in width, and overlapping adjacent costae; intercostal

striae narrow (only 8-18 % width of a costa); corallum roughly rectangular and

more highly compressed basally +... 5..--4.. 0225. S. imbricaticostatus sp. nov.

Sphenotrochus (S.) aurantiacus von Marenzeller, 1904

Fig. 6D-E, G-H

Sphenotrochus aurantiacus von Marenzeller, 1904: 280-281, pl. 18 (fig. 15). Wells, 1935: 531.

?Boshoff, 1981: 38-39. Cairns, 1989b: 38.

New records

AB-358A, 13, USNM 77185, 1, SAM-—H4589, 1, NM; AB-—358C, 2,

USNM 77182; MN-ZM8, 1, USNM 91719; 2 syntypes of Sphenotrochus aurantiacus

examined from Valdivia—104, ZMB 5095.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN DSS

Distribution

Agulhas Bank (Von Marenzeller 1904); off Durban and Zululand, South Africa;

?off south-eastern Mozambique (Boshoff 1981); 155-366 m. Bathymetric ranges of

Boshoff’s specimens not included because his specimens represent a mixed lot of two,

if not three, species.

Sphenotrochus (S.) evexicostatus sp. nov.

Fig. 8A-H

?Sphenotrochus intermedius: Macnae & Kalk, 1958: 123.

?Sphenotrochus sp. Pichon, 1974: 176, text-fig. 5.

Sphenotrochus aurantiacus: Boshoff, 1981: 38-39 [part.—3 specimens from off Inhaca, 12 m].

Records

Holotype: AB-372B, 1, USNM 77186. Paratypes: AB-371G, 3, USNM 77194;

AB-372B, 43, USNM 91712, 1, SAM-—H4584; AB-391J, 6, USNM 77184; AB-392J,

12, USNM 77188; off Inhaca, 12,2 m, 3, USNM 77190.

Description

Corallum medium in size for the genus, the holotype measuring 5,25 x 3,20 mm

in calicular diameter and 7,88 mm in height, but larger specimens up to 10,6 mm in

height present in type series. Corallum cuneiform, the flat corallum faces diverging at

a constant angle of about 12°. Viewed from the side, the corallum is triangular with a

gently rounded base. Costae equal in width (0,15—0,17 mm), rounded, and smooth,

only the principal Ci and their adjacent Cs being somewhat granular (Fig. 8H). Most

costae continuous from calice to about 1,5 mm from base, where they either terminate

or transform into discontinuous ridges. Costae usually not vertical in arrangement, but

slanted inward toward the sand-grain substrate incorporated into its base. Intercostal

striae broad and open, about 0,06—0,10 mm wide, or 35-55 per cent width of a costa,

affording an easy view of underlying corallum theca (Fig. 83D-E). Freshly preserved

coralla light yellow to brown.

Septa hexamerally arranged in three cycles. Pairs of Cs occur in each half-system

adjacent to the two principal septa (total = 32 costae), but rarely in half-systems

beyond these, but septa (Ss) do not correspond to these costae. S: and S2 equal in size,

moderately exsert, and extend about three-quarters distance to columella, merging

with columella only lower in fossa. Inner edges of S: and Sz slightly sinuous. S3 slightly

less exsert and only about one-third width of Si and S2. Septa and columella bear low,

inconspicuous granules, producing relatively smooth faces.

Columella a solid, sharp-edged lamella rising almost to height of exsert septa, and

of equal thickness to an S: (i.e. 0,15—0,16 mm).

Remarks

Although S. evexicostatus sp. nov. is of approximately the same size as S. imbri-

caticostatus sp. nov., has the same arrangement of septa and costae, similar columella,

and they are often found living together, S. evexicostatus is distinguished by its

rounded, non-imbricate costae of equal width, wider intercostal striae, smaller S3 in

relation to the S: and S2, and smoother septal and columellar faces. Sphenotrochus

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

cee ee

Fig. 7. A-B. Sphenotrochus gilchristi. A. S. dentosus (nomen nudum) of Boshoff (1981), South

Africa, South African Museum. B. Verma 19-28, USNM 87610, costal granulations near calice. X 24.

C. Tropidocyathus lessoni, MN—-ZH18, USNM 91698, lateral view showing alate end costae. x 2,5.

D-E, G-H. Sphenotrochus aurantiacus. D-E. Syntype from Valdivia—104, ZMB 5095, lateral and

calicular views. D x 6,6, E x 10. G-H. AB-358A, USNM 77185, lateral and calicular views. G x 6,2,

Hx 8,0. F, I. Thrypticotrochus multilobatus, AB-365D, USNM 91709, corallum fragment. x 10,5,

and enlargement of costae. Xx 35.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN Di).

Fig. 8. A-H. Sphenotrochus evexicostatus sp. nov. (B—C, holotype; A, D-H, paratype from

AB-392J). A. Lateral view of corallum. x 9,0. B. Stereo view of calice. x 11. C. Oblique view of

calice. 10. D-E. Longitudinal views of costae showing broad intercostal furrows and rounded

costae. D%*24, Ex71. F-G. Costal details at corallum base and near calice. FX 16s G X32-

H. Granulated edge costa. x 175.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

evexicostatus sp. nov. bears some resemblance to S. andrewianus Milne Edwards &

Haime, 1848 (eastern Atlantic, 12-105 m), in size and shape, but differs in having

some pairs of Cs, a more compressed corallum, and a consistently lamellar columella.

Etymology

The species name evexicostatus (from the Latin evexus, rounded at the top + cos-

tatus, having rib-like ridges) refers to the costae of this species, which are evenly

rounded on top.

Distribution

Off South Africa from Durban to south-eastern Mozambique; ?off Madagascar

(Pichon 1974); 12,2-73 m. Type locality: 24°48°S 34°59’E (off south-eastern Mozam-

bique), 42 m.

Sphenotrochus (S.) imbricaticostatus sp. nov.

Fig. 9A—H

Sphenotrochus aurantiacus: Boshoff, 1981: 38-39 [part.—9 specimens from off Inhaca, 12,2 m].

Records

Holotype: MN—ZB27, 1, USNM 91715. Paratypes: AB—370G, 12, USNM 77183;

AB-370H, 21, USNM 77181; AB-372B, 36, USNM 91716, 2, SAM—H4586;

AB-372L, 29, USNM 77187; AB-372P, 20, USNM 77189; AB-391J, 2,

USNM 91717; AB-392J, 7, USNM 91718; Inhaca, Mozambique, 12,2m, 9,

USNM 77190, and 24, currently deposited at Station Marine d’Endoume, Marseille.

Description

Corallum small, largest specimen (holotype) only 3,68 x 2,47 mm in calicular

diameter and 6,55 mm in height. Corallum cuneiform: highly compressed in lower

corallum where opposite faces are essentially parallel; upper corallum slightly flared.

Viewed from the side, corallum is often rectangular (Fig. 9A, C), with only a slight

rounding of the basal edges. Costae smooth, flat, and oriented vertically, most con-

tinuous from calice to base. In well-preserved coralla, 12-14 costae slightly overreach

the base and meet their counterparts from the opposite face. Costae alternate in width

near calicular edge, C3; (0,30-0,35 mm wide) being broader than the Ci and GQ

(0,14-0,18 mm wide, Fig. 9F—G). Intercostal striae narrow (0,030—0,035 mm wide),

or only 8-18 per cent of costal width, not affording a view of underlying theca. Each

costa has a thin (0,018—0,020 mm) lateral extension (Fig. 9G), which overlaps the

extension of its two adjacent costae in alternating fashion such that the Cs extensions

are invariably higher than those of the Ci and C2 near the calicular edge. Lower on

theca, the situation is reversed. These lateral costal extensions are particularly broad

(up to 0,1 mm) on the lower, outer edges of the four C2 near the principal C: and their

adjacent C; near the base of the corallum (Fig. 9H). Colour of a fresh corallum light

brown; theca procellaneous.

Septa hexamerally arranged in three complete cycles (24 septa). Pairs of Cs occur

in each half-system adjacent to the two principal S: (total = 32 costae), but S; do not

correspond to them inside calice. $i and S2 equal in size, moderately exsert, and

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 259

extend about four-fifths distance to columella, joining with columella only lower in

fossa. Inner edges of S: and &: slightly thickened. S3 equally exsert but only three-

quarters as wide as S: and S2. All septa, as well as columella, covered with tall (up to

0,060 mm), slender spines.

Columella a solid, sharp-edged lamella rising almost to height of exsert septa,

equal to or slightly thicker (about 0,1 mm) than Si and S2. Viewed from the side, the

columella is rectangular.

Remarks

Sphenotrochus imbricaticostatus sp. nov. is distinguished from the nine other

Recent species in the genus (eight listed by Cairns 19895: 38, plus S. evexicostatus sp.

nov.) by its extremely compressed rectangular—cuneiform corallum and its wide, flat

costae, which alternate in width and imbricate with edges of adjacent costae.

Etymology

The species name imbricaticostatus (from the Latin imbricatus, overlapping +

costatus, having rib-like ridges) is an allusion to the distinctive imbricate costae of this

species.

Distribution

Off Durban, South Africa to south-eastern Mozambique; 37-347 m. Type local-

ity: 27°03,9’S 32°53,0’E (off Zululand, South Africa), 44 m.

Sphenotrochus (Eusthenotrochus) gilchristi Gardiner, 1904

Fig. 7A-B

Sphenotrochus gilchristi Gardiner, 1904: 98-99, pl. 1 (figs a-g). Zibrowius & Gili, 1990: 44.

Eusthenotrochus moseri Wells, 1935: 530-532, pl. 18 (figs 5-6).

Sphenotrochus dentosus Boshoff, 1981: 39 (nomen nudum).

Sphenotrochus sp. (incertae sedis) Boshoff, 1981: 39 [= S. ‘dentosus’].

New records

AB-358A, 3, USNM 77191; AB-392J, 2, USNM 77192; Vema 19-28, 1,

USNM 87610; Sardinops CD32, 1, USNM 91714, 1, SAM—H4590.

Remarks

Three specimens identified as S. dentosus by Boshoff (1981) were borrowed from

the ORI and found to be typical S. gilchristi.

Distribution

Agulhas Bank (Wells 1935); off Durban and Simon’s Town; 24-165 m.

Genus Peponocyathus

Peponocyathus australiensis (Duncan, 1870)

Deltocyathus italicus var. australiensis Duncan, 1870: 297.

Deltocyathus minutus Gardiner & Waugh, 1938: 198, text-fig. 15. Scheer & Pillai, 1983: 156.

Peponocyathus australiensis: Cairns, 1989b: 30-32, pl. 14 (figs d—j), pl. 15 (figs a-d) [synonymy].

ANNALS OF THE SOUTH AFRICAN MUSEUM

iene Se

}

Fig. 9. A-H. Sphenotrochus imbricaticostatus sp. nov. (A-B, D-E, holotype; C, F—H, paratypes

from AB-372P). A, C. Lateral views. A X 8,3, C X11. B. Stereo view of calice at oblique angle,

x 14,5. D. Oblique lateral view of corallum, x 9,7. E. Calicular view, x 15,5. F-G. Imbricate

costae near calicular edge (exterior and cross sectional views). F x 41, Gx 110. H. Imbricate costae

near corallum base. x 60.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 261

New record

AB-3908S, 2, USNM 91711.

Distribution

Indian Ocean: off Natal, South Africa; off Zanzibar, Tanzania (Gardiner &

Waugh 1938); Red Sea (Gardiner & Waugh 1938); 101-366 m. Elsewhere: widely dis-

tributed in Atlantic and Indo-West Pacific; 44-635 m (Cairns 1989b).

Family Flabellidae

Genus Flabellum

Flabellum (Ulocyathus) japonicum bythios subsp. nov.

Fig. 1OA-B

Flabellum japonicum Wood Mason & Alcock, 1891a: 449-450. Alcock, 1898: 23. Gardiner, 1929:

306-308, pl. 13 (figs 1-6) [part.—not station 242].

Flabellum apertum: Keller, 1974: 205-208 [part.—V—4620, pl. 6 (figs 6—-7)].

Flabellum japonicum sensu Alcock, 1898: Cairns, 1989b: 54, 56.

Records

Holotype: V—2815, 1, IOM. Paratypes: V-—2653, 1, IOM; V—2814, 1, IOM;

V-—2815, 1, IOM; Marion Dufresne 27-4, 9, USNM 77203, 1, SAM-—H4575; Investi-

gator—133, 2, USNM 18156.

Description

Corallum elliptical in cross-section (GCD : LCD about 1,3), having a full, cam-

panulate shape, the 12 C: and C2 only slightly ridged at point of upward inflection of

corallum. Corallum fragile and thus often collected damaged. Slightly damaged holo-

type 46,7 x 36,0 mm in calicular diameter (estimated) and slightly over 25 mm in

height. Pedicel circular, about 2,5 mm in diameter. Freshly preserved coralla light

reddish-brown, sometimes more intensely pigmented near thecal edges of S: and S:.

Lower corallum—that part probably submerged in soft substrate—usually corroded

to a white or light grey colour.

Septa hexamerally arranged in five complete cycles; however, one large specimen

(V—2653) of 55 mm GCD has 14 primary septa and a corresponding number of higher

cycle septa for a total of 112 septa. S: and S2 equal in size, their straight inner edges

almost meeting in centre of fossa. S3 about half width of S: and S2; Ss about half width

of Ss; Ss rudimentary, extending only several millimetres down thecal wall and having

irregular, dentate inner edges. Calicular margins scalloped, each S: and S2 and adja-

cent Ss forming a triangular thecal extension about 3,5 mm tall, and each S;3 a smaller

extension of about 1,5 mm in height. Inner edges of all septa straight and the septa

themselves are planar; septal faces relatively smooth, covered with low, inconspicuous

granules. All septa quite thin (about 0,3 mm) and well separated from adjacent septa

by a distance of 4—5 septal thicknesses.

Fossa deep and elongate. Columella an elongate, concave fusion of lower, inner

edges of S:-S:.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Cairns (19896: 56) suggested that the Indian Ocean specimens previously

reported as F. japonicum (see synonymy) be considered as a new species, and listed

four differences between them and typical F. japonicum. To reiterate, the Indian

Ocean specimens have a campanulate corallum, higher thecal extensions, white

coralla, and lack thecal edge crests. These differences still appear valid except for

corallum colour, well-preserved Indian Ocean specimens having the same pigmen-

tation as F. japonicum. However, another difference noted here is that the pedicel of

F. japonicum bythios subsp. nov. is circular and larger than that of the typical sub-

species. Gardiner (1929) noted small differences between F. japonicum and Indian

Ocean specimens, but did not consider them to be of specific importance. Given the

geographic and bathymetric isolation of the two forms (see Distribution) and the

several minor but consistent morphological differences, the Indian Ocean specimens

are considered to represent a subspecies of F. japonicum.

Etymology

The subspecies name bythios (from the Greek bythios, of the deep) is an allusion

to the apparent deeper range of this subspecies than the typical subspecies.

Distribution

Off south-western Madagascar; Mascarene Plateau; Laccadive Sea (Gardiner

1929; Keller 1974); Gulf of Manaar (Gardiner 1929); Bay of Bengal (Alcock 1898;

Gardiner 1929); 1 095-1 720 m. Type locality: 9°40’S 60°31'E (Mascarene Plateau),

1 520-1 720 m. Distribution of typical subspecies: off Japan; Philippines; Indonesia;

128-1 141 m (Cairns 1989b).

Flabellum (Ulocyathus) lowekeyesi Squires & Ralph, 1965

Fig. 10D-E

Flabellum lowekeyesi Squires & Ralph, 1965: 259 (figs 1-2). Squires & Keyes, 1967: 27, pl. 6

(figs 1-2). Cairns, 1989b: 54.

New records

V-—2637, 1, IOM; V—2668, 11, IOM, 1, USNM 91734; V—2706, 4, IOM; V—2721,

95, IOM; V-—2764, 2, IOM; V—2816, 30, IOM.

Remarks

The south-west Indian Ocean specimens are indistinguishable from typical speci-

mens from off New Zealand.

Distribution

Indian Ocean: off south-eastern Mozambique; Mascarene Plateau; Madagascar

Plateau; 835-1 030m. Elsewhere: Campbell Plateau and off New Zealand;

278-732 m (Squires & Keyes 1967).

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 263

Flabellum (Ulocyathus) messum Alcock, 1902c

Fig. 10G-H

Flabellum lacianiatum var. messum Alcock, 1902c: 31.

Flabellum (U.) messum: Cairns, 1989b: 58-59, pl. 30 (figs f-i, k) [synonymy].

New record

V-—2816, 1, USNM 91735.

Remarks

Flabellum messum is very similar to F. lowekeyesi, but differs in having a reddish-

brown corallum; a rough, granular thecal texture; better defined thecal extensions;

and independent S: (see also Cairns 19895: 58).

Distribution

Indian Ocean: Mascarene Plateau; 430-835 m. Elsewhere: off Philippines (Cairns

1989b); Indonesia (Alcock 1902c); 368-949 m.

Flabellum (Flabellum) pavoninum Lesson, 1831

Flabellum pavoninum Lesson, 1831: 2. Gardiner, 1902: 123-125, pl. 4 (figs 18-21); 1904: 98. Gardiner

& Waugh, 1938: 174. Zibrowius et al., 1975: 98-99, pl. 2 (figs D-E). ?Boshoff, 1981: 35. Zibro-

wius & Grygier, 1985: 122. Cairns, 1989b: 46-50, pl. 23 (figs g—l), pl. 24 (figs a—d) [synonymy].

Flabellum pavoninum sensu Gardiner, 1902. Cairns, 1989b: 47, pl. 24 (figs g—h).

Flabellum sp. 6 Cairns, 1989a: 63, 67, text-fig. 2.

New records

V-2662, 1, IOM; V—2686, 1, IOM; V—2724, 16, IOM; V—2767, 8, IOM; V—2804,

3, IOM; AB-371E, 2, USNM 91724; AB-372L, 10, USNM 91726; AB-390S, 24,

USNM 91727; AB-421G, 1, USNM 91728; Manihine 381-63, 1, USNM 91722;

MN-ZD4, 1, USNM 91738; MN-ZCC1, 2, USNM 91732; MN-ZU13, 6,

USNM 91729; MN-ZU15, 11, USNM 91730, 1, SAM-H4582; MN-ZW6, 1,

USNM 91731. Reference specimens: Gardiner & Waugh’s (1938) specimens from:

JM-104, 1, BM 1950.1.9.1; JM-106, 3, BM 1950.1.9.11-35; JM-123, 1,

BM 1950.1.9.2—3; JM-—157, 2, 1950.1.9.36—40.

Remarks

Although I (Cairns 1989a, 1989b) previously distinguished the western Indian

Ocean specimens of Flabellum from F. pavoninum, based on discriminant analysis,

additional specimens of typical F. pavoninum recently examined from off Japan con-

vince us that these specimens are conspecific. Almost all characters analysed of

F. pavoninum and F. pavoninum sensu Gardiner (1902) in the discriminant analysis

(Cairns 1989a, table 5) were overlapping.

Distribution

Indian Ocean: off Durban (Gardiner 1902) to Zululand, South Africa; off Zanzi-

bar and Pemba, Tanzania (Gardiner & Waugh 1938); off Kenya (Cairns 1989b); off

western Madagascar (Zibrowius et al. 1975); Madagascar Plateau; Mascarene Plateau;

Arabian Sea (Gardiner & Waugh 1938); Maldive Islands (Gardiner & Waugh 1938);

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

98-665 m. Elsewhere: off Japan; South China Sea; Hawaii; 223-271 m (Cairns

1989b).

Genus Truncatoflabellum

KEY TO THE SOUTH-WEST INDIAN OCEAN SPECIES OF TRUNCATOFLABELLUM

1A. Corallum small (GCD < 13 mm); septa few (some Ss may be present, but never a

tullieycle yan Sx eae Tees Pole eee oe le oe oe Bee ee re 2

1B. Corallum medium- to large-sized, robust (adult GCD usually over 15 mm); septa

more numerous (usually > 80; often with Sc) ........-..22... 5) see 4

2A. Coralla with only three cycles of septa, often with some Su (i.e. 32 septa)........

a iM Aten ee eae ar eee em Oa elo Mere dh Mae T. pusillum Cairns, 1989b

2B. Coralla with four cycles of septa, often with some Ss (i.e. 56 septa)........... 8)

3A. Thecal edges carinate (not spinose); anthocyathus and anthocaulus invariably

SEM AT AGC ie ora ay tice ansines Ons. 3o. eis ab eam cle Metra ee T. gardineri sp. nov.

3B. Thecal edges rounded (also usually non-spinose); anthocyathus and anthocaulus

oitenmemiamprattache den rcs Cains a natn ay giclee arenas T. zuluense sp. nov.

4° Thecal edges rounded, no thecal spines). .......5. 2.2... J... ere 5)

4B. Thecal edges not rounded: either spinose or carmate . >. ..-.. =. ose eee 6

5A. Thecal edge angle large (about 68); anthocyathus basal scar small (3,5 xX 2,5 mm)

PS area Po gee eh cet ler Aon A nee i crane T. stabile (von Marenzeller, 1904)

5B. Thecal edge angle less than 60; anthocyathus scar larger, 5,1-6,7 x 4,0—4,8 mm

A MMT h/t get eo aks aah NN T. inconstans von Marenzeller, 1904

6A. Thecal edges of anthocyathus spinose, with up to 7 pairs of spines...............

Ree ed Naya crer east Banas Oe ce ois Pe eta a RARE Ee trae T. multispinosum sp. nov.

6B. Thecal edges of anthocyathus acute, often carinate, having no spines ............

Bea aes ay staf eRe he ah) Aesth anes Pr pa T. formosum Cairns, 19896

Truncatoflabellum sp. cf. T. stabile (von Marenzeller, 1904)

Fig. 10C, F

New record

V-2629, 1, IOM.

Diagnosis

A unique specimen measures 47,6 X 25,8 mm in calicular diameter and 61,3 mm

tall. Angle of thecal edges a constant 68°; inclination of thecal faces 32°. Theca worn,

although septa are freshly preserved. Thecal edges rounded; no thecal edge spines or

crests present. Basal scar small, only 3,5 x 2,5 mm in diameter. Corallum dense and

white, with an evenly arched upper calice. Septa hexamerally arranged according to

the formula: Si-S3>Ss>Ss. Four pairs of Ss occur in four random half-systems, for a

total of 104 septa. Inner septal edges straight; septa concave near junction with calice.

Fossa quite deep, containing a rudimentary columella composed of lower, inner edges

of Si-Ss.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 265

Remarks

This specimen corresponds to the description and illustrations of Truncatoflabel-

lum stabile (von Marenzeller, 1904: 273-274, pl. 17 (fig. 12)); however, no com-

parative specimens have been examined of this species, since the type apparently is

missing (Zibrowius 1980). Although similar to the Truncatoflabellum reported by

Zibrowius & Gili (1990) from Walvis Ridge in shape and in having a small basal scar,

it differs in having five cycles of septa, not four. It is most similar to 7. inconstans,

especially in corallum size, number of septa, and in lacking thecal edge spines, but

appears to differ in having a higher thecal edge angle (and thus a much more open

calice) and a smaller anthocyathus basal scar (see Key).

Distribution

Off south-eastern Mozambique; 1 250-1 520 m. Distribution of T. stabile: Cape

Verde, Selvagems, and Madeira islands; 1 450-3 010 m (Zibrowius & Gili 1990).

Truncatoflabellum formosum Cairns, 1989b

Fig. 101, 11A

Truncatoflabellum formosum Cairns, 1989b: 69-70, pl. 35 (figs j-k), pl. 36 (figs a—b) [synonymy].

New records

V-2635, 2, IOM, 1, USNM 91757; MN—ZCC2, 1, USNM 91760.

Remarks

The Indian Ocean specimens were compared to the types of 7. formosum and

found to be indistinguishable. The illustrated specimen measures 20,0 X 13,6 mm in

calicular diameter and 23,7 mm in height.

Distribution

Indian Ocean: off Zululand, South Africa; off south-eastern Mozambique;

150-230 m. Elsewhere: Korea Strait; Philippines; Celebes; 37-933 m (Cairns 1989b).

Truncatoflabellum pusillum Cairns, 1989b

Fig. 11E

Truncatoflabellum pusillum Cairns, 1989b: 71-72, pl. 37 (figs a-e).

New records

AB-371E, 1, USNM 91755; AB-371F, 35, USNM 91756, 1, SAM—H4587.

Remarks

The specimens from Mozambique agree in all respects with the type specimens of

T. pusillum from the Philippines, the largest anthocyathus measuring 8,39 x 6,22 mm

in calicular diameter and 12,2 mm in height.

Distribution

Indian Ocean: off south-eastern Mozambique; 110-132 m. Elsewhere: Philip-

pines; 143-146 m (Cairns 1989b).

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Truncatoflabellum gardineri sp. nov.

Fig. 11B-—D

Records

Holotype: AB-390S, 1, USNM 91736. Paratypes: AB—390S, 132, USNM 91737,

3, SAM-—H4577.

Description

Lower half of anthocyathi compressed, their lateral edges carinate in a series

of low, thin, discontinuous ridges projecting outward up to 1,3 mm; upper half of

anthocyathi less compressed (GCD : LCD = 1,30-1,47), their edges evenly

rounded and essentially parallel. Inclination of convex thecal faces 14-18°. Coralla

small and elongate: holotype (an anthocyathus) 10,9 x 8,3 mm in calicular diameter

and 17,4 mm in height, with a basal scar diameter of 5,5 x 3,2 mm. Largest antho-

cyathus 12,7 mm in GCD and 18,7 mm in height. Basal anthocyathus scar elliptical:

4,1-5,3 X 2,8-3,3 mm in diameter. Corallum white. Anthocauli also have carinate

edges, a pedicel diameter of 2,1-2,5 mm, four cycles of septa, and rarely exceed

10 mm in height.

Septa hexamerally arranged in four complete cycles (Si-S2>S3> S.), only the

largest specimens having four pairs of rudimentary Ss in the end quarter-systems

(56 total septa). Inner edges of Si and S2 vertical and highly sinuous, their lower, inner

edges fusing deep in fossa to form a rudimentary columella. S3 half to three-quarters

width of Si and S2 and less sinuous; Ss much smaller than S3, with dentate to laciniate

inner edges. All septal faces covered with prominent granules. Fossa deep and

elongate.

Remarks

T. P. Lowe observed living specimens of this species shortly after capture and

observed that each anthocyathus had two tentacular rings, the outer composed of

small, white tentacles, the inner composed of larger tentacles with white tips and

blood-red bases. Furthermore, he noted six orange stripes on the calice extending

from the calicular edge to the outer tentacular ring.

Of the 28 species of Truncatoflabellum listed by Cairns (1989a), almost all have

one or more pairs of thecal edge spines on their anthocyathi. In the few non-spinose

species, the lateral edges are either rounded or sharp-edged. Truncatoflabellum gardi-

neri sp. nov. is therefore unique in having non-spinose, carinate thecal edges. It is

further distinguished by its relatively small size, parallel thecal edges, and low number

of septa. It is similar to 7. pusillum in corallum and scar size, but can be distinguished

by its greater number of septa (48 vs 32) and in having non-spinose, parallel thecal

edges.

Etymology

Named in honour of John Stanley Gardiner, who published significant papers on

the deep-water corals of the south-west Indian Ocean (see Introduction and Table 1).

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 267

Distribution

Known only from the type locality: 29°34’S 31°42’E (off Durban, South Africa),

138 m.

Truncatoflabellum zuluense sp. nov.

Fig. 11F-G

Records

Holotype: MN—ZK21, 1, USNM 91747. Paratypes: MN—ZC9, 2, USNM 91748,

1, SAM-H4581; MN-ZC10, 2, USNM 91749; MN-ZC11, 1, USNM 91750;

MN-ZDD2, 1, USNM 91753; MN-ZDD3, 1, USNM 91754; MN-ZK20, 2,

USNM 91751; MN-—ZK21, 2, USNM 91752.

Description

In all 13 specimens examined, the anthocyathi and anthocauli remained together

as one corallum, but a difference in corallum colour and an incipient fracture line

usually indicated where the separation would occur. Corallum compressed

(GCD : LCD = 1,4~-1,65-1,8), with rounded thecal edges diverging at a constant

angle of 28-—38°. Inclination of thecal faces 18-—22°. Coralla small: holotype

13,2 x 7,9 mm in calicular diameter and 17,2 mm in height, with a pedicel diameter of

1,9 mm. Lower 6 mm of corallum (presumed anthocaulus) white and usually non-

spinose but sometimes bearing one pair of short thecal edge spines. At corallum

height of about 6 mm and a calicular diameter of about 6,0-6,5 x 4,5 mm, the coral-

lum bears a transverse crease, which is presumed to be the incipient fracture line,

above which the corallum (anthocyathus) is reddish-brown. Anthocyathi also usually

non-spinose but occasionally bear one pair of short thecal edge spines basally.

Septa hexamerally arranged in four cycles (Si-S2>S3>>Su,), larger specimens also

having pairs of Ss in the end half-systems (total of 56 septa). Inner edges of Si and Sz

vertical and sinuous, defining a deep and narrow fossa. S3 about two-thirds width of

larger septa; S1 much smaller, usually with finely dentate inner edges. All septa

covered with tall granules.

Remarks

Truncatoflabellum zuluense sp. nov. is distinguished from most other species in

the genus by its small size and by having only 48-56 septa; rarely having thecal spines;

and in maintaining the anthocaulus—anthocyathus connection long into ontogeny.

Truncatoflabellum zuluense is most similar to T. gardineri sp. nov., another relatively

shallow-water, south-west Indian Ocean species, especially in corallum size and septal

number, but can be distinguished by its rounded, sometimes spinose, divergent thecal

edges (not carinate and parallel); retention of the anthocaulus stage; and reddish-

brown corallum colour.

Etymology

The species name zuluense is an allusion to the area of capture of the type series:

off Zululand, South Africa.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Known only from off Zululand, South Africa; 62-84 m. Type locality: 27°47,6'S

32°39,1’E, 62-84 m.

Truncatoflabellum multispinosum sp. nov.

Figs 11H, 12A—C

Records

Holotype: V—2634, 1, USNM 91739. Paratypes: AB—372L, 55, USNM 91741, 1,

SAM-H4580; AB—409E, 2, USNM 91742; MN-ZB11, 1, USNM 91743; MN-—ZB17,

1, USNM 91744; MN-ZDD4, 1, USNM 91746; MN-ZH18, 2, USNM 91745;

CH11-329-10, 2, USNM 91740.

Description

Angle of acute thecal edges 41—-56°; inclination of convex thecal faces 19—32°.

Largest anthocyathus (holotype) 32,2 x 15,7 mm in calicular diameter and 28,1 mm in

height, with a basal scar diameter of 7,3 x 4,8 mm. Up to seven pairs of slender spines

project from thecal edges, usually regularly occurring from scar to calice, the num-

ber determined by size of corallum. Basal scar of anthocyathus elliptical:

5,6-7,3 X 4,2—4,8 mm in diameter. Calice elliptical, GCD : LCD = 1,67-1,83-2,05.

Theca of anthocyathus reddish-brown, often more intensely striped vertically; septa of

anthocyathus and entire anthocaulus white. Anthocauli rarely greater than 7,5 mm in

height, having the calicular diameter of the anthocyathus basal scar. Anthocauli have

a pedicel diameter of 1,8—2,0 mm, 24 septa, and one pair of thecal edge spines.

Septa of large coralla (e.g. over 23 mm GCD) hexamerally arranged in five cycles

according to the formula: Si—S3>Ss>Ss, often with four pairs of Ss in the four quarter-

systems adjacent to the two principal Si; however, the large holotype lacks two pairs

of Ss in lateral half-systems, resulting in 100 septa. In medium-sized anthocyathi (e.g.

GCD = 10-22 mm), symmetry appears to be decameral, coralla often having the

formula: 20 : 20 : 40 (80 septa), with irregular development of tertiary septa in lateral

half-systems and accelerated development in the end half-systems. S:i—S3 have finely

sinuous, vertical inner edges, which define a narrow, elongate fossa. Ss only slightly

smaller, their inner edges somtimes also reaching the columella. Ss about one-third

width of Ss; Ss rudimentary. Columella a trabecular fusion of lower, inner edges of

Si—Ss and occasionally the Ss.

Remarks

Among the 30 described species of Truncatoflabellum (see Cairns 1989b), T. mul-

tispinosum sp. nov. most closely resembles T. candeanum (Milne Edwards & Haime,

1848), particularly in thecal colour, number of septa, and in having relatively widely

divergent thecal edges. Truncatoflabellum multispinosum sp. nov. differs consistently,

however, in having a smooth calicular edge (that of T. candeanum is scalloped); larger

Sin relation to the S:-Ss; a larger anthocyathus scar; and slightly less divergent thecal

edges and faces. Truncatoflabellum vanuatu (Wells, 1984) is the only species known to

have up to five pairs of thecal spines, but differs in having a much smaller anthocyathus

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 269

Fig. 10. A-B. Flabellum japonicum bythios subsp. nov., V-2815, IOM, basal and calicular views.

Both *1,0. C, F. Truncatoflabellum sp. cf. T. stabile, V—2629, IOM, lateral and calicular views.

Cx0,8, FX1,1. D-E. Flabellum lowekeyesi, V-2668, USNM 91734, lateral and calicular views.

Dx1,2, Ex1,7. G-H. Flabellum messum, V-2816, USNM 91735, lateral and calicular views.

G 1,0, H 1,3. I. Truncatoflabellum formosum, V-2635, USNM 91757, lateral view. x 1,8.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. A. Truncatofiabellum formosum, V—2635, USNM 91757, calicular view. x 2,4. B—D. Trun-

catoflabellum gardineri sp. nov. from AB-390S. B-C. Calicular and lateral views of holotype.

B x 3,9, CX 2,9. D. Lateral view of two paratypes. x 2,2. E. Truncatoflabellum pusillum, AB-371F,

USNM 91756, lateral view of anthocyathus and anthocaulus. x 5,2. F—G. Truncatoflabellum zuluense

sp. nov., MN—ZK21 (holotype), lateral and calicular views. F x 2,9, G x 3,8. H. Truncatoflabellum

multispinosum sp. nov., aberrant paratype from AB-372L. x 2,2.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN Deal

Wi lac

Fig. 12. A-C. Truncatoflabellum multispinosum sp. nov. A. Lateral view of paratype from AB-372L

(USNM 91741) showing seven pairs of lateral spines. X 1,7. B-C. V-2634, lateral and calicular

views of holotype. B x 1,2,C x 1,5. D, G. Placotrochides scaphula, AB-389C, USNM 91772, lateral

and calicular views. Dx8,1, Gx8,4. E-F. Stenocyathus vermiformis, V-2722, USNM 91775,

lateral and calicular views. E x 3,7, FX 7,4. H-I. Guynia annulata, AB-390S, USNM 77201, lateral

and calicular views. H x 17, I x 23.

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

scar; having a much narrower anthocyathus (edge angle only 20—27°); and in having

fewer septa.

Etymology

The species name multispinosum (from the Latin multus, more + spina, spine) is

an allusion to the numerous thecal edge spines characteristic of this species.

Distribution

Off Zululand, South Africa; off south-eastern Mozambique; Mozambique

Channel off western Madagascar; off Zanzibar, Tanzania; 62-183 m. Type locality:

25°05'S 34°50’E (south-eastern Mozambique), 90-92 m.

Genus Javania

Javania insignis Duncan, 1876

Javania insignis Duncan, 1876: 435, pl. 39 (figs 11-13). Von Marenzeller, 1907b: 23, pl. 2 (fig. 6).

Zibrowius, 1974b: 8-9, pl. 1 (figs 1-6). Fricke & Schuhmacher, 1983: 184. Scheer & Pillai, 1983:

165-166, text-fig. 4, pl. 37 (figs 9-12). Cairns, 1989b: 77-78, pl. 40 (figs d—e, h, j, k) [synonymy].

Flabellum weberi Alcock, 1902a: 107. Cairns, 1989b: 77. [syn. nov.]

New records

AB-371F, 1, SAM-—H4591; MN-ZC11, 1, USNM 91768; MN-ZDD3, 2,

USNM 91770; MN-—ZQ8a, 1, USNM 91769. Reference specimen: holotype of Flabel-

lum weberi, Siboga—310, ZMA 1232.

Distribution

Indian Ocean: off Natal and Zululand, South Africa; off south-eastern Mozam-

bique (Cairns, 1989b); off north-western Madagascar (Zibrowius 1974b); Red Sea

(Von Marenzeller 1907b; Fricke & Schuhmacher 1983; Scheer & Pillai 1983);

74-255 m. Elsewhere: off Japan; Philippines; Indonesia; Hawaiian Islands; 46-825 m

(Cairns 1989b).

Genus Placotrochides

Placotrochides scaphula Alcock, 1902b

Fig. 12D5G

Placotrochides scaphula Alcock, 1902b: 121-122; 1902c: 34, pl. 4 (fig. 32, 32a). Cairns, 1989b: 78-79,

pl. 40 (fig. 1), pl. 41 (figs a—e) [synonymy].

New records

AB-365D, 1, USNM 91771; AB-389C, 4, USNM 91772; AB-389E, 1,

USNM 91773.

Remarks

The south-west Indian Ocean specimens are similar to the holotype of P. sca-

phula from Indonesia, more so than those specimens reported by Cairns (1989b) from

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 273

the Philippines, which are considerably larger and have more septa. The figured speci-

men measures 6,43 X 4,97 mm in calicular diameter and has 18 major septa.

Distribution

Indian Ocean: off Natal, South Africa; off south-western Madagascar;

475-1 360 m. Elsewhere: Philippines; Indonesia; 462-1 628 m (Cairns 19895).

Family Guyniidae

Genus Guynia

Guynia annulata Duncan, 1872

Fig. 12H-I

Guynia annulata Duncan, 1872: 32, pl. 1 (figs 1-8). Cairns, 1984: 23, pl. 5 (figs A-B); 1989b: 42,

pl. 21 (fig. f), pl. 22 (figs a—e) [synonymy]. Cairns & Parker, 1992: 42-43, pl. 14 (figs g—h).

Pyrophyllia inflata Hickson, 1910: 1-7, text-figs 1-4; 1911: 1 038-1 042. Harrison, 1911: 1 020, pl. 57

(figs 8-11), pl. 58 (figs 18-19).

New records

AB-372L, 3, USNM 77202; AB-390S, 2, USNM 77201.

Distribution

Indian Ocean: off Durban, South Africa; off south-eastern Mozambique; Gulf of

Oman (Hickson 1910, 1911; Harrison 1911); 112-286 m. Elsewhere: probably cosmo-

politan—amphi-Atlantic; Indonesia; Philippines; New Caledonia; Hawaiian Islands;

southern Australia; 28-653 m (Cairns & Parker 1992).

Genus Stenocyathus

Stenocyathus vermiformis (Pourtalés, 1868)

Fig. 12E-F

Coenocyathus vermiformis Pourtalés, 1868: 133.

Stenocyathus vermiformis: von Marenzeller, 1904: 298-300, pl. 18 (fig. 16). Zibrowius, 1974a:

769-770. Cairns, 1979: 168-170, pl. 32 (figs 8-10), pl. 33 (figs 1-2) [synonymy]; 1982: 52, pl. 16

(figs 8-11); 1984: 23-25, pl. 5 (fig. c). Cairns & Parker, 1992: 43-44, pl. 14 (figs b-c).

New records

V-2722, 1, USNM 91775; V—2723, 1, IOM.

Distribution

Indian Ocean: Madagascar Plateau near Walters Shoal; St Paul and Amsterdam

Islands (Von Marenzeller 1904; Zibrowius 1974a); 80-672 m. Elsewhere: amphi-

Atlantic, off New Zealand; Tasmania; Great Barrier Reef; 80-1 229 m (Cairns &

Parker 1992).

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Suborder DENDROPHYLLIINA

Family Dendrophylliidae

Genus Balanophyllia

Balanophyllia stimpsonii (Verrill, 1865)

Eupsammia stimpsonii Verrill, 1865: 150.

Rhodopsammia socialis: Alcock, 1893: 147.

Balanophyllia socialis: Bourne, 1905: 210, pl. 2 (fig. 8, 8a). Harrison & Poole, 1909: 902.

Balanophyllia imperialis: van der Horst, 1926: 48.

Balanophyllia affinis: van der Horst, 1931: 10. Gardiner & Waugh, 1939: 240.

Balanophyllia cumingii: Gardiner & Waugh, 1939: 238, pl. 1 (fig. 1).

Balanophyllia stimpsonii: Zibrowius, 1985: 234-235 [synonymy]. Zibrowius & Grygier, 1985: 126-127,

figs 27-29.

New records

V-2634, 1, IOM; AB-372L, 2, USNM 77228.

Distribution

A common, free-living, shallow-water Indo-West Pacific species. Indian Ocean:

False Bay and off Natal, South Africa (Zibrowius 1985); off Mozambique; off Somalia

(Zibrowius & Grygier 1985); Seychelles and Reunion (Zibrowius 1985); Gulf of Oman

(Zibrowius & Grygier 1985); off Sri Lanka (Van der Horst 1926; Gardiner & Waugh

1939); Gulf of Manaar (Bourne 1905); Maldive Islands (Gardiner & Waugh 1939);

Andaman Sea (Alcock 1893); Mergui Archipelago (Harrison & Poole 1909);

11-112 m. Elsewhere: Philippines; Indonesia; Chesterfield Islands; eastern Australia

(Zibrowius 1985).

Balanophyllia ponderosa van der Horst, 1926

?Eupsammia regalis Alcock, 1893: 144-145, pl. 5 (fig. 8, 8a).

Balanophyllia ponderosa van der Horst, 1926: 49-50, pl. 3 (figs 6-7). Eguchi, 1968: C54 [part.—

pl. C17 (figs 6-11, 13-14)]. ?Boshoff, 1981: 41.

New record

MN-ZA48, 1, USNM 91777. Reference specimen: holotype of B. ponderosa

(BM 1939.7.20.62).

Remarks

This specimen, measuring 20,4 x 15,8 mm in calicular diameter and 26,7 mm in

height, is identical to the holotype of B. ponderosa van der Horst (1926), except that

the specimen lacks epitheca and is unattached, perhaps both the result of damage

early in ontogeny. In this regard, however, it is very similar to the original illustration

of B. regalis (Alcock, 1893), which may prove to be the senior synonym if the type of

B. regalis can be obtained.

Distribution

Indian Ocean: ?off Durban (Boshoff 1981); off Zululand, South Africa; Sey-

chelles (Van der Horst 1926); Maldive Islands (Van der Horst 1926); ?off Sri Lanka

(Alcock 1893); 51-59 m. Elsewhere: off Japan; 10-250 m (Eguchi 1968).

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 275

Balanophyllia diffusa Harrison & Poole, 1909

Fig. 13A, D

Balanophyllia diffusa Harrison & Poole, 1909: 906, pl. 85 (fig. 4a, b). Gardiner & Waugh, 1939:

239-240. ?Pillai & Scheer, 1976: 16.

Non Balanophyllia diffusa Harrison & Poole. Scheer & Pillai, 1983: 168.

New records

AB-372L, 2, USNM 91781; AB-390S, 1, USNM 91782; EAMFRO-C13, 2,

USNM 78637; Cr. 329, Sta. 15, 2, USNM 78594 and 78639; MN-ZC10, 1,

USNM 91780. Reference specimens: B. diffusa of Gardiner & Waugh (1939):

JM-112, 4, BM 1939.7.13.67; JM-157, 10, BM 1950.1.6.35.

Remarks

Eight specimens appear to be conspecific with B. diffusa, based on the original

description and figures of this species. The quasi-colonial nature of the B. diffusa

reported by Gardiner & Waugh (1939) from Tanzania, Red Sea, and Maldive Islands,

is simply due to specimens settling close to one another.

Distribution

Off Durban and Zululand, South Africa; off south-eastern Mozambique; off

Pemba (Gardiner & Waugh 1939) and Zanzibar, Tanzania; north Kenya Banks;

Mergui Archipelago (Harrison & Poole 1909); Maldive Islands (Gardiner & Waugh

1939; Pillai & Scheer 1976); 6-274 m.

Genus Trochopsammia

Trochopsammia togata (van der Horst, 1927) comb. nov.

Balanophyllia togata van der Horst, 1927: 5—6, pl. 2 (figs 10-11), text-fig. 3.

Thecopsammia togata: Wells, 1935: 531.

New records

AB-358A, 1, USNM 77237; MN-SM226, 1, USNM 91792, 1, SAM-—H4592;

MN-SM232, 4, USNM 91791.

Remarks

Trochopsammia togata cannot be placed in the genus Balanophyllia, because its

septa are not arranged in a Pourtalés Plan. For the same reason, it cannot be placed in

Thecopsammia, even though Vaughan & Wells (1943) and Wells (1956) incorrectly

assumed that Thecopsammia socialis (type species of that genus) has normally

arranged septa (see Cairns 1979). Trochopsammia togata is more similar to Trochop-

sammia, which is known from only the type species, T. infundibulum Pourtales, 1878,

differing primarily in not having well-defined costae. Trochopsammia togata can be

diagnosed as having a slender, epithecate elongate corallum; fewer than four cycles of

normally arranged septa, the four lateral S: being wider than the other two; and

lacking a columella.

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Agulhas Bank (Wells 1935) to Durban, South Africa (Van der Horst 1927);

155-775 m.

Genus Endopachys

Endopachys grayi Milne Edwards & Haime, 1848b

Endopachys grayi Milne Edwards & Haime, 1848b: 82-83, pl. 1 (fig. 2). Wan der Horst, 1926: 51;

1927: 6-7, pl. 2 (fig. 12). Gardiner & Waugh, 1939: 241. Boshoff, 1981: 42 [part.]. Cairns, 1984:

27, pl. 5 (fig. E) [synonymy]. Zibrowius & Grygier, 1985: 128, figs 39-42. Cairns, 1989b: 34.

Endopachys weberi Alcock, 1902a: 109-110 [syn. nov.].

New records

V-2634, 12, IOM; V—2809, 6, IOM, 1, USNM 91811; AB-371F, 1, USNM

77247; AB-372J, 3, USNM 77251; AB-372L, 82, USNM 77245; AB-390S, 15,

USNM 77248, 2, SAM-—H4593, 2, NM; AB-391H, 1, USNM 77252; AB-391J, 3,

USNM 77246; Vema 14-SAT6, 5, USNM 77254; MN-ZDD2, 1, USNM 91813;

MN-ZWS8, 3, USNM 91812.

Remarks

The specimen from V—2809, measuring 38,9 x 32,7 mm in calicular diameter and

34,0 mm in height, is believed to be the largest recorded, yet only slightly larger than

the holotype. It nonetheless has only five cycles of septa (96).

Distribution

Indian Ocean: off South Africa from off Bisho (Van der Horst 1927) to Zululand

(Van der Horst 1927; Zibrowius & Grygier 1985); off south-eastern Mozambique; off

Zanzibar, Tanzania (Cairns 1989b); off Mauritius (Van der Horst 1926) and Saya de

Malha; north-eastern Arabian Sea (Gardiner & Waugh 1939); 57-274 m. Elsewhere:

Western Pacific; Hawaiian Islands; Gulf of California; 37-274 m (Cairns 1984).

Genus Rhizopsammia

Rhizopsammia annae (van der Horst, 1933)

Fig. 13C

Balanophyllia annae van der Horst, 1933: 156-158, pl. 7 (figs 14); 1938, pl. 5 (fig. 1). Boshoff, 1981:

40.

Rhizopsammia annae: Zibrowius & Gili, 1990: 44.

New record

AB-391J, 1 colony, USNM 91790.

Distribution

Known only from off South Africa from the Cape of Good Hope (Van der Horst

1933) to Port Shepstone (Boshoff 1981); 0-80 m.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN PHT

Rhizopsammia compacta Sheppard & Sheppard, 1991

Fig. 13B, E

Rhizopsammia compacta Sheppard & Sheppard, 1991: 153, fig. 179.

New records

AB-357B, 2, USNM 91794; AB-371F, 6, USNM 91795; MN-ZA43, 1,

USNM 91796; MN-ZA48, 3, USNM 91797, 1, SAM-—H4578; MN-ZA49, colony of

9 corallites, USNM 91793; MN—ZB11, 1, USNM 91798; MN-ZB14, 1, USNM 91799;

MN-ZB17, 1, USNM 91800; MN-ZB18, 1, USNM 91801; MN-ZB20, 2,

USNM 91802; MN-—ZB23, 1, USNM 91803; MN-ZB25, 1, USNM 91804; MN-ZDS,

1, USNM 91805; MN-ZD7, 4, USNM 91806; MN-ZD10, 2, USNM 91807;

MN-ZDD2, 2, USNM 91808; MN-ZDD3, 3, USNM 91809, 2, SAM-H4579;

MN-ZDDS, 1, USNM 91810.

Redescription

Colonies irregularly shaped, composed of clumps of corallites interconnected by

narrow stolons or consisting of small corallites budded from lower edge zone of larger

corallites. Individual corallites broken from the substrate are indistinguishable from

Balanophyllia. Corallites cylindrical to ceratoid in shape, the calice usually circular in

young corallites but often becoming highly compressed in larger corallites, with a

GCD : LCD up to 3,0. Largest colony examined (MN-—ZA49) composed of 9 coral-

lites, the largest 21,1 x 7,3 mm in calicular diameter and 27,2 mm in height, with

slightly concave thecal faces. Costae equal in width, covered with fine spines; no epi-

theca. Corallum white, but often encrusted basally with red foraminifera.

Septa hexamerally arranged in five cycles (96 septa), only the largest corallites

having pairs of Se, e.g. the large corallite from MN-—ZA49 has 8 pairs of Se in three of

its half-systems for a total of 112 septa. Si-S3 equal in size, only slightly exsert, and

have vertical inner edges descending into a deep fossa. S:—Ss slightly less exsert than

S:-Ss, the Ss being the narrowest of the septa, flanked by pairs of Ss, which are slightly

less wide than Si—S; in the upper fossa, but equal in width to S:—Ss in lower fossa,

where each pair of Ss joins before its enclosed Ss. Septa closely spaced, giving a

crowded aspect; septal faces covered by tall, robust granules. All inner septal edges

straight: entire in upper fossa, finely dentate in lower fossa. Paliform lobes absent.

Fossa contains an elongate, spongy, flat columella that fuses to lower, inner edges

of S:-S:, and Ss.

Remarks

This recently described species is redescribed herein, the original account being

based on only one specimen. The equally wide, vertically-edged Si—S; and Ss of

R. compacta produce a well-delineated fossa, which is characteristic of the species and

helps to distinguish it from all others. Furthermore, it differs from the 11 other valid

species in the genus (see Wells (1982) for listing of 8 species; plus R. annae (van der

Horst, 1933); R. wettsteini Scheer & Pillai, 1983; and R. eguchi (Wells, 1982)) by

having relatively large corallites with compressed calices; five full cycles of septa;

S:i—Ss equal in size; and no pali. It is perhaps most similar to R. manuelensis Cheva-

lier, 1966, particularly in corallum shape and corallite size, but R. manuelensis differs

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

in having paliform lobes; exsert septa; one less cycle of septa; and a discrete colu-

mella. Rhizopsammia compacta differs from R. wettsteini, known from the Red Sea,

in having larger, compressed corallites and in having equally-sized S:—Ss.

Distribution

Known only from the Indian Ocean off Durban to south-eastern Mozambique

(43-110 m) and the Gulf of Oman (35 m). Type locality: off Musandam, Gulf of

Oman, 35 m.

Genus Dendrophyllia

Dendrophyllia sp. cf. D. horsti Gardiner & Waugh, 1939

Fig. 13F, I

?Dendrophyllia arbuscula van der Horst, 1922: 53, pl. 8 (fig. 6).

?Dendrophyllia horsti Gardiner & Waugh, 1939: 237-238, pl. 2 (figs 5-6). Pillai & Scheer, 1976: 16.

Fricke & Schuhmacher, 1983: 184, fig. 14d. Scheer & Pillai, 1983: 171-172, pl. 39 (figs 11-12).

Dendrophyllia coccinea: van der Horst, 1926: 45-46, pl. 3 (figs 1-3). Gardiner & Waugh, 1939: 236.

New records

AB-372G, 1 colony, USNM 91814; MN-XX153, 1 colony, USNM 91815; MN-

ZB11, 3 colonies, USNM 91816; MN-ZB18, 1 colony, USNM 91817; MN-ZB20,

2 colonies, USNM 91818; MN-—ZC11, 1 colony, USNM 91819; MN-ZD6, 1 colony,

USNM 91820. Reference specimens of Gardiner & Waugh’s (1939) D. coccinea:

JM-112, 1 colony, BM 1939.7.13.83.

Remarks

The south-west Indian Ocean specimens may be characterized as small irregular

colonies, usually less than 5 cm in height, composed of 2—5 thick (about 12 mm in

diameter), cylindrical corallites budded at angles of 45—80° from the parent corallite.

Septa arranged in four cycles (Si>S2>Ss>Ss), with occasional Ss; fossa deep; colum-

ella discrete, convex. Because the types of D. horsti were not examined, the identi-

fication of these specimens must remain tentative. Dendrophyllia arbuscula may well

be a larger colony of the same species.

Distribution

Off South Africa from Natal to Zululand; off Pemba, Tanzania (Gardiner &

Waugh 1939); 50-113 m. Distribution of D. horsti: Red Sea (Fricke & Schuhmacher

1983; Scheer & Pillai 1983); Maldive Islands (Gardiner & Waugh 1939; Pillai &

Scheer 1976); ?Indonesia (Van der Horst 1922); 45-229 m.

Dendrophyllia dilatata van der Horst, 1927

Dendrophyllia dilatata van der Horst, 1927: 2-3, pl. 1 (figs 2-4), text-fig. 1. Zibrowius & Gili, 1990:

44.

New records

AB-371E, 4 colonies, USNM 71865; AB-371F, 8 colonies, USNM 71866,

1 colony, SAM-—H4571.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 279

Remarks

Dendrophyllia dilatata is similar to the previous species in corallum size, septal

arrangement, and geographic distribution, but can be distinguished by having a

slightly larger (up to 7 cm), usually arborescent colony composed of up to a dozen

sympodially arranged corallites; only four cycles of septa; smaller, ceratoid corallites;

and a shallower fossa.

Distribution

Off Durban, South Africa (Van der Horst 1927) to off south-eastern Mozam-

bique; 97-132 m.

Dendrophyllia cladonia van der Horst, 1927

Dendrophyllia cladonia van der Horst, 1927: 3-4, pl. 1 (figs 5-6), pl. 2 (fig. 7), text-fig. 2. Zibrowius

& Gili, 1990: 44.

New records

AB-357B, 2 corallites, USNM 91826; AB-357E, 7 corallites, USNM 91825;

AB-371F, 3 corallites, USNM 91824; AB-390S, 48 colonies or isolated corallites,

USNM 91823.

Remarks

Dendrophyllia cladonia is very similar to Dendrophyllia sp. cf. D. horsti diag-

nosed above, especially in colony shape, size, and budding, number of septa, and in

having cylindrical corallites. Dendrophyllia cladonia differs in having smaller corallites

(about 8 mm in GCD), endothecal dissepiments, and in having a highly developed

Pourtalés Plan, in which the two Ss in every system that are adjacent to the Si are

quite large and unite before the S2, merging with the columella as one lamella. The S:

of this species do not quite reach the columella. In Dendrophyllia sp. cf. D. horsti, Ss

do not join one another and the S: are the widest septa.

Distribution

Known only from Port Shepstone, South Africa (Van der Horst 1927) to off

south-eastern Mozambique; 49-457 m.

Dendrophyllia gaditana (Duncan, 1873)

Balanophyllia gaditana Duncan, 1873: 333.

?Dendrophyllia minuscula: Bourne, 1905: 213, pl. 2 (fig. 11, 11A).

Dendrophyllia praecipua Gardiner & Waugh, 1939: 240, pl. 1 (fig. 2).

?Dendrophyllia cf. gaditana: Best et al., 1980: 621.

Non Dendrophyllia minuscula: Gardiner & Waugh, 1939: 237 (= Enallopsammia rostrata).

Non Dendrophyllia minuscula: Boshoff, 1981: 42 (= Dendrophyllia ijimai).

Non Dendrophyllia cf. minuscula: Fricke & Schuhmacher, 1983: 184, fig. 14a.

Dendrophyllia gaditana: Cairns, 1979: 181-182, pl. 36 (figs 5-10); 1984: 25, pl. 4 (fig. I) [synonymy].

New records

AB-371F, 1 branch, USNM 91821; AB-401B, 3 colonies, USNM 91822;

V-2697, 1 colony, IOM; V—2753, 1 colony, IOM.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Indian Ocean: off south-eastern and central Mozambique; off Pemba, Tanzania

(Gardiner & Waugh 1939); ?Seychelles (Best et al. 1980); Madagascar Plateau near

Walters Shoal; ?Gulf of Manaar (Bourne 1905); 65-480 m. Elsewhere: amphi-

Atlantic, Java Sea, off Queensland, Hawaiian Islands (Cairns 1984), off Japan (Cairns

in prep.); 73-505 m.

Dendrophyllia ijimai Yabe & Eguchi, 1934

Fig. 13G

Dendrophyllia ijimai Yabe & Eguchi, 1934: 2026. Eguchi, 1968: C65—C66, pl. C16 (figs 1-2), pl. C22

(fig. 1), pl. C30 (figs 4-5).

Dendrophyllia cf. minuscula: Scheer & Pillai, 1983: 170-171, pl. 39 (figs 5-10).

Dendrophyllia minuscula: Boshoff, 1981: 42.

New records

AB-394B, 1 colony, USNM 91844; Cruise 333, sta. 14, 1 colony, USNM 78543;

off Zanzibar, 84m, 1 colony, USNM 78544; MN-ZK21, 1 colony, USNM 91843.

Reference material: 2 colonies from off Moroisa, Sagami Bay, USNM.

Diagnosis

Coralla large and arborescent, up to 23 cm wide and 20 cm in height, with a

massive base up to 4 cm in diameter. Branches relatively straight and circular in cross-

section, each gradually attenuating to a distal axial corallite 5-6 mm in diameter.

Non-axial corallites circular to slightly elliptical and smaller (4,5-5,5 mm in diameter)

than axials, arranged in three or four irregular rows along branch, each corallite pro-

jecting 2-8 mm perpendicular to branch or slightly tilted anteriorly. Theca striate,

with well-delineated costae. Corallum white; dried tissue dark brown. Septa hexa-

merally arranged in four cycles; however, fourth cycle rarely complete in non-axial

corallites, whereas some Ss do occur in axials. S: independent and by far the largest

septa. S2 also independent but much smaller. S3 smallest of septa, each enclosed by a

larger pair of Ss. Columella large and spongy.

Remarks

These specimens appear to be identical to those reported as Dendrophyllia cf.

D. minuscula by Scheer & Pillai (1983) from the Red Sea. Dendrophyllia minuscula

Bourne, 1905, has much smaller corallites. Among the approximately 30 valid Recent

species of Dendrophyllia, these south-west Indian Ocean specimens are indistinguish-

able from D. ijimai, previously known only from off Japan at 37-366 m. Superficially,

this species resembles Tubastraea micrantha in colour and size, but differ in having

septa arranged in the Pourtalés Plan.

Distribution

Off Durban (Boshoff 1981) and Zululand, South Africa; off Zanzibar, Tanzania;

off Kenya; Red Sea (Scheer & Pillai 1983); 62-223 m. Elsewhere: off Japan,

37-366 m (Eguchi 1968).

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 281

Dendrophyllia fistula (Alcock, 1902a)

Balanophyllia (Thecopsammia) fistula Alcock, 1902a: 109; 1902c: 42, pl. 5 (fig. 36, 36a).

Thecopsammia fistula: von Marenzeller, 1907a: 8—9, text-fig. 6; 1907b: 16-17, pl. 1 (figs a-h).

Dendrophyllia fistula: Gardiner & Waugh, 1939: 237. Pillai & Scheer, 1976: 16. Scheer & Pillai, 1983:

170 [synonymy].

Non Dendrophyllia fistula: Wells, 1954: 472-473, pl. 180 (figs 1-3).

Non Dendrophyllia fistula: Eguchi, 1968: C63, pl. C12 (figs 4-6).

New record

V-2635, 1 colony, IOM. Reference material: 3 syntypes of B. fistula from

Siboga-105, ZMA Coel. 564.

Remarks

Like D. cornucopia Pourtalés, 1871, D. fistula is an unattached, recumbent coral-

lum bearing numerous buds of variable length and position on the primary corallite,

rarely, if ever, having an intact third generation. The specimen reported by Wells

(1954) from the Marshall Islands differs in colony and costal shape and in having

higher generation buds present.

Those species of Dendrophyllia having an unattached, recumbent corallum with

irregular budding of smaller coralla from its theca (as opposed to attached, sympo-

dially branching colonies more typical of Dendrophyllia) were placed in the new

subgenus Dendrophyllia (Alcockia) by Eguchi (1968), but the name Alcockia is a

junior homonym of a fish genus (Goode & Beane 1895) and thus will require a

replacement name.

Distribution

Indian Ocean: off south-eastern Mozambique; off Zanzibar, Tanzania (Gardiner

& Waugh 1939); Red Sea (Von Marenzeller 1907a, 1907b; Gardiner & Waugh 1939;

Scheer & Pillai 1983); Maldive Islands (Gardiner & Waugh 1939); 210-900 m. Else-

where: Sulu Sea (Alcock 1902a, 1902c); 270-275 m.

Genus Enallopsammia

Enallopsammia rostrata (Pourtalés, 1878)

Amphihelia rostrata Pourtalés, 1878: 204, pl. 1 (figs 4-5).

Anisopsammia rostrata: von Marenzeller, 1904: 314-315, pl. 18 (fig. 23).

Madrepora ramea: Gardiner & Waugh, 1939: 226-227.

Dendrophyllia minuscula: Gardiner & Waugh, 1939: 237.

Non Dendrophyllia amphelioides: Gardiner & Waugh, 1939: 238.

Enallopsammia rostrata: Cairns, 1982: 57, pl. 18 Neg 4) [synonymy]; 1984: 27-28. Zibrowius &

Gili, 1990: 39-42, pl. 6 (figs A—-F), pl. 7 (figs A—F).

Enallopsammia amphelioides: Zibrowius, 1973: 45-48, figs 16-20 [synonymy]; 1980: 203-2 4, pl. 106

(figs D-I) [synonymy]. Zibrowius & Grygier, 1985: 131, 134, fig. 51.

New record

V-2731, 1 branch, IOM.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Indian Ocean: off Comores Islands (Zibrowius 1982; Zibrowius & Grygier 1985);

off Reunion (Zibrowius & Grygier 1985); Madagascar Plateau off Walters Shoal

(Zibrowius 1982); off Maldive Islands (Gardiner & Waugh 1939); off Nicobar Islands

(Von Marenzeller 1904); 229-805 m. Elsewhere: amphi-Atlantic, west and central

Pacific, New Zealand region; 229-2 165 m (Cairns 1982).

Genus Jubastraea

Tubastraea micrantha (Ehrenberg, 1834)

Oculina micranthus Ehrenberg, 1834: 304.

Dendrophyllia nigrescens Dana, 1846: 387.

Coenopsammia viridis Milne Edwards & Haime, 1848b: 110.

Dendrophyllia micranthus: van der Horst, 1926: 43-44, pl. 2 (figs 6-7). Scheer & Pillai, 1974: 63,

pl. 29 (fig. 3). Pillai & Scheer, 1976: 16.

Dendrophyllia cf. micrantha: Best et al., 1980: 621.

Tubastraea micranthus: Macnae & Kalk, 1958: 123. Scheer & Pillai, 1983: 175-176, pl. 41 (figs 7-8)

[synonymy]. Schuhmacher, 1984: 94-98, fig. la—b [synonymy]. Zibrowius & Grygier, 1985: 1039.

Tubastraea micrantha: Pichon, 1978: 441. Rosen, 1979: 20. Wells, 1983 [synonymy].

New records

Aldabra, 15 colonies, USNM 79137, 79138, 79140, 79143, 91832; Nossi Bé,

Madagascar, 2 colonies, USNM 91833-34; Rodriques Island, 1 colony, USNM 22018.

Remarks

This species is easily distinguished from the five other valid species in the genus

(T. coccinea Lesson, 1829; T. diaphana (Dana, 1846); T. tagusensis Wells, 1982;

T. floreana Wells, 1982; and T. faulkneri Wells, 1982) by having arborescent, sympo-

dially branched coralla, the others having phaceloid or plocoid colonies.

Distribution

Indian Ocean: off Mozambique (Macnae & Kalk 1958) to the Red Sea (Scheer &

Pillai 1983); Comores Islands (Schuhmacher 1984); Seychelles (Milne Edwards &

Haime 1848); Best et al. 1980); off Madagascar (Pichon 1978); off Mauritius, Maldive

and Nicobar islands (Scheer & Pillai 1974); 0,5-55 m. Elsewhere: western Pacific

(Schuhmacher 1984).

Tubastraea coccinea Lesson, 1829

Tubastraea coccinea Lesson, 1829: 93. Wells, 1983: 243-244, pl. 18 (figs 1-2) [synonymy]. Cairns,

1991: 26-27, pl. 12 (figs c—e).

Lobophyllia aurea Quoy & Gaimard, 1833: 195, pl. 15 (figs 7-11).

Coenopsammia ehrenbergiana Milne Edwards & Haime, 1848b: 109, pl. 1 (fig. 12).

Non Dendrophyllia coccinea: van der Horst, 1926: 45 (= Dendrophyllia sp. cf. D. horsti).

Non Dendrophyllia aurea: van der Horst, 1926: 46-48 (= T. faulkneri).

Dendrophyllia aurea: Macnae & Kalk, 1958: 123.

Non Tubastraea coccinea: Scheer & Pillai, 1983: 175, pl. 41 (figs 5-6).

Tubastraea aurea: Pichon, 1964: 191; 1978: 441. Best et al., 1980: 621. Scheer & Pillai, 1983: 173-174,

pl. 40 (fig. 8) [synonymy]. Schuhmacher, 1984: 94-95.

?Tubastraea sp. Zibrowius & Grygier, 1985: 130.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 283

Fig. 13. A, D. Balanophyllia diffusa, Cr. 329-15, USNM 78594, lateral and calicular views. A X 2,7,

D x 3,6. B, E. Rhizopsammia compacta, MN-ZA49 (USNM 91793) lateral view of a colony and

calice of largest corallite. Bx1,1, Ex1,8. C. Rhizopsammia annae, AB-391J, USNM 91790, a

small colony. x 1,4. F, I. Dendrophyllia sp. cf. D. horsti, MN-ZC11, USNM 91819, lateral view of

colony and calice of a corallite. F x 1,2, 13,7. G. Dendrophyllia ijimai, AB-394B, USNM 91844,

colony with dried black tissue. 0,25. H. Tubastraea diaphana, Nossi Bé, Madagascar,

USNM 91838, colony. x 0,8.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

New records

Aldabra, 0-2 m, 6 colonies, USNM 79139 and 79141; Nossi Bé, Madagascar,

2 colonies, USNM 91836—37.

Remarks

This species is distinguished from the other two species known from the Indian

Ocean by having a plocoid corallum, not dendroid or bushy. Tubastraea coccinea was

compared to the three eastern Pacific species by Cairns (1991, table 4).

Distribution

Indian Ocean: off south-eastern Mozambique (Macnae & Kalk 1958); Madagas-

car (Pichon 1974, 1978); Seychelles (Milne Edwards & Haime 1848b); Red Sea

(Lesson 1829); Chagos Islands; Maldive Islands; Gulf of Manaar; Mergui Archipelago

(Dana 1846); reef depth. Elsewhere: Pacific (Scheer & Pillai 1983); off Japan (Eguchi

1968); Galapagos and Cocos islands (Cairns 1991); western Atlantic (Cairns 1979);

1,5-110 m.

Tubastraea diaphana (Dana, 1846)

Fig. 13H

Dendrophyllia diaphana Dana, 1846: 389, pl. 27 (fig. 3).

Tubastraea diaphana: Scheer & Pillai, 1983: 174, pl. 41 (figs 1-4) [synonymy]. Wells, 1983: 243.

Tubastraea coccinea: Pillai & Scheer, 1976: 17. Scheer & Pillai, 1983: 175, pl. 41 (figs 5-6).

New records

Off Zanzibar, Tanzania, 1 colony, USNM 83697; Nossi Bé, Madagascar, 2 col-

onies, USNM 91838-—39; southern coast of Natal, 9-15 m, 1 colony, USNM 91840.

Remarks

Similar to 7. coccinea in septal arrangement, but colony forms small clumps or

bushes with frequent budding from corallite walls instead of basal coenosteum.

Distribution

Indian Ocean: off Natal, South Africa; off Zanzibar, Tanzania; off north-western

Madagascar; Red Sea (Scheer & Pillai 1983); shallow. Elsewhere: off Singapore

(Dana 1846); central Pacific; Great Barrier Reef (Scheer & Pillai 1983); reef depths.

ACKNOWLEDGEMENTS

We would like to thank the following people who have generously loaned us

specimens used in this study: David G. Herbert (Natal Museum), Michelle van der

Merwe (SAM), D. H. H. Kiithlmann (ZMB), M. H. Schleyer (ORI, Durban), and

Charles Hussey (BM). We are also very grateful to Helmut Zibrowius, who identified

many of the R.V. Anton Bruun specimens and edited an early draft of the manuscript.

Molly Ryan, Smithsonian staff illustrator, prepared Figures 1 and 2. The SEM was

done by SDC in the SEM laboratory of the NMNH, Smithsonian Institution.

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN 285

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290 ANNALS OF THE SOUTH AFRICAN MUSEUM

APPENDIX

STATION LIST

Station Latitude Longitude Depth

number (°S) GE) (m)

R.V. Anton Bruun (AB)

357B PSA 32202; 69

357E 29°10' 32505) 168

358A 299: 32°00" 366

358C 29721 31°58' 366

365C 23°19; 43°33! 439

365D 23205 43°32" 475-695

369J 24°12' 36°01" 1 140

370D 24°28' 35236: 880

370G 24°20' 35°28' 347

370H 24°41" B5528) 311-320

SE 24°46' 35,220! 132

371K 24°46' Som 18: 110

371G 24°49" Saal 73

372B 24°48' 34°59" 42

372G DASd3i 34°56' 55

372J 25°07' 34°34" 106

372L 2200: 34°34" 112

372P Zao: 33°02" 37

373B 26°00' 33°05; 135

S/3H 26°58' 33°54! 896

373] 26°58' 331793" 880

374D 27°08' 34°05' 1 326

389C 30°12' 32°01' 1 360

389E 30°09' S37 930

390S 29S) 31°42' 138

391H 2952 31835) Sil

391J PA SI): Sif

392J 29°19" 31°26' 38

394B 2982s SSH 68-70

399A 2233) 36°10' 925

399B 22230! 36°07' 850-960

399C 21°18" 36°18" 1 510-1 600

401B 19°50' 36°21' 65

400C 20°30' 35°43' 62

408D 16°42' 43°19" 150-300

409E IK UE 43°42" 62

421A 02°54' 40°23' 34

421G 02°S56' 40°28' 240

R.V. Vityaz (V)

2608 12°28' 48°18" 780

2626 24°42" Sara: 320

2629 25528: 351736; 1 250-1 520

2631 25°30' 35°08' 490-535

2634 25505: 34°50' 90-92

2635 25505) 85715! 110-230

2637 252113) S592) 980-1 000

2644 22°19" 43°06' 330-335

2650 2222: 42°59" 950-960

2653 222100" 42°50' 1 440-1 510

2662 22°14" 43°07' 310-315

2668 33°01" 44°09" 1010

Date

30 July 1964

12 August 1964

17 August 1964

18 August 1964

19 August 1964

22 August 1964

23 August 1964

7 September 1964

9 September 1964

10 September 1964

25 September 1964

1 October 1964

2 October 1964

4 October 1964

3 October 1964

15 October 1964

18 October 1964

7 November 1964

8 November 1964

14 November 1988

22 November 1988

23 November 1988

25 November 1988

26 November 1988

2 December 1988

3 December 1988

4 December 1988

8 December 1988

AZOOXANTHELLATE SCLERACTINIA FROM THE SOUTH-WEST INDIAN OCEAN

Station Latitude Longitude

number = (°S) (°E)

R.V. Vityaz (V)_ (cont.)

2671 32256) 45°01"

2674 38°01" 45°27'

2686 33°04" 43°52!

2697 3315) 43°55!

2699 S3e 18) 43°54"

2706 33°03' 44°33'

2716 Slip 44°55"

2721 322255 43°37"

2722 B2725) AS ii

2724 33°09" 43°50'

2731 SB yalliey: 44°08"

2733 33223) 44°06'

2753 SBI 43°52"

2764 33°10' 43°41"

2767 33°08' 45°49"

2803 Lie 61°44"

2804 OG; 62°14'

2809 10°29' 61°09"

2814 09°38" 60°50'

2815 09°40' 60°30'

2816 09°31' Spb)

R.V. Meiring Naude (MN)

SM=129_-30%53;4" -30231;7'

SM=1162)5-322959:0—~ 28231,0:

SM-174 33°19,6' = -27°52,4'

SM-226 32°28,6' 28°58,8'

SM-232 32°14,9' = 29° 10,4"

DEX 193i 2 9932-2 S198:

ZA-43 Z2OLS4 Op 3250550

ZA-48 ZOZ9339) 3255950"

ZA-49 2679453) 132099;))

ZA-51 20-92-81, _ 32;56,0!

ZB-11 DA O02 eS e543)

ZB-14 27T20058" 3255453).

ZB-17 ZOOS 2) Be S2255; 3),

ZB-18 2ISO00;4") ~ 32255"2)

ZB-20 PAV SIE -8y2 Spe

ZB-23 21203;0% 3225457"

ZB-24 DiS; On Ol O45)

ZB-25 DIZ03 7 | @32293.4"

ZB-27 DIgOS9Le "32255.0;

ZC-9 PAPA et OPO RPa

ZC-10 2i1206:0, ~ 32253,3)

ZC-11 DOG Se, 5292.9)

ZD-4 Dislele 325 50:9)

ZD-5 Died 8 320s

ZD-6 Del atOr eS 2249 7:

ZD-7 Di NOU 3224957):

ZD-8 PUN eae aS VIEL Bays

ZD-10 Zeus 3225054;

ZH-18 DI3Y.8" 3242-8)

ZH-19 LICSL,8:, ~ 32 442;8)

ZH-23 DISS 8y 1) SL c42D)

ZH-26 PH hes 1. 9 Sa a. PO

32°41,6'

ZED A 27350:

Depth

(m)

LETS

1 600-1 610

650-665

160

530-610

970-980

630-680

1 000-1 030

680-720

280-360

750-755

410-480

910-920

260

87-110

230-235

110-115

1 650-1 700

1 520-1 720

430-835

Date

9 December 1988

10 December 1988

12 December 1988

13 December 1988

14 December 1988

15 December 1988

16 December 1988

17 December 1988

18 December 1988

22 December 1988

24 December 1988

7 January 1989

8 January 1989

9 January 1989

11 May 1977

25 May 1978

28 May 1978

24 June 1979

25 June 1979

1 June 1990

3 June 1990

4 June 1990

5 June 1990

6 June 1990

7 June 1990

8 June 1990

2 June 1990

7osh\|

DOD

ANNALS OF THE SOUTH AFRICAN MUSEUM

Station Latitude

number

R.V. Meiring Naude (MN) (cont.)

(°S)

Longitude

(°E)

ZK-20 DISAT As 32°34,0'

ZK-21 27°47,6' 32°39 1"

ZK-25 27°46,4' 32°39,4'

ZL-3 DAD4. 3% 32°38,0'

ZQ-8a 29°05,3' 32°08,3'

ZU-13 29°31,8' 3122729!

ZU-15 29°39,8' 31230) 1

ZV-5 29°44,3' S276)

ZV-20 29°41,6' Silae2 180)

ZV-21 29°41,4' Shp

ZW-6 29°45 ,9' S176)

ZW-8 29°45 ,9' SSD

ZCC-1 27°08,3' S224

ZCC-2 27°09,9' B2 25140);

ZDD-2 QT N46! 32°48,8'

ZDD-3 DANS ES! 32°49,5'

ZDD-4 DilgAlSeOn 32°49 ,3'

ZDD-5 DIN eS. 32°49 ,8'

ZDD-7 Py Aleyey 32°49,3'

Miscellaneous stations

Vema

14-SAT6 29°48' BiloGy

19-28 35°40' D259"

Marion Dufresne (Cruise 27)

Sta. 4 08°29'N 79°19'

Sardinops

CD32 34°13,9' 18°29,3'

Depth

(m)

Date

9 June 1990

9 June 1988

17 June 1989

19 June 1989

20 June 1989

21 June 1989

7 June 1990

8 June 1990

23 April 1958

28 July 1981

10 April 1971

EAMFRO (East African Marine Fisheries Research Organization, Zanzibar)

Station ro Latitude

number Locality (°S)

Cis north Kenya Banks —

Unknown vessel

Cr. 329-10 off Zanzibar a

Cr. 329-15 off Zanzibar —

Cr. 333-14 — 02°40,5'

Manihine

381-1 04°13'

381-3 05°27'

381-63 07°32'

Longitude Depth

(°E) (m)

122

183

273

O07 223

CAeSe 250

°09' 302

25256) 240

Date

25 Feb 1971

11 August 1971

9 December 1971

24 October 1974

31 October 1974

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

larly Articles 22 and 51).

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

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

etc.

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

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

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

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

from the scientific name by a colon.

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

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

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

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, 1861: 87.

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

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

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

dash, not comma, separates consecutive numbers.

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

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

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

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

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

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

Holotype

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

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

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

7. SPECIAL HOUSE RULES

Capital initial letters

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

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

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

initials or full names

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

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

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

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

article, such as

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

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

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

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

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

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