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Monograph
Azooxanthellate Scleractinia (Cnidaria, Anthozoa) from South Africa
expand article infoZoleka N. Filander§, Marcelo V. Kitahara|, Stephen D. Cairns#, Kerry J. Sink¤§, Amanda T. Lombard§
‡ Biodiversity and Coastal Research, Oceans and Coasts, Department of Environment, Forestry, and Fisheries, Cape Town, South Africa
§ Nelson Mandela University, Port Elizabeth, South Africa
| Universidade Federal de São Paulo, Santos, Brazil
¶ Universidade de São Paulo, São Sebastião, Brazil
# Department of Invertebrate Zoology, Smithsonian Institution, Washington, United States of America
¤ South African National Biodiversity Institute, Cape Town, South Africa
Open Access

Abstract

Globally, South Africa ranks in the top five countries regarding marine species richness per unit area. Given the high diversity, it is not surprising that many invertebrate taxa in the region are poorly characterised. The South African azooxanthellate Scleractinia (Anthozoa) is one such taxonomic group, and was last reviewed by Boshoff in 1980. Although more recent regional publications have reported on some species, there has not been a faunistic review that accounts for the country’s species diversity since then. Moreover, numerous unidentified specimens representing more than three decades of sampling effort have accumulated. In this study the authors update the state of knowledge of South African azooxanthellate coral species. Specimens, particularly those within the extensive collections of the Iziko South African and Smithsonian museums, were morphologically examined and identified. Other data considered included historic data represented as imagery data, associated species data from recent research surveys, and the scientific literature. To date, the study has increased the total number of known species from 77 to 108 across eleven families, 28 new South African records, and three are new species with one new genus.

Keywords

Ahermatypic, corals, hermatypic, revision, taxonomy

Introduction

The South African marine environments host a variety of fauna that encompasses at least 12,000 species (Griffiths et al. 2010), although many benthic taxa remain poorly understood (Gibbons 1999; Griffiths et al. 2010). This diversity, and a high level of endemism of species, is influenced by the dynamic nature of the oceanographic regimes that surround the country (Brown and Jarman 1978; Thandar 1989; Awad et al. 2002; Griffiths et al. 2010). These regimes are primarily distinguished by water temperature. The cold, upwelling nutrient-rich Benguela Current flowing northwards along the Atlantic margin results in high biological productivity, which in turn supports high species abundance (Shannon and Nelson 1996), but low species diversity (Griffiths et al. 2010). Along the eastern margin the warm fast southward-flowing Agulhas Current brings nutrient-poor waters, which drive high species diversity in this region (Heydorn et al. 1978). Both these currents interact in the Southern margin region resulting in a unique environment which promotes high endemism patterns for most benthic invertebrate marine fauna (Lutjeharms et al. 2000; Awad et al. 2002).

This heterogenous physical environment underpins South Africa’s high marine species per unit area, which the country is reported to rank amongst the top five globally (Costello et al. 2010). However, it is estimated that more than a third of the country’s fauna remains to be characterised (Costello et al. 2010). This paucity of taxonomic knowledge is particularly acute for the deeper waters (> 200 m), which represent 90% of the marine territory (Lombard et al. 2004; Costello et al. 2010; Griffiths et al. 2010; Sink et al. 2012, 2019). In addition to the existing challenges in sampling efforts across the shallow and offshore regions (Griffiths et al. 2010), the lack of regional taxonomists (Gibbons et al. 1999) has also contributed to the high estimates of undescribed species (Costello et al. 2010; Griffiths et al. 2010). The importance of regional guides has been emphasised by Costello et al. (2010), who demonstrated how published monographs and guides within a region correlate with the state of knowledge of known taxa. These findings are in keeping with the observations of Gibbons et al. (1999) and Griffiths et al. (2010) that South Africa has low taxonomic efforts despite its high biodiversity. Although regional expertise has recently expanded within invertebrate groups such as Echinodermata (Olbers 2016; Filander and Griffiths 2017), Crustacea (Biccard 2012; Landschoff 2011), and one other Cnidaria class, the Actiniaria (Laird 2013), there is still a notable gap in taxonomy of several other cnidarian orders.

One specific group with a paucity in regional taxonomic understanding is the azooxanthellate scleractinian corals and this is the focus of the work presented here. Our knowledge on the South African azooxanthellate scleractinian corals is based on ten publications (Duncan 1876; Gardiner’s 1902a, b, 1904; van der Horst 1927, 1938; Boshoff 1981; Zibrowius and Gili 1990; Cairns and Keller 1993; Cairns and Zibrowius 2016) none of which holistically details the South African fauna across the oceanographic regimes (Agulhas and Benguela regions). A significant contribution to the South African azooxanthellate fauna is documented in the 20th century (Table 1).

Table 1.

Significant papers contributing to the South African azooxanthellate coral fauna to date. References are presented chronologically (denoted by capital letters: A = Duncan 1876, B = Gardiner 1902a, C = Gardiner 1902b, D = Gardiner 1904, E = von Marenzeller 1904, F = van der Horst 1927, G = van der Horst 1933, H = van der Horst 1938, I = Boshoff 1981, J = Zibrowius and Gili 1990, K = Cairns and Keller 1993, L = Cairns and Zibrowius 2016, and M = present study) and then following family and species alphabetically; for which the first column lists the accepted species name at time of publication followed by an authority column. New species records are identified by a numeric superscript.

Species Authority References
A B C D E F G H I J K L M
1. Anomocora fecunda (Pourtalès, 1871) X
Anomocora marchadi (Chevalier, 1966) X X X
2. Aulocyathus sp. cf. matricidus (Kent, 1871) X
3. Aulocyathus juvenescens von Marenzeller, 1904a X
4. Caryophyllia (A.) dentata (Moseley, 1881) X
5. Caryophyllia (A.) grayi (Moseley, 1881) X
Caryophyllia (C.) cinticulata (Alcock, 1898) X
6. Caryophyllia (C.) diomedeae von Marenzeller, 1904b X
Caryophyllia (C.) ephyala Alcock, 1891 X
Caryophyllia (C.) grandis Gardiner & Waugh, 1938 X
Caryophyllia (C.) lamellifera Moseley, 1881 X
Caryophyllia (C.) profunda Moseley, 1881 x X
Caryophyllia (C.) quadragenaria Alcock, 1902a X
Caryophyllia (C.) rugosa Moseley, 1881 X
7. Caryophyllia (C.) sarsiae Zibrowius, 1974b X
8. Caryophyllia (C.) scobinosa Alcock, 1902a X
Caryophyllia (C.) stellula Cairns, 1998 X
Caryophyllia (C.) valdiviae Zibrowius & Gili, 1990 X X
9. Crispatotrochus cornu (Moseley, 1881) X
Desmophyllum dianthus (Esper, 1794) X X X
Desmophyllum pertusum (Linnaeus, 1758) X X
Goniocorella dumosa (Alcock, 1902c) X
Heterocyathus aequicostatus Milne–Edwards & Haime, 1848a X
Heterocyathus alternatus Verrill, 1865 X
Heterocyathus monileseptatum sp. nov. X
Heterocyathus sulcatus (Verrill, 1866) X
Labyrinthocyathus delicates (von Marenzeller, 1904) X X X X
Monohedotrochus capensis comb. nov. (Gardiner, 1904) X X X
10. Polycyathus sp X
Rhizosmilia robusta Cairns in Cairns & Keller, 1993 X
Solenosmilia variabilis Duncan, 1873 X X
Stephanocyathus (Acinocyathus) explanans (von Marenzeller, 1904a) X X
11. Stephanocyathus (Odontocyathus) campaniformis (von Marenzeller, 1904a) X
12. Stephanocyathus (Odontocyathus) nobilis (Moseley, 1873) X
13. Tethocyathus virgatus (Alcock, 1902a) X
Trochocyathus (T.) sp. 1 Gardiner,1904 X X X
Trochocyathus (T.) sp. 2 X
Trochocyathus (T.) sp. 3 X X
Trochocyathus (T.) sp. cf. rawsonii sensu Cairns in Cairns and Keller 1993 X X
14. Vaughanella concinna Gravier, 1915 X
Deltocyathus italicus (Michelotti, 1838) X
Deltocyathus rotulus (Alcock, 1898) X
Atlantia denticulata sp. nov. X
Balanophyllia (B.) bonaespei van der Horst, 1938 X
Balanophyllia (B.) capensis Verrill, 1865 X X
Balanophyllia (B.) diademata van der Horst, 1927 X
Balanophyllia (B.) diffusa Harrison & Poole, 1909 X
15. Balanophyllia (B.) sp. cf. malouinensis Squires, 1961 X
Balanophyllia (B.) vanderhorsti Cairns, 2001 X
Balanophyllia (E.) stimpsonii (Verrill, 1865) X
Dendrophyllia arbuscula van der Horst, 1922 X
Dendrophyllia cladonia van der Horst, 1927 X X
Dendrophyllia cornigera (Lamarck, 1816) x X
Dendrophyllia dilatata van der Horst, 1927 X X
Dendrophyllia ijimai Yabe & Eguchi, 1934 X X
Dendrophyllia sp. 1 X
Ednapsammia columnapriva sp. nov. X
16. Enallopsammia pusilla (Alcock, 1902a) X
17. Enallopsammia rostrata (Pourtalès, 1878) X
Endopachys grayi Milne–Edwards & Hamie, 1848b X X X
18. Endopsammia philippensis Milne–Edwards & Haime, 1848b X X
Heteropsammia cochlea (Spengler, 1781) X
19. Heteropsammia eupsammides (Gray, 1849) X
Pourtalopsammia togata (van der Horst, 1927) X X
Rhizopsammia annae (van der Horst, 1933) X X X
Rhizopsammia compacta Sheppard & Sheppard, 1991 X X
20. Rhizopsammia verilli van der Horst, 1922 X
Tubastraea sp. cf. diaphana (Dana, 1846) X X
Tubastraea coccinea Lesson, 1829 X
Tubastraea micranthus (Ehrenberg, 1834) X
Flabellum (F.) leptoconus Cairns & Zibrowius, 2016 X X X
Flabellum (F.) pavoninum Lesson, 1831 X X X X
21. Flabellum (F.) politum Cairns, 1989a X
Flabellum (U.) alabastrum Moseley, 1873 X
22. Flabellum (U.) lowekeyesi* Squires & Ralph, 1965 X
23. Javania antarctica (Gravier, 1914) X
Javania insignis Duncan, 1876 X
Placotrochides scaphula Alcock, 1902c X
Rhizotrochus typus Milne–Edwards & Haime, 1848a X X X
Truncatoflabellum formosum Cairns, 1989a X
Truncatoflabellum gardineri Cairns in Cairns & Keller, 1993 X
Truncatoflabellum inconstans (von Marenzeller, 1904a) x X
24. Truncatoflabellum multispinosum Cairns in Cairns & Keller, 1993 X
Truncatoflabellum pusillum Cairns, 1989a X
Truncatoflabellum sp. X
Truncatoflabellum zuluense Cairns in Cairns and Keller, 1993 X X
Fungiacyathus (B.) hydra Zibrowius & Gili, 1990 X
Fungiacyathus (B.) sibogae (Alcock, 1902a) X
Fungiacyathus (F.) stephanus (Alcock, 1893) X
Fungiacyathus (F.) sp X
Guynia annulata Duncan, 1872 X
Letepsammia formosissima (Moseley, 1876) X
Letepsammia franki Owens, 1994 X X
25. Rhombopsammia niphada Owens, 1986a X
Stephanophyllia fungulus Alcock, 1902b X
Madrepora oculata Linnaeus, 1758
Culicia excavata (Milne–Edwards & Haime, 1849) X
26. Culicia sp. cf. australiensis Hoffmeister, 1933
Culicia sp. cf. tenella natalensis Dana, 1846 X X
27. Stenocyathus vermiformis (Pourtalès, 1868) X
28. Cyathotrochus pileus (Alcock, 1902a) X
Deltocyathoides orientalis (Duncan, 1876) X X
Deltocyathoides sentus Kitahara & Cairns, 2021 X
Sphenotrochus (E.) gilchristi Gardiner, 1904 X X X
Sphenotrochus (S.) aurantiacus von Marenzeller, 1904a X X X
Sphenotrochus (S.) evexicostatus Cairns in Cairns & Keller, 1993 X X
Sphenotrochus (S.) imbricaticostatus Cairns in Cairns & Keller, 1993 X X
Tropidocyathus lessonii (Michelin, 1842) X

Prior to this, just one rare solitary form (Culicia natalensis Dana, 1846) was reported by Duncan in 1876 (Table 1). Subsequently, Gardiner’s (1902a, b, 1904) research significantly improved the knowledge base of the South African fauna and he reported 16 species (four representing new species). Gardiner’s (1902a, b, 1904) flabellids and caryophylliid contribution was then later complemented by von Marenzeller 1904a reported six species (two of which were new to science). Following this, van der Horst (1927, 1933, 1938) documented ten dendrophylliid species. Both the Gardiner and van der Horst accounts are mainly based on specimens collected through the University of Cape Town Ecological Surveys (UCTES) and remain the foundation of azooxanthellate coral research in South Africa. Five decades later, Boshoff (1981) published a checklist on 54 azooxanthellate scleractinian corals from the south-west Indian Ocean. Subsequent authors (Zibrowius and Gili 1990; Cairns and Keller 1993) highlighted the need for the re-analysis of Boshoff’s account. Other papers which include South African records are biogeographic reviews such as Zibrowius and Gili’s (1990) south-east Atlantic paper documenting five species, Cairns and Keller’s (1993) south-west Indian Ocean review that accounts for 45 species, and Cairns and Zibrowius’s (2016) new flabelliid species (Flabellum (Flabellum) leptoconus Cairns & Zibrowius’s, 2016). Since then, numerous specimens have remained unidentified in the Iziko Museum of South Africa collection. Moreover, additional specimens are being collected by offshore research collaborative programs involving the Department of Environment, Fisheries and Forestry (DEFF), the African Coelacanth and Ecosystem Program (ACEP), the South African Environmental Observation Network (SAEON), and the Ecosystem Approach to Fisheries (EAF)-Nansen Programme. Here we present the first faunistic review of South African azooxanthellate scleractinian corals, increasing the number of known species from 77 to 108 (Table 1).

Materials and methods

This study is primarily based on the azooxanthellate Scleractinia deposited at the Iziko South African Museum (Cape Town) and at the National Museum of Natural History (Smithsonian, Washington DC), comprising more than 600 samples collected through six historical expeditions (RV ‘Anton Bruun’, Benguela IV, RV ‘Meiring Naude’, RV ‘Pieter Faure’, ‘Sardinops’, and University of Cape Town Ecological Surveys). Other collections considered include 38 samples from the Boshoff Collection housed at the Oceanographic Research Institute (Durban), 71 samples from the African Coelacanth Ecosystem Programme (Deep-Secrets and Imida surveys), 31 samples from the Department of Environment, Forestry and Fisheries/South African Environmental Observation Network demersal surveys, ten samples collected on the 2018 Nansen survey, and five from the Department of Environment, Forestry and Fisheries offshore benthic surveys. Overall, these samples comprise ca. 3100 specimens, all of which originated within the South African territorial sea and Economic Exclusive Zone (Suppl. material 1), and covering a depth range from the intertidal zone to 1,420 m depth. Although the Prince Edwards Islands constitute the South African territory, samples from this locality were not considered in the current study owing to the region exhibiting distinctive oceanographic patterns as compared with the surroundings of mainland South African surroundings. Therefore, the Prince Edwards Islands fauna will be reviewed subsequently.

Morphological descriptions follow the terminology used by Cairns (1989a, b, 1997, 2001), and Cairns and Kitahara (2012). Diagnoses for genera were amended from existing literature (Cairns 2001; Cairns and Kitahara 2012). Species are presented in alphabetical order according to family, genus, and species. Each species entry is followed by a synonym list, type locality, and type material (if known). The section on material examined provides the catalogue number or a sample identifier term (arranged in chronological order), followed by the number of specimens (in brackets), sampling location and depth. Both catalogue and sample identifier numbers are presented as written on the labels (e.g., BMNH #). Previously reported South African records are highlighted in bold. Furthermore, refined regional localities are given in which records with coordinate information (Suppl. material 2: Fig. S1) follow a hierarchical approach that gives distance of coral records to the closest gazetted coastal town (in uppercase) and estuary system (Suppl. material 2: Figs S2–5). The methodology undertaken to standardize place names caters for international and national/indigenous communities, whilst locality given on the catalogue label is presented for records without coordinate information. Species descriptions are based mostly, but not solely, on the South African material examined. Imagery data of some South African specimens, shared by Dr Helmut Zibrowius, were also considered and presented. Although these specimens may represent misplaced records as the specimens with Iziko accession numbers were not traceable in the museum’s collection (Suppl. material 1). For all species, following their morphological descriptions, the local and global distribution is provided with a depth range. South African (local) distribution are presented, in lower cases, within the following regions: western margin – from the Namibian border to Cape Point; the southern margin – eastwards from Cape Point to the Mbashe River; and the eastern margin – the region extending from the Mbashe River to the Mozambique border. A locality range is also presented in this section and limited to the closest coastal town. Remarks containing morphological comparisons and, when pertinent, a brief history of the taxonomy of the species in South Africa, is provided. Furthermore, new knowledge on the regional distribution is also added in the remarks section, where applicable. Finally, images from most species identified are presented.

List of abbreviations

Museums and collection institutes

BMNH British Museum of Natural History London (now NHMUK);

IM Indian Museum, Calcutta;

IO Institute of Oceanology, Moscow;

MCZ Museum of Comparative Zoology, Harvard University, Cambridge;

MNHN Muséum national d’Histoire naturelle, Paris;

MOM Muséum Océanographique de Monaco, Monaco;

MoNZ Museum of New Zealand Te Papa Tongarewa, Wellington;

NHMUK Museum of Natural History London;

NMNH National Museum of Natural History, Smithsonian, Washington DC;

ORI Oceanographic Research Institute, Durban;

SAMC South African Museum, Cape Town;

SAMH South African Museum Hydroids;

WAM Western Australian Museum, Perth;

YPM Yale Peabody Museum, New Haven;

ZMA Zöologisch Museum, Amsterdam;

ZMB Zoologisches Museum, Berlin.

Expeditions and institutions

AB RV ‘Anton Bruun’;

ACEP African Coelacanth Ecosystem Programme;

CCS Cape Canyon Survey (Department of Environment, Forestry and Fisheries: DEFF);

DSC Deep Secrets Cruise (African Coelacanth Ecosystem Programme: ACEP);

DTE Deutschen Tiefsee-Expedition;

MN RV ‘Meiring Naude’;

PF RV ‘Pieter Faure’;

SAEON South African Environmental Observation Network;

SVMEC Southern margin Vulnerable Marine Ecosystem Cruise (Department of Environment, Forestry and Fisheries: DEFF);

UCTES University of Cape Town Ecological Survey.

Morphological terms

BD Basal diameter;

GCD:H Ratio of greater calicular diameter to height of a solitary corallum;

GCD:LCD Ratio of greater calicular diameter to lesser calicular diameter;

H:D Ratio of height to diameter of a solitary corallum;

LCD Lesser calicular diameter;

PD Pedicel diameter;

GPD Greater pedicel diameter;

PD:GCD Ratio of pedicel diameter to greater calicular diameter of a solitary corallum;

Sx, Cx, Px Septa, costae, or pali (respectively) of cycle designated by the number;

Sx > Sy Septa of cycle × wider than those of cycle y’

Systematic account

Order Scleractinia

Family Caryophylliidae Dana, 1846

Anomocora Studer, 1877

Diagnosis

Solitary, subcylindrical, free. Tendency to bud new coralla from margin zone with subsequent loss of organic connection. Wall thin. Columella trabecular, no pali.

Type species

Coelosmilia fecunda Pourtalès, 1871, by monotypy.

Anomocora fecunda (Pourtalès, 1871)

Fig. 1A, B

Coelosmilia fecunda Pourtalès, 1871: 21–22 (in part: pl. 1, fig. 12, pl. 6, figs 14–15).

Coenosmilia fecunda. –Zibrowius 1980: 131–133 (in part: pl. 67, figs A–K).

Parasmilia fecunda. –Gardiner and Waugh 1939: 229.

Anomocora fecunda. –Cairns 1979: 127–129, pl. 24, figs 6–8, Map 35. –Cairns 2000: 128. –Reyes et al. 2009: 25–26, fig. 4L, M. –Kitahara and Cairns 2021: 452–454, figs 244E–G, 245.

Type locality

Off Southern Straits, Florida; 124–576 m (Cairns 1979).

Type material

Six syntypes are deposited at the MCZ (Cairns 1979).

Material examined

SAMC_A073042 (1 specimen): 53 km from Shaka’s Rock/46 km off Zinkwasi Estuary, 29°32'53.88"S, 31°47'12.11"E; 200 m.

Description

Corallum cylindrical, straight to gently curved, tapering towards a broken base. Axial corallite of examined specimen measures 8.1 mm in CD and 48.9 mm in H, and bears 18 secondary corallites. Secondary corallites bud irregularly, usually perpendicular to axial corallite. Calices circular to slightly elliptical (GCD:LCD = 1.00–1.1), calicular margin slightly serrated. Specimen examined has two scars of former buds on theca, with secondary corallites rarely exceeding 5 mm in CD. Costae well developed, particularly at calicular margin, corresponding with septa in size. C1 most prominent, higher costae progressively smaller. Intercostal striae shallow and narrow. Theca thin. Corallum white, with C1–2 having light greenish brown tints.

Septa hexamerally arranged in four cycles, the last cycle being incomplete, according to formula: S1 > S2 > S3 > S4 (46 septa). S1 most exsert, reaching columella with straight and vertical axial margins. S2 less exsert and half the width of S1, also with straight axial margins bearing slender ribbons. S3 not exsert, 2/3 the size of S2, also bearing ribbons that intermingle with those of S2. S4 rudimentary, with sinuous to slightly dentate axial margins. Septa mostly smooth, with growth lines along septal margin, and faint granulation perpendicular to septal margin. Fossa deep, with poorly developed trabecular columella, which is indistinguishable from S2 and S3 intermingled slender ribbons.

Distribution

Regional: Eastern margin of South Africa, off Shaka’s Rock; 200 m. Elsewhere: Bahamas; Caribbean (Reyes et al. 2009); eastern Gulf of Mexico; northeastern Brazil (off Maranhão); St. Peter and St. Paul Rocks (Cairns 1979); Azores; Madeira; Canary Islands (Zibrowius 1980); New Caledonia (Kitahara and Cairns 2021); 37–640 m.

Remarks

According to Cairns (1979), Anomoroca fecunda closely resembles Coenosmilia arbuscula Pourtalès, 1871, but differs in having a more elongated corallum with thin costae forming continuous ridges. Further differences are highlighted in budding: in A. fecunda buds appear randomly and detach from the theca of parent corallum before the third generation appears. However, in C. arbuscula, new buds appear equally spaced around the calicular perimeter and remain firmly attached, resulting in a small bushy colony of which the main corallite gives rise to five successive generations. Differences in columella are also noted between the two species, with A. fecunda having a faint trabecular columella and C. arbuscula a well-developed papillose columella. Examined specimen represents a new record to the southwest Indian Ocean.

Figure 1. 

A, B Anomocora fecunda (SAMC_A073042, off Shaka’s Rock, 200 m) A calicular view B lateral view C, F Anomocora marchadi C, D (SAMC_A090093, off Durban, 49 m) C calicular view D lateral view E, F (SAM_H3100, off Shaka’s Rock, 66 m) E calicular view F lateral view G, H Aulocyathus juvenescens (Mortensen25, off Durban, 424 m) G calicular view H lateral view I, J Aulocyathus sp. cf. matricidus (DIa1, locality data unknown) I calicular view J lateral view K, L Caryophyllia (Acanthocyathus) grayi (USNM 91541, off Kosi Bay, 98 m) K calicular view L lateral view M, N Caryophyllia (Caryophyllia) diomedeae (MN_SM129, off Margate, 850 m) M calicular view N lateral view N, O Caryophyllia (Caryophyllia) ephyala (SAMC_A072974, off Paternoster, 440 m) O Calicular view P Lateral view. Scale bars: 10 mm.

Anomocora marchadi (Chevalier, 1966)

Fig. 1C–F

Ceratotrochus johnsoni. –Gardiner 1904: 118–119, pl. 1, figs 5A–C, pl. 2, fig. M. –Gardiner and Waugh 1938: 188.

Dasmosmilia marchadi Chevalier, 1966: 944–949, pl. 5, figs 3, 4.

Asterosmilia marchadi. –Cairns and Keller 1993: 249, fig. 6A, B.

Balanophyllia capensis. –Boshoff 1981: 40 (in part).

Anomocora marchadi. –Cairns 2000: 130–131. – Cairns 2004: 276. –Reyes et al. 2009: 26, 27, fig. 4A–Q.

Type locality

Off Cape Verde, Senegal (RV ‘Gerard Tréca’ stn 18–2–1954); 97–98 m (Chevalier 1966).

Type material

The holotype is deposited at the MNHN (Cairns 1979).

Material examined

SAMC_A090093 (1 specimen): Eastern margin, 14 km off Durban/12 km Mbokodweni Estuary, 29°58'00.00"S, 31°01'59.99"E; 49 m. SAM_H1683 (1 specimen): Western margin, 22 km off Cape Town/13 km off Elsies Estuary, 34°04'59.99"S, 18°19'59.99"E; depth unknown. SAM_H3098 (2 specimens): Eastern margin, 6 km off Durban/9 km off Umgeni Estuary, 29°52'59.99"S, 31°03'05.00"E; 86 m. SAM_H3099 (7 specimens): Eastern margin, 19 km from Shaka’s Rock/3 km off Mdloti Estuary, 29°38'59.99"S, 31°07'59.99"E; 71–73 m. SAM_H3100 (1 specimen): Eastern margin, 9 km from Shaka’s Rock/2 km off Tongati Estuary, 29°34'00.00"S, 31°10'59.99"E; 66 m. ORI_EIa4 (1 specimen): no locality data. USNM 91561 (1 specimen): Eastern margin, 41 km south of Ponta Do Ouro/26 km off Kosi Bay Estuary, 27°13'03.60"S, 32°49'18.00"E; 60 m.

Imagery data

BMNH 1950.1.10.118 (2 specimens), BMNH1950.3.22.13 (2 specimens): no locality data. SAM_H1456 (1 specimen): Eastern margin, 9 km from Shaka’s Rock/2 km off Tongati Estuary, 29°34'00.00"S, 31°10'59.99"E; 66 m. SAM_H2806 (1 specimen): no locality data. MN_ZH17 (1 specimen): Eastern margin, 59 km off Cape Vidal/10 km of Mgobezeleni Estuary, 27°37'00.00"S, 32°40'54.00"E; 65–70 m.

Description

Corallum solitary, ceratoid, tapering to a free pointed base. Axial corallite with ≤ two secondary corallites. Corallites bud from margin zone of parent corallum. Budding extra-tentacular. Calices round to slightly elliptical (GCD:LCD = 1.0–1.1), with serrated calicular margin. Largest specimen examined (SAMC_A90093) 9.8 × 8.9 mm in CD and 16.1 mm in H. Costae prominent and unequal in width. C1–2 wider than C3–4. All costae granulated and separated by broad intercostal striae extending from calicular margin to base. Corallum white to reddish brown.

Septa hexamerally arranged in four cycles according to formula: S1 > S2 > S3 > S4 (48 septa). S1 highly exsert, with straight axial margin that fuse to columella deep in fossa. S2 slightly less wide and less exsert than S1, but otherwise similar in profile. S3 ½ width and less exsert than S2, with dentate axial margin, each bearing a paliform lobe (P3). In each system, a pair of P3 fuse before S2 near columella. S4 as exsert as S3, but rudimentary in development. Septal faces bear granules arranged perpendicular to septa margin. Fossa of moderate depth, containing a crispate columella which is usually indistinguishable to pali.

Distribution

Regional: Western and eastern margin of South Africa, from Cape Town towards the Kosi Bay estuary, (41 km south of Ponta Do Ouro: Mozambique); 49–86 m. Elsewhere: off Pensacola; Florida; southern Caribbean from Colombia to Peninsula de Paria Venezuela (Cairns 1979; Reyes et al. 2009); from Spanish Sahara to Gabon (Zibrowius 1980); Philippines; Indonesia (Cairns and Zibrowius 1997); South China Sea; Mozambique; off Pemba; Tanzania; Maldives (Gardiner and Waugh 1938; Cairns and Keller 1993); 32–229 m.

Remarks

Anomocora marchadi differs from A. prolifera (Pourtalès, 1871) in having P3 fusing before S2 near the columella (Cairns 1979). However, A. prolifera has not yet been reported from South Africa, thus A. marchadi differs from the only other species reported in the region (A. fecunda), in having paliform lobes indistinguishable from the columellar elements. Anomocora marchadi was first reported from South Africa by Gardiner (1904), off Cape Natal. Subsequently, Boshoff (1981) incorrectly identified this species as Balanophyllia capensis. Nonetheless, material examined herein extends the known South African distribution of this species further north towards Durban.

Aulocyathus von Marenzeller, 1904a

Diagnosis

Corallum solitary, ceratoid, and free. Evidence of budding from a longitudinally fragmented of the parent corallum common. Costae poorly defined. Upper, distal septal margins join theca below upper thecal margin usually forming a calicular thecal rim. Slender paliform lobes occasionally present before S1–3. Columella trabecular.

Type species

Aulocyathus juvenescens von Marenzeller, 1904a, by monotypy.

Aulocyathus juvenescens von Marenzeller, 1904a

Fig. 1G, H

Aulocyathus juvenescens von Marenzeller, 1904a: 301–302, pl. 18, fig. 17. –Zibrowius 1980: 107. –Cairns and Keller 1993: 247. –Cairns 1994: pl. 26, figs H, I. –Cairns and Zibrowius 1997: 130. –Cairns 1999a: 104, fig. 15H. –Kitahara and Cairns 2021: 409–411, figs 221, 222G–I.

Type locality

Off Pemba and Zanzibar Island, Tanzania (SS ‘Valdivia’ stn. 243 and 245: 6°39'1"S, 39°30'8"E, 5°27'9"S, 39°18'8"E, respectively); 400–463 m (von Marenzeller 1904a).

Type material

Syntypes are deposited at the ZMB (Cairns and Keller 1993).

Material examined

None.

Imagery data

‘Galathea Expedition’ stn. 196 (2 specimens): Eastern margin, 33 km off Durban/31 km off Beachwood Mangroves, 29°55'00.00"S, 31°19'59.99"E; 425–430 m. ‘Mortensen-Java Expedition’ stn. 25 (1 specimen): Eastern margin, off Durban, 424 m.

Description

Corallum solitary, ceratoid, and usually attached to a fragment of the parent corallum. Calice circular, with finely serrated calicular margin. Largest imaged specimen (Galathea Expedition stn. 196) 6.5 × 6.5 mm in CD, and ≤ 11.0 mm in H. Costae restricted to calicular margin, with low intercostal striae. Theca smooth and porcelaneous. Corallum white, with longitudinal light brown pigmentation.

Septa hexamerally arranged in four cycles, the last cycle being incomplete, according to the formula: S1 > S2 > S3 > S4 (30 septa). S1 most exsert septal cycle. S2 slightly less exsert than S1, being 3/4 the width of S1. S3–4 progressively less exsert. S3 dimorphic in size: those in half systems lacking S4 only ½ the width of S2, and those flanked by a pair of S4 attain the same width as S2. If present, S4 ½ the size of S3. All septa with vertical and slightly sinuous axial margin, S3–4 bearing dentate axial margins deeper on fossa. All septal faces with fine granulations. Fossa deep containing a rudimentary columella.

Distribution

Regional: Eastern margin of South Africa, off Durban; 424–430 m. Elsewhere: Vanuatu; Philippines (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); Tanzania (von Marenzeller 1904a; Cairns and Keller 1993); 182–790 m.

Remarks

The imagery records of Aulocyathus juvenescens represent a distributional range extension for this species further south of Tanzania and are also new records for South Africa. Although these specimens have no more than 30 septa, their morphology is consistent with the taxonomic diagnosis of the species detailed by von Marenzeller (1904a). Amongst the four recent species of the genus, A. juvenescens has the smallest CD, least number of septa, and a porcelaneous theca (Cairns 1999a).

Aulocyathus sp. cf. matricidus (Kent, 1871)

Fig. 1I, J

Flabellum matricidum Kent, 1871: 276, pl. 23: fig. 2A–C.

Fragilocyathus conotrochoides Yabe & Eguchi, 1932a: 388, 389, fig. 1. –Yabe and Eguchi 1941a: 101. –Yabe and Eguchi 1942b: 116, 145. pl. 9: fig. 15. –Eguchi 1965a: 288, 4 figs. –Eguchi and Miyawaki 1975: 57.

Aulocyathus cf. matricidus. –Yabe and Eguchi 1942b: 112, 116.

Aulocyathus matricidus. –Cairns 1999a: 104. –Cairns 1994: 60, pl. 26, figs C–G, pl. 42, figs B–D.

Type locality

Off Japan, 84 m (Zibrowius 1980; Cairns 1994).

Type material

Two syntypes are deposited at the NHM (Cairns 1994).

Material examined

ORI_DIa1 (2 specimens): no locality data.

Description

Corallum solitary, attached, and conical to elongate. Calice circular, calicular margin smooth. Largest of two specimens examined (ORI_DIa1) 5.2 × 5.1 mm in CD, and ≤ 15.4 mm in H. Costae wide. Theca and costae granular. Corallum light brown.

Septa hexamerally arranged in four cycles, the last cycle being incomplete, according to the formula: S1 > S2 > S3 > S4 (32 septa). S1–2 most exsert septa. S1 extend almost towards centre of fossa, with vertical to slightly sinuous axial margin. S2 ¼ less wide than S1, also has vertical axial margin, and being granular deeper in fossa. S3–4 progressively less exsert (if at all). S3 dimorphic in size: those half-systems lacking S4 only ½ the width of S2, while those flanked by S4 attain almost the same width of S2. If present, S4 ½ the width of S3. Axial margins of S3–4 dentate. Septal faces finely granular. Fossa deep containing a rudimentary columella.

Distribution

Regional: Eastern margin of South Africa; depth unknown. Elsewhere: Tsugara Strait; and Japan Sea (Cairns 1994); 84–207 m.

Remarks

Among all congeners, specimens examined closely resemble Aulocyathus matricidus in having a smooth calicular margin, upper peripheral septa not notched, rudimentary columella, and a slender pedicel. However, A. matricidus is only known from Japan and, therefore, its occurrence in the southwestern Indian Ocean would represent a disjunct distribution.

Caryophyllia Lamarck, 1816

Diagnosis

Corallum solitary, attached or free: if attached, corallum cylindrical, trochoid, or ceratoid; if free, corallum usually cornute. Calice circular, elliptical, or compressed; thecal margin spines present on species having compressed coralla. Septal symmetry variable, but hexameral symmetry with four cycles of septa most common. One crown of paliform lobes present before penultimate or rarely the antepenultimate cycle of septa. Columella fascicular, composed of several twisted laths. Exclusively azooxanthellate and common in deep water.

Caryophyllia (Acanthocyathus) Milne-Edwards & Haime, 1848

Diagnosis

Caryophyllia with thecal margin spines or crests.

Type species

Acanthocyathus grayi Milne-Edwards & Haime, 1848a, by subsequent designation (Milne-Edwards and Haime 1850b).

Caryophyllia (Acanthocyathus) dentata (Moseley, 1881)

Acanthocyathus sp. –Moseley 1876: 550.

Acanthocyathus dentatus Moseley, 1881: 143, pl. 2, fig. 7A–C.

Caryophyllia (Acanthocyathus) dentata. –Cairns and Zibrowius 1997: 98–99, fig. 8A–D. –Kitahara et al. 2010: 92, 112.

Type locality

Off Kandavu Islands, Fiji (HMS ‘Challenger’ stn. 174D: 19°05'50"S, 178°16'20"E); 384 m (Moseley 1881).

Type material

The holotype is deposited at the NHMUK (Kitahara et al. 2010).

Material examined

USNM 91540 (1 damaged specimen): Eastern margin, 23 km from Shaka’s Rock/off Tongati Estuary, 29°41'14.39"S, 31°21'10.80"E; 85 m.

Distribution

Regional: Eastern margin of South Africa, off Shaka’s Rock; 85 m. Elsewhere: Fiji (Moseley 1881); and Indonesia (Cairns and Zibrowius 1997); 90–384 m.

Remarks

The specimen reported herein is severely damaged and therefore tentatively added to the South African coral fauna- thus representing a new record for the region. Nonetheless, representatives from Indonesia have been well described by Cairns and Zibrowius (1997), who noted the discrepancies in the septal symmetry between the holotype (hexameral) and the specimens they examined (decameral). Apart from the symmetry, the same authors also pointed out a distinctively ridged C1–2 of their studied specimens in relation to the holotype.

Caryophyllia (Acanthocyathus) grayi (Moseley, 1881)

Fig. 1K, L

Acanthocyathus grayi Milne-Edwards & Haime, 1848a: 293, pl. 9, fig. 2. –Alcock 1898: 15. –van der Horst 1931: 6. –Umbgrove 1938: 264–265. –Umbgrove 1950: 641–642, pl. 81, figs 27–32. –Wells 1984: 209, pl. 2, figs 5–9. –Zou 1988: 76, figs 8–9.

Caryophyllia (Acanthocyathus) grayi . –Cairns 1994: 49, pl. 21, figs I–K. –Cairns and Zibrowius 1997: 97–98, figs 7C, F, I. –Cairns 1998: 377. –Cairns 1999a: 76. –Cairns 2004: 276.

Caryophyllia grayi. –Kitahara et al. 2010: 102, figs 53–55, 58–59.

Type locality

Unknown (Cairns and Zibrowius 1997; Kitahara et al. 2010).

Type material

One syntype is deposited at the NHMUK (Cairns and Zibrowius 1997; Kitahara et al. 2010).

Material examined

USNM 91541 (1 specimen): Eastern margin, 32 km south of Ponta Do Ouro/20 km off Kosi Bay Estuary, 27°08'10.79"S, 32°52'07.20"E; 98 m.

Description

Corallum ceratoid, curved, unattached, with a slender and slightly curved pedicel (PD:GCD = 0.09). Calice compressed (GCD:LCD = 1.6). Calicular margin serrate. Only specimen examined: 19.9 × 12.4 mm in CD, 1.8 mm in PD, and 22.1 mm in H. Thecal margins rounded, with three thecal spines. Costae rounded, equal in width, and extending towards pedicel. C1 more prominent than remaining costae. Intercostal striae narrow. Examined specimen eroded with a light brown appearance.

Septa in 15 sectors arranged in four cycles, the last cycle being incomplete, according to the formula: 15:15:30:8 (68 septa). Primary septa most exsert, and extend ¾ distance to columella, with straight to slightly sinuous axial margin. Higher cycle septa less exsert (if at all). Secondary septa ¼ the width of primaries, with sinuous axial margins. Tertiary septa ¾ less the width of secondaries also having sinuous axial margins. S4 rudimentary. A total of 15 (P3) with sinuous axial margins encircle an elongated fascicular columellar. Fossa of moderate depth.

Distribution

Regional: Eastern margin of South Africa, off Kosi Bay Estuary (32 km south of Ponta Do Ouro: Mozambique); 98 m. Elsewhere: Japan (Cairns 1994); Philippines; Indonesia (Cairns and Zibrowius 1997); Wallis and Futuna Islands; Vanuatu (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); Australia (Cairns 2004; Kitahara et al. 2010); Andamans Islands (van der Horst 1931); 37–490 m.

Remarks

Caryophyllia (Acanthocyathus) grayi differs from C. dentata, the only other Acanthocyathus from the region, in having 15 primary septa and in bearing spines on both thecal edges only on one side in C. dentata. Although C. grayi was previously reported from South Africa (Cairns and Zibrowius 1997; Cairns 1999a; Kitahara et al. 2010), none of the authors presented the locality information. Nonetheless, South African representatives of C. grayi are similar to the Australian specimens in S1 bearing a sinuous axial margin, whereas they are both different from the Japanese specimens in that regard. However, this South African specimen differs from both the Japanese and Australian representatives in its septa arranged in 15 sectors, with four pairs of S4 (15:15:30:8); instead of 14 or 18 sectors.

Caryophyllia (Caryophyllia) Lamarck, 1806

Diagnosis

Caryophyllia without thecal margin spines or crests.

Type species

Madrepora cyathus Ellis & Solander, 1786, by subsequent designation (Stokes and Broderip 1828).

Caryophyllia (Caryophyllia) cinticulata (Alcock, 1898)

Thecocyathus cinticulatus Alcock, 1898: 17–18, pl. 2, figs 5, 5A.

Trochocyathus cinticulatus. –Gardiner 1904: 99, 103–104, pl. 2, fig. 2. –Cairns et al. 1999: 24.

Caryophyllia cinticulata. –Kitahara et al. 2010a: 98, 113, figs 17–21.

Caryophyllia (Caryophyllia) cinticulata. –Kitahara and Cairns 2021: 465–466, 468, figs 252, 253A–F.

Type locality

Off the Maldives (HMS ‘Investigator’); 84 m (Alcock 1898; Kitahara et al. 2010).

Type material

The holotype is purportedly deposited at the IM (Kitahara et al. 2010).

Material examined

None.

Distribution

Regional: Eastern margin of South Africa, off East London; 59 m (Gardiner 1904). Elsewhere: Maldives (Alcock 1898); and New Caledonia (Kitahara et al. 2010; Kitahara and Cairns 2021); 282–384 m.

Remarks

Caryophyllia cinticulata representatives from New Caledonia are well described by Kitahara et al. (2010), in which these authors mention that it displays circumferential transverse ridges on the theca, and differs from its congeners by having decameral symmetry, adults attaining > 10 mm in GCD, S3 smaller than S2, and extremely sinuous S1 and S2 axial margins. No specimens of this species were found in the material examined and therefore the entry is based on Gardiner’s (1904) record, who reported the species off East London.

Caryophyllia (Caryophyllia) diomedeae von Marenzeller, 1904

Fig. 1M, N

Caryophyllia diomedeae von Marenzeller, 1904b: 79–80, pl. 1, fig. 2. –Durham and Barnard 1952: 10, 82, pl. 9, fig. 43. –Cairns 1991: 11–13, pl. 4, figs C–E. –Cairns 1995: 49–50, pl. 9, figs A–D. –Cortès 1997: 330. –Cairns and Zibrowius 1997: 88. –Koslow and Gowlett-Holmes 1998: 38. –Cairns 1999a: 74. –Cairns et al. 1999: 20. –Piñón 1999: 20, 81. –Cairns 2004: 264, 277, 328. –Cairns et al. 2005: 17, 25, 28, fig. 2D–E. –González-Romero et al. 2008: 1–2, fig. 1. –Kitahara et al. 2010: 100, 102, figs 37–46.

Caryophyllia profunda. –Cairns 1982: 17–19 (in part: ‘Eltanin’–1403).

Caryophyllia sarsiae . –Cairns and Parker 1992: 20, figs 5C, E, F.

Type locality

Off Panama (USS ‘Albatross’ stn. 3358: 6°30'N, 81°44'W); 1043 m (von Marenzeller 1904b).

Type material

One syntype is deposited at NHMUK (Kitahara et al. 2010).

Imagery data

MN_SM 85 (1 specimen): Eastern margin, 20 km off Cape Vidal/23 km off St Lucia Estuary, 27°59'30.00"S, 32°40'47.99"E; 550 m. MN_SM 129 (7 specimens): Eastern margin, 17 km off Margate/ km off Boboyi Estuary, 30°53'24.00"S, 30°31'41.99"E; 850 m. MN_SM 226 (1 specimen): Southern margin, 32 km off Mazeppa Bay/24 km off Kobole Estuary, 32°28'36.00"S, 28°58'48.00"E; 710–775 m.

Description

Corallum straight to slightly curved, and attached to substrate by a broad pedicel (PD:GCD ~ 0.5) that expands into an encrusting base. Largest imaged specimen (MN_SM 85) 13.0 × 12.0 mm in CD, 5.0 mm in PD, and 29.0 mm in H. Calice elliptical (GCD:LCD = 1.08–1.14), calicular margin jagged. Costae prominent at calicular margin, with shallow intercostal striae, sometimes bearing granules. Theca thick and porcelaneous. Corallum white to beige.

Septa hexamerally arranged in four cycles according to the formula: S1 ≥ S2 > S3 > S4 (48 septa). S1–2 most exsert, equal in width, and almost reach columella with vertical to slightly sinuous axial margins. S3–4 progressively narrower and less exsert. S3 ¾ the width of S1–2, each bearing a sinuous pali (12 P3), and consistently 1 mm in width. S4 slightly smaller than S3. Higher cycle septa with sinuous axial margin, especially S3. Septal faces somewhat smooth near calicular margin, but becoming granulated towards fossa. Fossa of moderate depth containing a fascicular columella, encircled by a P3 crown.

Distribution

Regional: Southern and Eastern margins of South Africa, off Mazeppa Bay extending towards Cape Vidal; 550–850 m. Elsewhere: Australia (Cairns and Parker 1992, 1998, 2004a; Kitahara et al. 2010); New Zealand (Cairns 1982, 1995); New Caledonia (Kitahara et al. 2010; Kitahara and Cairns 2021); Cocos and Galapagos Islands (Cairns 1991); Philippines; Indonesia (Cairns and Zibrowius 1997); Vanuatu (Cairns 1999a); Mediterranean to the Azores (Cairns et al 1999); Bermuda; Cook Island (Cairns 1995); Chile (Cairns et al. 2005); northern Pacific (González-Romero et al. 2008); 225–2200 m.

Remarks

Among congeners that are attached and display hexamerally arranged septa in four complete cycles, Caryophyllia (C.) diomedeae closely resembles C. sarsiae Zibrowius, 1974a (Cairns 1991). It is therefore no surprise that Cairns and Parker (1992) mistook their records of C. diomedeae for C. sarsiae, in which they mention the latter to occur in South Africa but did not provide the associated locality data for their South African records. Nonetheless, the imaged specimens (listed under imagery data) seem to have been mixed with C. sarsiae and have therefore been separated based on: (i) having a smooth theca and costae restricted to calicular margin (Cairns 1995), as compared with C. sarsiae which has costae prominent throughout corallum, and (ii) pali consistently being 1 mm in width as compared with C. sarsiae which bears pali being three times less the width of the pali bearing septa (key in Kitahara et al. 2010).

Caryophyllia (Caryophyllia) ephyala Alcock, 1891

Fig. 1O, P

Caryophyllia ephyala Alcock in Wood-Mason & Alcock, 1891: 6–2. –Alcock 1898: 13–14, pl. 1, fig. 4, 4A. –Gardiner 1904: 117–118. –Yabe and Eguchi 1932a: 388–389. –Cairns and Keller 1993: 219.

Type locality

Off the western margin of Andaman Sea, India (HMS ‘Investigator’ stn. 56); 439–402 m (Wood-Mason and Alcock 1891).

Type material

Types are presumably deposited at the IM.

Material examined

SAMC_A072974 (2 specimens): Western margin, 168 km off PATERNOSTER/173 km off Brak Estuary, 32°05'41.99"S, 16°19'47.99"E; 440 m.

Description

Corallum ceratoid, attached, tapering to a slightly curved and slender pedicel (PD:GCD = 0.20). Calice slightly elliptical (GCD:LCD = 1.1), with slightly lancet calicular margin. Largest specimen examined (SAMC_A072974) 11.2 × 9.9 mm in CD, 2.4 mm in PD, and 23.0 mm in H. Costae poorly developed, but C1–2 more prominent and double the width of C3–4. C3–4 equal in width. All costae prominent at calicular margin, disappearing towards base, and separated by shallow and thin intercostal furrows. Theca glistening, with faint costal ridges. Corallum white to light brown.

Septa hexamerally arranged in four complete cycles according to the formula: S1 ≥ S2 > S3 > S4 (48 septa). S1 highly exsert, and extend towards columella with slightly sinuous axial margins. S2 equal to or slightly less wide and exsert than S1, but otherwise similar in profile. Higher cycle septa progressively less exsert, but S4 more exsert than S3. S4 joining neighbouring S1–2 and forming a slightly lanceted calicular margin. S3 2/3 the width of S2, but have a more sinuous axial margin. Each S3 bears a thin and sinuous pali (12 P3). S4 rudimentary, with straight to slightly sinuous axial margins. All septa and pali covered in granules. Fossa of moderate depth, containing a fascicular columella composed of five or six ribbon-like elements.

Distribution

Regional: Western and eastern (Gardiner 1904) margin of South Africa, off Paternoster extending towards Buffalo River mouth (Gardiner 1904); 146–567 m. Elsewhere: Japan (Yabe and Eguchi 1932a); Andaman Sea (Alcock 1891); 146–1289 m (Cairns and Keller 1993).

Remarks

Caryophyllia (C.) ephyala belongs to the largest morphological group of the genus, having septa hexamerally arranged in four complete cycles, and when keyed out comes closest to C. huinayensis Cairns, Häussermann & Försterra, 2005. However, C. ephyala is distinguished from C. huinayensis in its calicular margin being slightly lanceted and having S3 larger than S4. Furthermore, these species differ in distributional range, of which C. ephyala is known from the Indian Ocean and C. huinayensis recorded in the South Pacific. Among the South African caryophylliids, C. ephyala superficially resembles C. scobinosa Alcock, 1902 in having a slightly lanceted calicular margin, poorly developed costae, and a thin pedicel, but can be differentiated by having a circular calice, corallum attached, and consistently having four complete septa cycles. This species was previously reported from South Africa, off Mossel Bay extending north of Richards Bay (Gardiner 1904 – specimens could not be traced). Thus, the new record reported herein extends its regional distribution to the South Atlantic, off Lamberts Bay.

Caryophyllia (Caryophyllia) grandis Gardiner & Waugh, 1938

Fig. 2A, B

Caryophyllia clavus. –von Marenzeller 1904a: 281 (in part: pl. 16, figs 9–9I).

Caryophyllia grandis Gardiner & Waugh, 1938: 177, pl. 1, fig. 2. –Zibrowius and Gili 1990: 32. –Cairns 1991: 12. –Cairns and Keller 1993: 234.–Cairns and Zibrowius 1997: 96, fig. 9G–H. –Cairns 1998: 376. –Cairns et al. 1999: 20. –Cairns 2004a: 277. –Kitahara et al. 2010: 102, figs 47–52.

Caryophyllia arcuata. –Boshoff 1980: 36.

Type locality

Off the west of Fadiffolu, Maldives (HEMS ‘Mabahiss’ stn. 145E: 4°58'42"N, 73°16'24"E); 494 m (Gardiner and Waugh 1938).

Type material

Four syntypes are deposited at the NHMUK (Cairns and Keller 1993; Kitahara et al. 2010).

Material examined

SAMC_A073150 (1 specimen): Southern margin, 26 km off Mazeppa Bay/33 km off Great Kei Estuary, 32°41'12.12"S, 28°43'54.12"E; 480–490m. SAM_H2813 (1 specimen): Eastern margin, 20 km off Cape Vidal/23 km off St Lucia Estuary, 27°59'30.00"S, 32°40'47.99"E; 550 m. ORI_DIIIa3 (1 specimen): Eastern margin, other locality information unknown. USNM 62497 (4 specimens): Eastern margin, 4 km from Shaka’s Rock/5 km off Mhlali Estuary, 29°29'23.46"S, 31°15'48.92"E; 183–220 m.

Description

Corallum large, trochoid, usually free, and curved with a narrow pedicel (GCD:PD = 0.08–0.13) Calice elliptical (GCD:LCD = 1.41–1.75), calicular margin jagged. Largest specimen examined (ORI_DIIIa3) 35.0 × 20.0 mm in CD, 3.0 mm in PD, and 30.0 mm in H. C1–3 slightly ridged or absent. Upper theca and septal faces light beige, with white or discoloured lower theca.

Septa hexamerally arranged in five complete cycles according to the formula: S1–3 > S4 > S5 (96 septa). S1–3 highly exsert, with straight to slightly sinuous axial margins. S4 less exsert than S1–3, each bearing a paliform lobe (24 P4). S5 more exsert than S4, and fuse to adjacent septa at the calicular margin forming lancets. Septal faces and pali bear granules arranged parallel to septal margin. Pali thin, with sinuous axial margin, usually narrower than S4, and forming crown encircling columella. Fossa moderately deep, composed of a fascicular columella composed of 6–8 broad twisted elements.

Distribution

Regional: Southern to eastern margin of South Africa, off Mazeppa Bay extending towards off Cape Vidal; 183–550 m. Elsewhere: Australia (Cairns 1998, 2004); New Caledonia (Kitahara et al. 2010); Indonesia (Cairns and Zibrowius 1997); Sumatra (von Marenzeller, 1904); Mozambique (Cairns and Keller 1993); and Maldives (Gardiner and Waugh 1938); 183–595 m.

Remarks

Caryophyllia (C.) grandis closely resembles C. ambriosa Alcock, 1898, for which similarities exist in the number of septa (< 96 septa), corallum shape, and substrate relationship (both unattached). However, C. grandis is distinguished by its narrower pedicel and in having S4 wider than S5. Caryophyllia grandis also has a brownish theca (Cairns and Keller 1993), as compared with C. ambriosa, which is uniformly white. Caryophyllia grandis was first recorded from South African (off the Southern margin, off Knysna, 500 m) by von Marenzeller (1904a), who reported several Caryophyllia under the name C. clavus Scacchi, 1835 (C. grandis being one of them). Subsequently, Boshoff (1981) identified C. grandis as C. arcuata (Milne-Edwards & Haime, 1848a) in his annotated checklist of the southern Africa fauna. Thus, Zibrowius and Gili (1990) represent the first historically reliable record of C. grandis in the South African region, and authors mention that this species is represented in various localities off Natal but did not list the South African material examined. Cairns and Keller (1993) built on Zibrowius and Gili’s (1990) also reported C. grandis in Mozambique. The new records of C. grandis (SAM_H2813 and SAMC_A073150) are within the previously reported localities (von Marenzeller 1904a a; Zibrowius and Gili 1990; Cairns and Keller 1993).

Figure 2. 

A, B Caryophyllia (Caryophyllia) grandis (SAMC_A073150, off Mazeppa Bay, 480–490m) A calicular view B lateral view C, D Caryophyllia (Caryophyllia) lamellifera (SAMC_A090155, off Sodwana Bay, 120 m) C calicular view D lateral view E, F Caryophyllia (Caryophyllia) profunda (DTE Valdivia Stn 108, off Agulhas, 126 m) E calicular view F lateral view G, H Caryophyllia (Caryophyllia) quadragenaria (USNM 91539, Eastern coast, off Shaka’s Rock, 350 m) G calicular view H lateral view I, J Caryophyllia (Caryophyllia) rugosa (SAMC_A090071, off Kosi-Kumpungwini (Sifungwe) Estuary, 370 m) I calicular view J lateral view K, N Caryophyllia (Caryophyllia) sarsiae K, L (MN_SM226, off Mazeppa Bay, 710–775 m) K calicular view L lateral view M, N (MN_SM174, off Kidds Beach, 760 m) M calicular view N lateral view O, P Caryophyllia (Caryophyllia) scobinosa (SAM_H1248, off Durban, 91m) O calicular view P lateral view. Scale bars: 10 mm.

Caryophyllia (Caryophyllia) lamellifera Moseley, 1881

Fig. 2C, D

Caryophyllia lamellifera Moseley, 1881: 140–141, pl. 1, fig. 7A, B. –Hutton 1904: 315. –Cairns 1991: 12. –Cairns 1995: 51–52, pl. 9, fig. I, pl. 10, figs A–C. –Cairns and Zibrowius 1997: 90. –Cairns 1999a: 74–75. –Cairns et al. 1999: 20. –Cairns 2004: 278. –Kitahara et al. 2010: 104, 105, 113, 115, figs 64–68.

Caryophyllia (Caryophyllia) lamellifera. –Kitahara and Cairns 2021: 483–485, figs 263F–G, 264.

Type locality

Kermadec Ridge (HMS ‘Challenger’ stn. 170: 29°55'S, 178°14'W); 1152 m (Moseley 1881; Kitahara et al. 2010).

Type material

Two uncatalogued syntypes are deposited at the NHMUK (Kitahara et al. 2010).

Material examined

SAMC_A090155 (1 specimen): Eastern margin, Sodwana, 120 m.

Description

Corallum trochoid and attached to substrate by a broad pedicel (PD:GCD = 1.5) that expands into an encrusting base. Specimen examined 12.8 × 10.7 mm in CD, 32.5 mm in H, and displays a case of rejuvenescence. Calice elliptical (GCD:LCD = 1.27), with a jagged calicular margin. Costae poorly developed, but prominent near calicular margin and progressively disappearing towards base. Theca covered with thin transverse ridges. Corallum mostly white, but brownish purple near calicular margin.

Septa hexamerally arranged in five cycles, the last incomplete, according to the formula: S1–2 > S3 > S4 > S5 (52 total septa). S1–2 equal in width and most exsert septa, with straight to slightly sinuous axial margins. S3–4 progressively less exsert, with sinuous axial margins, S3 being most sinuous. S3 1/3 less wide than S1–2, bearing a thick and sinuous pali (13 P3). S4 dimorphic in size: half systems with S5 the S4 neighbouring S1 being 1/3 less wide than S3, and S4 flanked by S5 being the same width as S3; whilst in half systems without S5 S4 is 1/5 less wide than S3. S5 slightly less wide than S4. Septal and palar faces covered with granules arranged in a perpendicular manner. Fossa moderately deep, containing a fascicular columella composed of 12 twisted elements.

Distribution

Regional: Eastern margin of South Africa, off Sodwana; 120 m. Elsewhere: Australia (Cairns 1995, 2004a); New Zealand (Cairns 1995); New Caledonia (Kitahara et al. 2010; Kitahara and Cairns 2021); Philippines; Indonesia (Cairns and Zibrowius 1997); Wallis and Futuna Islands; and Vanuatu (Cairns 1999a); 89–1152 m.

Remarks

As noted by Kitahara et al. (2010), Caryophyllia (C.) lamellifera differs from the other five congeners (C. rugosa Moseley, 1880, C. corrugata Cairns, 1979, C. cinticulata (Alcock, 1898), C. versicolorata Kitahara, Cairns & Miller, 2010a, and C. aspera Kitahara, Cairns & Miller, 2010a) that have theca covered with transverse ridges, by its septal arrangement and profile. Although C. lamellifera resembles C. corrugata and C. aspera in having hexamerally arranged septa, it differs from these two species in having S1 = S2 as compared with S1 > S2 as in C. corrugata and C. aspera. Septal exsertness is another distinguishing feature, particularly when comparing C. lamellifera with C. versicolorata, for which the former displays a highly exsert S1 (2 mm) as compared with 1.5 mm in C. versicolorata. Furthermore, all septa of C. lamellifera have sinuous axial margins while the S1 of C. aspera are straight. Caryophyllia lamellifera differs from the other two South African congeners (C. rugosa and C. cinticulata) in having septa hexamerally arranged in five incomplete cycles in contrast to septa octamerally (may sometimes be decamerally) arranged in three cycles in C. rugosa, and decamerally arranged in three cycles in C. cinticulata. The only examined South African specimen of C. lamellifera varies from those from New Caledonia in having 13 pali instead of 12, having septa arranged in five incomplete cycles (6:6:12:24:4), and in the dimorphic size shown by S4. This account represents a new record for South Africa and extends the previous Pacific distribution of C. lamellifera into the Indian Ocean.

Caryophyllia (Caryophyllia) profunda Moseley, 1881

Fig. 2E, F

Caryophyllia profunda Moseley, 1881: 138–139 pl. 1, figs 6, 6b.–von Marenzeller 1904a, 298. –Gardiner 1913: 688–689. –Gardiner 1929: 126. –Gardiner 1939: 331.–Ralph 1948: 108, fig. 2. –Squires 1964, pl. 11. –Squires 1969: 16–17, pl. 6, map 1. –Ralph and Squires 1962: 3, 6–7, pl. 1, figs 8–11. –Squires and Keyes 1967: 15, 17, 23, pl. 2, figs 1–4. –Zibrowius 1974b: 751–755, pl. 1, figs 1–10. –Cairns 1979: 206. –Cairns 1982: 17–19, pl. 5, figs 1–5. –Zibrowius and Gili 1990: 25–26, pl. 4, figs L–R. –Cairns and Keller 1993: 235–236. –Cairns 1995: 44–45, pl. 7, fig. I, pl.8, figs A–C. –Kitahara et al. 2010: 114.

Caryophyllia cyathus.–von Marenzeller 1904a: 295, pl. 16, figs 6, 6A.–Hoffmeister 1933: 14, pl. 4, figs 4–5.–Gardiner 1939: 330–331.–Squires 1961: 7.

Caryophyllia planilamellata . –Dennant 1906: 157–158, pl. 6, figs 4A, B. –Squires 1961: 18.

Caryophyllia clavus. –Wells 1958: 265, pl. 1, figs 12, 13.

Caryophyllia cf. C. maculata. –Ralph 1948: 108, fig. 2. –Ralph and Squires 1962: 3, 7, pl. 2, figs 1–2. –Squires and Keyes 1967: 15, 17, 23, pl. 2, figs 4, 5.

Type locality

Off Nightingale Island, Sub-Antarctica (HMS ‘Challenger’ stn. 135: 37°0'50"S, 12°19'10"W); 183–274 m (Moseley 1881).

Type material

Twenty syntypes are deposited at the NHMUK (Cairns 1982).

Material examined

None.

Imagery data

DTE Valdivia Stn. 108 (1 specimen): Southern margin, 60 km off AGULHAS/69 km off De Mond-Heuningnes Estuary, 35°19'18.00"S, 20°15'17.99"E; 126 m.

Description

Adapted from Cairns (1982) and von Marenzeller (1904a): Corallum trochoid, slightly curved, and attached to substrate by a broad pedicel. Calice slightly elliptical (GCD:LCD = 1.04), calicular margins slightly serrate. Imaged specimen 25.0 × 24.0 mm in CD and 54.0 mm in H. Costae flat, more prominent near calicular margin, and equal in width. C1–3 sometimes slightly ridged. Theca thick, porcelaneous, and finely granulated. Corallum white with brown theca.

Septa hexamerally arranged in five complete cycles according to the formula: S1–2 > S3 > S4 > S5 (96 septa). S1–2 moderately exsert, equal in width, and bearing axial margins that join columella deep in fossa. Higher cycle septa progressively less exsert. S3 1/5 less wide than S1–2. S4 ~ ¾ the width of S3, each bearing a broad pali (24 P4), with vertical inner margin. S5 ¾ the width of S4. All septal faces slightly granulated and septal margins straight, except for S4 which may be slightly sinuous. Fossa moderately deep containing a fascicular columella encircled by a paliform crown.

Distribution

Regional: Southern margin of South Africa, off Agulhas; 126 m. Elsewhere: St Paul and Amsterdam Islands (Zibrowius 1974b); Australia (Hoffmeister 1933); New Zealand (Ralph and Squires 1962; Squires and Keyes 1967; Cairns 1995); Sub-Antarctic Islands (Moseley 1881; von Marenzeller 1904a; Zibrowius 1974b; Cairns 1982); 35–1116 m.

Remarks

Caryophyllia (C.) profunda is one of the several Caryophyllia species reported under the name of Caryophyllia cyathus (Ellis & Solander, 1786) by von Marenzeller (1904a), a long-standing confusion that Zibrowius (1974b) discusses in his account of the corals of the St Paul and Amsterdam Islands. Caryophyllia profunda differs from C. cyathus in its S1 being straight as compared with slightly sinuous in the latter, and having 24–25 instead of 20 P4. Nonetheless, von Marenzeller’s (1904a) records form the basis of the occurrence of this species in South African territory, as no other known South African samples have been reported subsequently.

Caryophyllia (Caryophyllia) quadragenaria Alcock, 1902

Fig. 2G, H

Caryophyllia quadragenaria Alcock, 1902a: 91–92. –Alcock 1902b: 10, pl. 1, figs 4, 4A. –Keller 1981: 18. –Cairns 1991: 12. –Cairns 1994: 46–47, pl. 20, figs C–H, pl. 51, figs C–D. –Cairns 1995: 45–46, pl. 7, figs G–H. –Cairns and Zibrowius 1997: 88, 93. –Cairns 1998: 375, –Cairns 1999a: 73. –Cairns et al. 1999: 20. –Cairns 2004: 278. –Gonźalez-Romero et al. 2008: 1–2, fig. 2. –Kitahara et al. 2010a: 107, figs 78–81.

Caryophyllia scobinosa. –Yabe and Eguchi 1942b: 119 (in part).

Caryophyllia scobinosa decapali Yabe & Eguchi, 1942b: 120, 149, pl. 10, figs 6–7. –Eguchi 1968: C33–34. –Eguchi and Miyawaki 1975: 56. –Cairns 1991: 12.

Caryophyllia profunda. –Squires and Keyes 1967: 23 (in part).

Caryophyllia decapali . –Grygier 1983: 420. –Zibrowius and Grygier 1985: 120, figs 10–11.

Caryophyllia (Caryophyllia) quadragenaria. –Kitahara and Cairns 2021:490–491, 493, figs 268, 269A–C.

Type locality

Off Makassar Strait, Banda and Timor seas (Indonesia) (HMS ‘Siboga’ stns 90, 251, 289: 1°17'50"N, 12°19'10"W; 5°28'40"S, 132°02'00"W; and 9°0'30"S, 122°24'5"W respectively); 54–281 m (Alcock 1902a).

Type material

Two syntypes are deposited at the ZMA (Cairns 1995; Cairns and Zibrowius 1997; Kitahara et al. 2010a).

Material examined

USNM 91538 (3 damaged specimens): Eastern margin, 69 km off Durban/68 km off Beachwood Mangroves, 30°08'59.99"S, 31°37'12.00"E; 930 m. USNM 91539 (1 specimen): Eastern margin, 44 km from Shaka’s Rock/45 km off Mhlali Estuary, 29°41'59.99"S, 31°37'47.99"E; 350 m.

Description

Corallum small, ceratoid to subcylindrical, slightly curved, and attached to substrate by a robust but narrow pedicel (PD:GCD = 0.3). Calice circular to slightly elliptical (GCD:LCD = 1.1), with a slightly lanceted calicular margin. Largest examined specimen (USNM 91539) 8.2 × 7.4 mm, 9.3 mm in H, and 2.5 mm in PD. Costa equal in width, separated by moderately deep furrows that fade towards base. All costae covered with low granules. Theca thick. Corallum white to light brown.

Septa decamerally arranged in three cycles according to the formula: S1 > S2 > S3 (40 septa). S1 highly exsert and almost meet columella with straight to slightly sinuous axial margins. S2 ~ ¾ the width of S1, least exsert, but bear the most sinuous axial margins. Each S2 bears a tall (extending above columella elements) and sinuous pali (10 P2). S3 slightly less wide than S2 and bearing a less sinuous axial margin. All septal faces granular. Fossa moderately deep, with a fascicular columella composed of 6–8 ribbon-like elements.

Distribution

Regional: Eastern margin of South Africa, off Durban extending towards off Shaka’s Rock; 350–930 m. Elsewhere: Japan (Cairns 1994); South China Sea (Cairns and Zibrowius 1997); Indonesia (Alcock 1902a; Cairns and Zibrowius 1997); New Caledonia (Kitahara et al. 2010a; Kitahara and Cairns 2021); Vanuatu, Wallis, and Futuna (Cairns 1999a); New Zealand (Cairns 1995); Australia (Cairns 1998); North Pacific (Gonźalez-Romero et al. 2008); 54–1669 m.

Remarks

Among the Caryophyllia that have decameral septal symmetry, Caryophyllia (C.) quadragenaria most closely resembles C. perculta Cairns, 1991, but can be differentiated by having smooth or slightly granular instead of extremely granular or carinate septal faces . C. quadragenaria may also be mistaken for C. cinticulata (Alcock, 1898) and C. rugosa Moseley, 1881 but may be differentiated from these two South African congeners by lacking circumferential thecal transverse ridges reported. The specimens reported herein represent new records for the Indian Ocean.

Caryophyllia (Caryophyllia) rugosa Moseley, 1881

Fig. 2I, J

Caryophyllia rugosa Moseley, 1881: 141–143, pl. 1, fig. 8. –Wells 1954: 469, pl. 177, figs 5, 6. –Cairns 1984: 11–13, pl. 2, figs A, B, pl. 4, fig. I. –Cairns 1991: 20. –Cairns and Keller 1993: 236, fig. 3I. –Kitahara et al. 2010a: 108, figs 93–97.

Caryophyllia paraoctopali Yabe & Eguchi, 1942b: 120, 150, pl. 10, fig. 12.

Caryophylliia (Caryophyllia) rugosa . –Cairns 1994: 47, pl. 20, fig. I, pl. 21, fig. A. –Cairns and Zibrowius 1997: 91–92. –Cairns 1998: 375. –Kitahara and Cairns 2021: 495–496, 498, figs 269H–I, 271, 272A–C.

Type locality

Off Banda and Sulu Seas, Indonesia (HMS ‘Challenger’ stns 192 and 201); 187–230 m (Moseley 1881).

Type material

Syntypes are deposited at the BMNH (Cairns 1994).

Material examined

SAMC_A073073 (1 specimen): Eastern margin, 29 km off Richards Bay/20 km off Nhlabane Estuary, 28°44'23.99"S, 32°23'12.11"E; 320–340 m. SAMC_A073180 (1 specimen): Southern margin, 33 km from Mazeppa Bay/24 km off Cwili Estuary, 32°45'47.88"S, 28°36'24.12"E; 240–250 m. SAMC_A090071(1 specimen): Eastern margin, 15 km south of Ponta Do Ouro/17 km off Kosi-Kumpungwini (Sifungwe) Estuary, 26°55'30.00"S, 33°01'05.99"E; 370 m. SAMC_A090077 (1 specimen): Eastern margin, 34 km off Port Dunford/38 km off Kosi-Kumpungwini (Sifungwe) Estuary, 29°10'00.00"S, 32°04'59.99"E; 170 m. USNM 91529 (1 specimen): Eastern margin, 37 km south of Ponta Do Ouro/23 km off Kosi Bay Estuary, 27°11'05.99"S, 32°50'53.88"E; 100 m.

Description

Corallum small, ceratoid to trochoid, and attached through a robust pedicel (PD:GCD = 0.3–0.8) that expands into a large encrusting base. Calice circular to elliptical (GCD:LCD = 1.0–1.2), with serrated calicular margin. Largest examined specimen (USNM 91529) 6.9 × 5.8 mm in CD, 2.4 mm in PD, and 12.5 mm in H. Costae covered with well-defined transverse ridges; ridges split and re-join around corallum. Corallum white to light brown.

Septa octamerally arranged in three cycles according to the formula: S1 > S2 > S3 (32 septa). However, one specimen (SAMC_A090071) has decamerally arranged septa (40 septa). S1 highly exsert, extending almost to columella. S2–3 becoming progressively narrower and less exsert. All septa bear sinuous axial margins, sinuosity being extreme in S1–2. S2 bearing a highly sinuous and thick pali (8 P2). Fossa shallow containing a prominent fascicular columella.

Distribution

Regional: Eastern margin of South Africa, off Kei Mouth extending towards Kosi Bay Estuary, (15 km south of Ponta Do Ouro: Mozambique); 100–370 m. Elsewhere: New Caledonia (Kitahara et al. 2010a; Kitahara and Cairns 2021); Hawaii (Cairns 1984); Japan; East China Sea (Cairns 1994); Philippines; Indonesia (Cairns and Zibrowius 1997); Wallis and Futuna Islands; Vanuatu (Cairns 1999a); Australia (Cairns 1998, 2004; Kitahara et al. 2010); New Zealand (Cairns 1995); Mozambique; Kenya; and Maldives (Cairns and Keller 1993); 71–581 m.

Remarks

Kitahara et al. (2010) noted that Caryophyllia (C.) rugosa is easily distinguished from congeners by the presence of transverse ridges and its extremely sinuous septa and pali. Although some specimens have a hexameral or decameral symmetry, the most common septal symmetry is octameral (Kitahara et al. 2010). South African specimens have both septal symmetries (octameral and decameral), for which the specimen (SAMC_A090071) that have decamerally arranged septa also displays rejuvenescence of the corallum. Caryophyllia rugosa was previously reported from South African waters by Cairns and Keller (1993) off Lake Kosi (in KwaZulu-Natal), and the examined specimens herein extend its regional distribution further north towards the Mozambican border.

Caryophyllia (Caryophyllia) sarsiae Zibrowuis, 1974

Fig. 2K–N

Caryophyllia cylindracea . –Jourdan 1895: 11 (in part).

Balanophyllia cornu . –Jourdan 1895: 27, pl. 2, fig. 21A, B.

Caryophyllia clavus. –Gravier 1920: 16 (in part).

Caryophyllia arcuata. –Gravier 1920: 25 (in part), pl. 2, figs 26, 29.

Caryophyllia sarsiae Zibrowius, 1974b: 779–782, pl. 3, figs A–F. –Zibrowius et al. 1975: 95, fig. 3A–E. –Zibrowius 1980: 62–63, pl. 24, figs A–J.

Caryophyllia sp. –Zibrowius 1974c: 755–756, pl. 1, fig. 11, pl. 2, fig. 1.

Type locality

Southeast of Le Chapelle Bank, west of Brittany (RV ‘Sarsia’ stn. 1: 47°19'00"N, 06°36'00"W); 880–980 m (Zibrowius 1974b).

Type material

Types are deposited at the NHMUK (Zibrowius 1974b).

Material examined

MN_SM 162 (2 specimen): Southern margin, 40 km off Kei Mouth/29 km off Cwili Estuary, 32°55'00.00"S, 28°31'00.00"E; 630 m. MN_SM 174 (1 specimen): Southern margin, 26 km off Kidds Beach/27 km off Ncera Estuary, 33°19'36.00"S, 27°52'23.99"E; 760 m. MN_SM 226 (19 specimens): Southern margin, 32 km off Mazeppa Bay/24 km off Kobole Estuary, 32°28'36.00"S, 28°58'48.00"E; 710–775 m.

Description

Corallum ceratoid, straight to slightly curved, and attached to substrate through a robust pedicel (PD:GCD < 0.60). Calice circular to slightly elliptical (GCD: LCD = 1.0–1.1), with serrated calicular margin. Largest imaged specimen (MN_SM 226) 15.0 × 15.0 mm in CD, 8.0 mm in PD, and 42.0 mm in H. Costae equal, flat, and smooth. Theca granular. Corallum white, with beige theca.

Septa hexamerally arranged in four cycles according to the formula: S1–2 > S3 ≥ S4 (48 septa). S1–2 equal in width, highly exsert, thick, and extend to columella with straight (although S1 may be slightly sinuous in some specimens) axial margins. S3 ¾ the width of S1–2, less exsert, with extremely sinuous axial margins, which bear narrow and sinuous pali (12 P3). S4 equal to slightly smaller than S3, but equally exsert, bearing straight axial margins. All septal faces appearing smooth, with small and randomly arranged granules deeper in fossa. Fossa of moderate depth, with fascicular columella composed of 6–9 twisted laths.

Distribution

Regional: Southern margin of South Africa, off Kei Mouth extending towards Kidds Beach; 630–760 m. Elsewhere: Madeira and Azores, extending to off Portugal and Ireland (Zibrowius 1974b); the Mediterranean (Zibrowius 1980); 520–2200 m.

Remarks

All the imaged specimens represented here have: (i) septa hexamerally arranged in four cycles; (ii) S1–3 bearing sinuous axial margins, S3 being the most extreme; and (ii) S4 having straight axial margins. Further to the septa symmetry and profile, specimens have granulated costae throughout corallum. Zibrowius (1974b) noted the intraspecific variation that may exist in the corallum shape of C. sarsiae, which is highly dependent on the environmental conditions, particularly in relation to substrate attachment. Some of the examined specimens (for example MN_SM 229) are attached to Solenosmilia variabilis Duncan, 1873, and do indeed display a more curved corallum. This account of C. sarsiae in South African territory extends the species distributional range further south and, therefore, represents a new record for the region.

Caryophyllia (Caryophyllia) scobinosa Alcock, 1902

Fig. 2O, P

Caryophyllia scobinosa Alcock, 1902a: 90. –Alcock 1902b: 8, pl. 1, figs 2, 2A. –Gardiner and Waugh 1938: 177–178. –Cairns 1991: 12. –Cairns and Keller 1993: 235. –Cairns 1994: 45–46, pl. 20, figs A, B (in part). –Cairns 1995: 52–53, pl. 10, figs G–I, pl. 11, figs A–C. –Cairns and Zibrowius 1997: 94. –Cairns 1999a: 75. –Cairns et al. 1999: 20. –Kitahara 2007: 498, 507, 510, fig. 2K. –Kitahara et al. 2008: 16, fig. 2D. –Cairns 2004: 278. –Kitahara et al 2010a: 109, figs 113,117.

Caryophyllia cultrifera Alcock, 1902b: 7–8, figs 1, 1A. –Faustino 1927: 67–68, pl. 8, figs 8, 9. –Veron 1986: 905.

Caryophyllia clavus. –von Marenzeller 1904a: 281 (in part ‘Valdivia’–246), pl. 16, figs 9C–G.

Type locality

Off Flores and Sulu Seas (HMS ‘Siboga’ stns. 45 and 102: 7°24'00"S, 118°15'20"E and 6°04'10"N, 120°44'00"E, respectively); 535–794 m (Alcock 1902a).

Type material

Six syntypes are deposited at the ZMA (Cairns 1994; Kitahara et al. 2010).

Material examined

SAM_H1248 (1 specimen): Eastern margin, 17 km from DURBAN/8 km off Mdloti Estuary, 29°42'34.21"S, 31°05'50.82"E; 91 m.

Description

Corallum ceratoid, unattached, and curved to a narrow pedicel (PD:GCD = 0.2). Calice slightly elliptical (GCD:LCD = 1.1), with jagged calicular margin. Specimen examined 9.5 × 8.6 mm in CD, 2.0 mm in PD, and 11.3 mm in H. Costae granular, flat, and separated by narrow intercostal furrows that fade towards base. Corallum white.

Septa hexamerally arranged in four cycles according to the formula: S1–2 > S3 > S4 (48 septa). S1–2 thick, slightly exsert, and extend to columella deep in fossa with straight to slightly sinuous axial margins. S2–3 progressively less exsert. S3 axial margin more sinuous than remaining septa. S3 ~ 2/3 the width to S1–2, each bearing a sinuous pali (12 P3). S4 rudimentary, but joining adjacent S1 and S2 at calicular margin to form rectangular lancets. All septal faces bear small pointed granules. Fossa relatively shallow containing a fascicular columella composed of five ribbon-like elements.

Distribution

Regional: Eastern margin of South Africa, off Durban; 91 m. Elsewhere: Brazil (Kitahara 2007); New Zealand (Cairns 1995); New Caledonia (Kitahara et al. 2010; Kitahara and Cairns 2021); Australia (Cairns 1995, 2004; Kitahara et al. 2010); Tanzania (von Marenzeller 1904; Gardiner and Waugh 1938); Madagascar Plateau; Walter Shoal (Cairns and Keller 1993); Sulu Sea; Celebes Sea (Alcock 1902a); off Tonga and Samoa (Cairns 1995); Philippines; Indonesia (Cairns and Zibrowius 1997); Wallis and Futuna Islands; and Vanuatu (Cairns 1999a); 253–2450 m.

Remarks

As noted by Kitahara et al. (2010), Caryophyllia (C.) scobinosa can be distinguished from other unattached Indo-Pacific Caryophyllia in having 48–72 septa, 12–14 pali, and a jagged calicular margin. The South African representative of C. scobinosa superficially resembles C. stellula Cairns, 1998 but can be distinguished by its S4 being more exsert than S3, and also by joining neighbouring S1 and S2 at the calicular margin forming triangular lancets. Corallum size and density also differentiate the adult forms of these two species. Although C. scobinosa has been previously reported in neighbouring areas (Cairns and Keller 1993), the current study confirms the occurrence of this species further south in the South African territory.

Caryophyllia (Caryophyllia) stellula Cairns, 1998

Fig. 3A, B

Caryophyllia epithecata. –Gardiner 1904: 114–117, pl. 1, figs 3 A–C (in part: localities I–V). –Cairns and Keller 1993: 219.

Caryophyllia (Caryophyllia) stellula Cairns, 1998: 375–376, fig. 2A–C. –Cairns 2004a: 278.

Type locality

Off west of Rottnest Island, Australia (RV ‘Diamantina’ stn. 25: 31°48'00.0"S, 114°58'12.0"E); 402 m (Cairns 1998).

Type material

The holotype is deposited at the WAM (Cairns 1998).

Material examined

DEFF_SVMEC–INV190 (11 specimen): Southern margin, 54 km from Cape Point/56 km off Buffels Wes Estuary, 34°43'48.62"S, 18°07'14.02"E; 386–392m. DEFF/SAEON_A32776 (1 specimen): Southern margin, 200 km from KNYSNA/210 km off Ratels Estuary, 34°43'48.62"S, 18°07'14.02"E; 636 m. SAMC_A073140 (1 specimen): Eastern margin, 6 km off Cape Vidal/17 km off St Lucia Estuary, 28°08'17.88"S, 32°36'54.00"E; 200 m. SAMC_A088909 (8 specimens): Southern margin, 140 km off Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445 m. SAMC_A088922 (3 specimens): Southern margin, 140 km off Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A088923 (4 specimens): Southern margin, 140 km off Agulhas/144 km Ratels off Bulura Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A088928 (2 specimens): Southern margin, 65 km off Cape St. Francis/70 km off Slang Estuary, 34°47'05.01"S, 24°45'42.30"E; 392 m. SAMC_A090127 (1 specimen): Southern margin, 240 km off Agulhas/247 km off De Mond-Heuningnes Estuary, 36°45'34.13"S, 21°12'46.61"E; 513 m. SAMC_A090145 (1 specimen): Southern margin, 116 km off Knysna/off Goukamma Estuary, 35°07'11.27"S, 23°02'41.91"E; 333 m. SAM_H1378 (2 specimens): Southern margin: 11 km off East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H1394 (25 specimens): No locality data. SAM_H1396 (2 specimens): Western margin, 14 km off Saldanha/31 km off Berg River I Floodplain Estuary, 33°06'29.99"S, 18°01'59.99"E; 347 m. SAM_H1418 (8 specimens): Western margin, 1 km off Cape Point/8 km off Buffels Wes Estuary, 34°21'42.64"S, 18°30'12.06"E; 549 m. SAM_H1421 (2 specimens): Western margin1 km off Cape Point/8 km off Buffels Wes Estuary, 34°21'42.64"S, 18°30'12.06"E; 567–1024 m. SAM_H1433 (16 specimens): Western margin, 1 km off Cape Point/8 km off Buffels Wes Estuary, 34°21'42.64"S, 18°30'12.06"E; 574–732 m. SAM_H1436 (44 specimens): Southern margin, 2 km off Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 165–183 m. SAM_H1448 (92 specimens): Southern margin, 241 km off Agulhas/247 km off De Mond-Heuningnes Estuary, 36°40'00.00"S, 21°25'59.99"E; 200 m. SAM_H1485 (72 specimens): Southern margin, 241 km off Agulhas/247 km off De Mond-Heuningnes Estuary, 36°40'00.00"S, 21°25'59.99"E; 200 m. SAM_H3056 (21 specimen): Southern margin, 2 km off Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 229 m. SAM_H3059 (2 specimens): South Africa, no other locality data.

Imagery data

BMNH 1939.7.20.249–251 (2 specimens): South Africa, locality data unknown. RV ‘Galathea’ stn. 202 (3 specimens): Eastern margin, KwaZulu-Natal, depth unknown.

Description

Corallum ceratoid, unattached, curved, having a slender pedicel (PD:GCD = 0.1–0.2). Calice slightly elliptical (GCD:LCD = 1.1), with smooth to slightly serrated calicular margin. Largest specimen examined (SAM_H1485) 11.4 × 10.2 mm in CD, 1.8 mm in PD, and 22.4 mm in H. Costae poorly developed, slightly granular, and separated by shallow intercostal furrows. Costae extending towards base. Theca thick. Corallum white, with yellowish brown theca.

Septa hexamerally arranged in four cycles according to the formula: S1–2 > S3 > S4 (48 septa). S1–2 slightly exsert and extend to columella. S3 2/3 the width of S1–2, and bear a sinuous pali (12 P3). S4 ¼ the width of S3. All septal have sinuous axial margins, S3 being the most sinuous. Higher cycle septa (S3–4) progressively less exsert (if at all). Fossa relatively deep, with a fascicular columella composed of 4–6 ribbon-like elements.

Distribution

Regional: Western to eastern margin of South Africa, off Saldanha extending towards Cape Vidal; 200–567 m. Elsewhere: Western Australia (Cairns 1998); 240–402 m.

Remarks

Of the Caryophyllia reported from the South African territory, C. stellula may be confused with C. scobinosa, but differs in having a more robust corallum, a thicker calicular margin, and different pattern of septal exsertness. Caryophyllia (C.) stellula was first reported from South Africa as C. epithecata (Gardiner 1904) who elevated Duncan’s (1873) C. clavus var. epithecata to species level. However, this goes against the International Code of Zoological Nomenclature (1985: article 45g) and, therefore, the correct author of C. epithecata is Duncan (1873). Despite this, Duncan’s C. clavus var. epithecata is a junior synonym of C. smithi (Zibrowius 1974c). In South Africa, C. stellula shows a broad regional distribution and the several specimens examined herein enabled the observation of intraspecific morphological variation. Some representatives have curved corallum with a narrow pedicel while others display a more robust and thicker pedicel, features also illustrated by Gardiner (1904). Nonetheless, the species consistently have septa hexameral symmetry in four complete cycles, and a total of 12 well-defined P3, each positioned before S3. Unfortunately, Gardiner’s (1904) specimens were untraceable but his records are within the reported distribution proposed herein. The new material presented here increases the known depth range of this species by 75 m.

Figure 3. 

A, B Caryophyllia (Caryophyllia) stellula (SAM_H1485, off Agulhas, 200 m) A calicular view B lateral view C, D Caryophyllia (Caryophyllia) valdiviae (SAM_H3108, off Durban, depth unknown) C calicular view D lateral view E, F Crispatotrochus cornu (UCT_NAD 17 F, off Isipingo, 49 m) E calicular view F lateral view G, J Desmophyllum dianthus G, H (SAMC_A077974, off Paternoster, 440 m) G calicular view H lateral view I, J (BMNH.1939.7.20.218, locality data unknown) I calicular view J lateral view K, L Desmophyllum pertusum (SAM_H1605, off Melkbosstrand, depth unknown) K calicular view L lateral view M, N Goniocorella dumosa (SAM_H3190, off Kidds Beach, 760 m) M calicular view N lateral view O, P Heterocyathus aequicostatus (SAMC_A073186, off Durban, 150 m) O calicular view P lateral view Scale bars: 10 mm (A–I, K–P);100 mm (J).

Caryophyllia (Caryophyllia) valdiviae Zibrowuis & Gili, 1990

Fig. 3C, D

Caryophyllia clavus. –von Marenzeller 1904a: 281 (in part ‘Valdivia’–291–292), pl. 16, figs 9K–M.

Caryophyllia epithecata. –Boshoff 1981: 35.

Caryophyllia valdiviae Zibrowius & Gili, 1990: 28, 30, 32, pl. 2, figs P–R, pl. 3, figs A–T.

Type locality

Walvis Ridge, west of Namibia (‘Benguela VI Expedition’ stn. BB12: 25°34'00.0"S, 6°07'00.0"E); 886 m (Zibrowius and Gill 1990).

Type material

The holotype is deposited at the NHMUK (Zibrowius and Gill 1990).

Material examined

SAM_H3108 (2 specimens): Eastern margin, 35 km from Durban/33 km off Mbokodweni Estuary, 30°03'49.62"S, 31°15'30.89"E; depth unknown. ORI_DIIIa1 (7 specimens): Eastern margin, no other locality data.

Description

Corallum ceratoid, unattached, curved, and with a small pedicel (PD:GCD = 0.1). Calice subcylindrical, slightly compressed (GCD:LCD = 0.9–1.2), and with a jagged calicular margin. Largest specimen examined (SAM_H3108) 26.8 × 23.0 mm in CD, 2.8 mm in PD, and 55.3 mm in H. Costae poorly developed and flat, except for C1–2, which are prominent and high. C1–2 slightly wider than C3–4. All costae covered with fine granules, extending towards base, and separated by narrow intercostal striae. Theca thick. Corallum white, with beige theca.

Septa octamerally arranged in four cycles according to the formula: S1 > S2 > S3–4 (64 septa). S1 most wide, with straight axial margins that meet columella deep in fossa. S2 slightly less wide than S1, with straight to slightly sinuous axial margins. S1–2 most exsert. S3–4 1/3 less wide than S2, and bear sinuous axial margins. S3 the most sinuous and each bearing a tall and thin pali (16 P3). All septal faces covered with sparsely arranged granules. Fossa relatively deep, with a fascicular columella composed of 4–12 ribbon-like elements.

Distribution

Regional: Western (Zibrowius and Gili 1990) and eastern margins of South Africa, off Alexander Bay (Zibrowius and Gili 1990) and southeast of Durban; 442–882 m. Elsewhere: Walvis Ridge (Zibrowius and Gili 1990); 882–2670 m.

Remarks

The examined specimens match the characteristics described by Zibrowius and Gili’s (1990) Caryophyllia (C.) valdiviae specimens in having: (i) octamerally arranged septa, (ii) four cycles, and (iii) 16 P3. Caryophyllia valdiviae is similar to C. cornulum Cairns & Zibrowius, 1997 in adult specimens having 48–72 septa and bearing four to five septa size classes (Kitahara et al. 2010 – key). However, C. valdiviae can be distinguished by having septa highly exsert (1.8–4.2 mm) as compared with 0.7–1.1 mm in C. cornulum, S1–2 being equally exsert, and also in septal formula: S1 > S2 > S3–4 in C. valdiviae and S1 > S3 ≥ S2 in C. cornulum. Among the other three unattached Caryophyllia recorded in South Africa (C. grandis, C. scobinosa, and C. stellula), C. valdiviae overlaps in distribution and superficially resembles C. grandis, but may be distinguished by its adult corallum having septa octamerally arranged in four cycles (64 septa) as compared with septa hexamerally arranged in five cycles (96 septa). The number of resultant pali also differs between these two species, with C. valdiviae having 16 P3 and C. grandis 24 P3. Furthermore, C. valdiviae bears a robust and trochoid corallum, while C. grandis have a less robust and ceratoid corallum. Caryophyllia valdiviae is also one of the species reported by von Marenzeller (1904a) as C. clavus (Valdivia Expedition stn. 83) (Zibrowius and Gill 1990). Apart from von Marenzeller’s (1904a) misidentification of the Atlantic (Walvis Ridge) record, Boshoff (1980) reported an Indian ocean record of C. valdiviae as C. epithecata, a sample collected through UCTES and a sub-sample of SAM_H3108. Nonetheless, this KwaZulu-Natal record represents a disjunction in the previously known Atlantic distribution of this species, indicating that this species might occur all around the South African continental slope.

Crispatotrochus Tenison-Woods, 1878

Diagnosis

Corallum solitary, ceratoid to turbinate, and usually attached. Septotheca costate or covered with transverse ridges. Pali absent; columella fascicular composed of discrete, twisted elements.

Type species

Crispatotrochus inortatus Tenison-Woods, 1878, by monotypy.

Crispatotrochus cornu (Moseley, 1881)

Fig. 3E, F

Cyathoceras cornu Moseley, 1881: 156–157, pl. 4, fig. 7 (in part). –Cairns 1979: 67, pl. 12, figs 1, 3.

Crispatotrochus cornu. –Cairns 1991: 15. –Kitahara and Cairns 2008: 63. –Cairns and Polonio 2013: 70–71, figs 2L–M, 3A–B, 10.

Type locality

Off Rio de la Plata, Uruguay (HMS ‘Challenger’ stn. 320: 37°17'00"S, 53°52'00"W); 1097 m (Moseley 1881).

Type material

Lectotype and paralectotype are deposited at the NMNH (Cairns 1979).

Material examined

None.

Imagery data

UCTES_NAD 17 F (1 specimen): Eastern margin, off Isipingo; 49 m.

Description

Corallum ceratoid, attached through a robust pedicel (PD:GCD = 0.4) that expands into an encrusting base. Calice elliptical (GCD:LCD = 1.2) with a serrated calicular margin. Only imaged specimen examined (NAD 17 1F) 5.9 × 5.0 mm in CD, 2.2 mm in PD, and 7.0 mm in H. Theca granulated. Costae prominent at calicular margin, with C1––2 more prominent and distinctive. No intercostal striae. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1–2 > S3 > S4 (48 septa). S1–2 equal in size and exsertness, with slightly sinuous axial margins, and almost meet columella. Remaining septa (S3–4) becoming progressively less exsert. S3 ~ ¾ the width of S1–2, and have the most sinuous axial margins. S4 rudimentary, also with straight to slightly sinuous axial margins. Septal faces bear small and blunt granules. Fossa moderately deep, with a rudimentary fascicular columella composed of two twisted elements.

Distribution

Regional: Eastern margin of South Africa, off Isipingo; 49 m. Elsewhere: Seamount or ridge on Heezen fracture zone, South Pacific; 549 m (Cairns 1982).

Remarks

The imaged specimen, that forms the basis of this species report, is consistent with the known description of Crispatotrochus cornu in having a robust pedicel and septa hexamerally arranged in four complete cycles (S1–2 > S3 > S4), but differs in having an exceptionally sinuous as opposed to a straight S3 as previously reported (Cairns 1979, Kitahara and Cairns 2008). This species groups with the other Crispatotrochus that have septa hexamerally arranged in four cycles (C. inornatus, C. galapagensis Cairns, 1991, C. rugosus Cairns, 1995, and C. irregularis Cairns, 1982), all previously reported from the Pacific Ocean. Crispatotrochus cornu appears to be a juvenile form and is similar to the specimens collected by RV Gerda and RV Pillsbury stations which display a pedicel measuring ½ of the calicular diameter (Cairns 1979). However, Cairns (1979) highlights that these specimens were omitted from the description/account on the bases of immaturity and their small size. Species may have septa arranged in a decameral or hexameral fashion (Cairns 1979, 1991), the latter being the pattern observed in the imaged specimen reported herein (UCTES_NAD17F). Nonetheless, the South African record (UCTES_NAD17F) was collected at a much shallower depth than the previously known. Despite this, specimen is added to the account and represents a new record for the Crispatotrochus in the southwest Indian Ocean.

Desmophyllum Ehrenberg, 1834

Diagnosis

Solitary, trochoid, fixed. Pali absent. Columella absent or rudimentary. Sparse endothecal dissepiments.

Type species

Madrepora dianthus Esper, 1794, by subsequent designation (Cairns 1994).

Desmophyllum dianthus (Esper, 1794)

Fig. 3G–J

Madrepora dianthus Esper, 1794: pl. 69, figs 1–3. –Esper 1795: 85–86. –Scheer 1990: 406.

Desmophyllum cristagalli Milne-Edwards & Haime, 1848a: 253, pl. 7, figs 10, 10a. –Milne-Edwards and Haime 1857: 76. –Duncan 1873: 321. –Pourtalès 1878: 203 (in part: Blake Stn. 2). –Pourtalès 1880: 96, 106 (in part: BL–288). –Verrill 1885: 150. –Agassiz 1888: 151. –Alcock 1902c: 28. –von Marenzeller 1904a: 267–268, pl. 15, fig. 2A–B. –Vaughan 1907: 67, pl. 7, figs 3, 3A–B. –Verrill 1908: 494. –Stephens 1909: 25. –Döderlein 1913: 126, pl. 8, figs 45, 45A. –Gravier 1920: 72–76 (in part), pl. 8, figs 130–135. –Gardiner 1929: 125–126. –Zibrowius 1974a: 758–761, pl. 3, figs 1–10. –Zibrowius et al. 1975: 98, pl. 4, fig. A–B. –Zibrowius and Grieshaber 1977: 379. – Zibrowius 1978: 535. –Zibrowius 1979: 19, pl. 1, figs 5–6. –Cairns 1979: 117–119, pl. 21, figs 7–8, pl. 22, fig. 8. –Zibrowius 1980: 117–121, pl. 61, figs A–O, pl. 62, figs A–M. –Cairns 1981: 10. –Cairns 1982: 29, pl. 8, figs 9–12, pl. 9, figs 1–3. –Zibrowius 1988: 136. –Zibrowius and Gili 1990: 35–36. –Cairns 1991: 17, pl. 6, figs G–I. –Cairns and Parker 1992: 28–29, fig. 8B–C. –Tyler and Zibrowius 1992: 227. –Cairns and Keller 1993: 246.

Desmophyllum capense Gardiner, 1904: 96–97. –Gardiner 1939: 329–330. –Wells 1958: 262. –Cairns 1979: 206.

Desmophyllum capensis. –Squires 1961: 23, fig. 5.

Desmophyllum dianthus . –Ehrenberg 1834: 299–300. –Milne-Edwards and Haime 1848a: 254–255. –Milne-Edwards and Haime 1857: 77–78. –Yabe and Eguchi 1942b: 113–114, pl. 9, figs 1–3. – Eguchi 1965a: 290. –Cairns 1994a: 26–27, pl. 9, figs 9A–D. –Cairns 1995: 77, pl. 21, figs D–F. –Cairns and Zibrowius 1997: 131, fig. 17G–H. –Cairns 1998: 385–386. –Cairns 1999a: 104–105. –Cairns et al. 1999: 22. –Stolarski 2003: 508, fig. 7A–G. –Cairns 2004a: 281. –Cairns 2006: 47. –Kitahara 2007: 502, 503, fig. 3K–L. –Pires 2007: 269. –Cairns 2009: 13. –Kitahara and Cairns 2021: 520–523, figs 284D–E, 289.

Type Locality

Sagami Bay, Japan, depth unknown (Cairns 2004a).

Type material

The neotype is deposited at the NMNH (Cairns 1994a).

Material examined

SAMC_A072968 (2 specimens): no locality data. SAMC_A072974 (2 specimens): Western margin, 168 km off Paternoster/173 km off Brak Estuary, 32°05'41.99"S, 16°19'47.99"E; 440 m. SAMC_A073013 (2 specimens): Eastern margin, 34 km from Coffee Bay/18 km off Ntlonyane Estuary, 32°17'23.99"S, 29°05'35.87"E; 340–450 m. SAMC_A073015 (2 specimens): Southern margin, 32 km from Mazeppa Bay/19 km off Mendu Estuary, 32°25'00.11"S, 28°58'18.11"E; 330–340 m. SAMC_A073263 (1 specimen): Eastern margin, Wright Canyon; 171 m. SAMC_A088918 (3 specimens): Southern margin, 23 km from Plettenberg Bay/25 km off Piesang Estuary, 34°16'15.60"S, 23°24'50.40"E; 95 m. SAMC_A088919 (2 specimens): Southern margin, 172 km from Agulhas/182 km off De Mond-Heuningnes Estuary, 36°20'22.20"S, 20°24'06.11"E; 166 m. SAMC_A088929 (1 specimen): Southern margin, 140 km off Agulhas/144 km Ratels off Bulura Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A090122 (1 specimen): Eastern margin, 15 km south of Ponta Do Ouro/17 km off Kosi-Kumpungwini (Sifungwe) Estuary, 26°55'30.00"S, 33°01'05.88"E; 370 m. SAM_A090128 (1 specimen): Western margin, 46 km from Paternoster/71 km off Berg River V Estuary Estuary, 32°56'32.93"S, 17°25'14.16"E; 325 m. SAM_A090130 (1 specimen): Western margin, 147 km from Groen River/144 km off Brak Estuary, 31°40'45.59"S, 16°23'07.80"E; 360 m. SAM_A090131 (1 specimen): Southern margin, 74 km from Agulhas/79 km off Ratels Estuary, 35°27'43.79"S, 19°51'20.99"E; 154 m. SAM_A090132 (1 specimen): Western margin, 86 km from Hondeklipbaai/84 km off Spoeg Estuary, 30°47'55.25"S, 16°34'37.98"E; 235 m. SAM_A090133 (1 specimen): Western margin, 113 km from Groen River/114 km off Groen Estuary, 31°13'55.85"S, 16°34'37.98"E; 310 m. SAM_A090134 (1 specimen): Western margin, 195 km from Port Nolloth/194 km off Buffels Estuary, 30°04'07.79"S, 15°05'26.69"E; 393 m. SAM_A090135 (1 specimen): Western margin, 165 km from Port Nolloth/165 km off Holgat Estuary, 29°45'08.82"S, 15°16'20.22"E; 183 m. SAMC_A090151 (1 specimen): Southern margin, 116 km from Knysna/ off Goukamma Estuary, 35°07'11.27"S, 23°02'41.91"E; 333 m. SAM_H1475 (1 specimen): Western margin, 3 km from Pringle Bay/6 km off Buffels Oos Estuary, 34°23'11.29"S, 18°49'49.39"E; 80 m. SAM_H3049 (2 specimens): Eastern margin, 20 km off Cape Vidal/23 km off St Lucia Estuary, 27°59'30.00"S, 32°40'47.99"E; 550 m. SAM_H3050 (1 specimen): Eastern margin, 16 km from Margate/off Boboyi Estuary, 30°52'59.99"S, 30°31'00.00"E; 850 m.

Imagery data

BMNH 1939.7.20.218 (2 specimens), MCZ (1 specimen): locality data unknown. SS ‘Pickle’ stn. 1480 (1 specimen): Western margin, off Hout Bay; 131 m.

Description

Corallum variable, ranging from serpentine to ceratoid, attached to substrate by a pedicel (PD:GCD = 0.3–0.8) that expands into an encrusting base. Calice elliptical (GCD:LCD = 1.1–2.5), calicular margin serrate. Largest specimen examined (SAMC_A088919) 57.4 × 22.8 mm in CD, 30.0 mm in PD, and 119.3 mm in H. Theca granular. Costae more prominent near calicular margin, disappearing towards base. Corallum white to beige.

Septa hexamerally arranged in five complete cycles according to formula: S1–2 > S3 > S4 > S5 (96 septa). S1–2 most exsert, extend furthest to fossa (sometimes almost meeting opposite septa), with vertical and straight axial margins. S3 half as exsert and ⅓ smaller than S1–2, also with vertical and straight axial margin. S4 least exsert septa and 4/5 the size of S3, with straight to slightly sinuous axial margin. S5 more exsert than S4 and fuses to adjacent S13 at calicular margin. S5 ~ 1/2 the width of S4, with straight upper axial margins that become sinuous deeper in fossa. Fossa deep, columellar absent.

Distribution

Regional: Western to eastern margin of South Africa, off Lambert’s Bay extending towards Kosi-Kumpungwini (Sifungwe) Estuary (15 km south of Ponta Do Ouro: Mozambique); 80–850 m. Elsewhere: Cosmopolitan except from continental Antarctica and Boreal Pacific (Cairns 1994a); 8–2460 m.

Remarks

Desmophyllum dianthus is the most well-studied azooxanthellate solitary coral. The species was first reported in South Africa by Gardiner (1904) sample SAM_H1475 off Cape Hangklip (False Bay area) at 80 m deep.

Desmophyllum pertusum (Linnaeus, 1758)

Fig. 3K, L

Madrepora pertusa Linnaeus, 1758: 797.

Madrepora prolifera Pallas, 1766: 307.

Lophelia prolifera. –Milne-Edwards and Haime 1850a: 81. –Cecchini 1917: 149. –Laborel 1970: 156. –Cairns 1979:125–127, pl. 24: figs 1–5. –Cairns 1982: 30–31, pl. 9: fig. 6. –Cairns 1991a:17–18, pl. 6: fig. J.

Lophohelia prolifera. –Milne-Edwards and Haime 1857: 117. –Pourtalès 1871: 24–25, pl. 1, figs 3–5. –Duncan 1873: 328–332, pl. 42, figs 7–8. –Moseley 1881: 178–179, pl. 8, figs 7–8 (not Challenger–109). –Verrill 1883: 63–64. –Agassiz 1888: 151, fig. 472. –Jordon 1895: 25. –von Marenzeller 1904a: 307, pl. 15, figs 3, 3A. –Gourret 1906: 121, pl. 11, fig. 10, pl. 12, fig. 10A. –Gravier 1920: 87–89 (in part: not pl. 10, fig. 157). –Nobre 1931: 67–68, pl. 19–20.

Lophelia affinis Pourtalès, 1868: 135.

Lophohelia tubulosa Studer, 1878: 631, pl. 1, fig. 8A–E

Bathelia candida . –Jourdan 1895: 27.

Lophelia californica Durham, 1947: 36, pl. 1: figs 13, 16; pl. 2: fig. 11. –Cairns 1991: 17.

Dendrosmilia nomlandi . –Durham and Barnard 1952: 85, pl. 10: fig. 47. –Cairns 1979: 126. –Bythell 1986:16, pl. 10: fig. F.

Desmophyllum cristagalli. –Squires 1959a: 18–22 (in part: figs 8–10).

Lophelia pertusa. –Zibrowuis 1974b: 761, pl. 2, figs 6–9. –Zibrowuis 1980: 126–130, pl. 66, figs A–L. –Zibrowius and Gill 1990: 36–38. –Cairns and Keller 1993: 218. –Cairns 1994: 27–28: pl. 9. figs E–I. –Cairns 2000: 100–102.

Desmophyllum pertusum Addamo et al., 2016: 10–11, fig. 1A, B, D, E, fig. 3F–I.

Type locality

Southern California, depth unknown (Cairns 1994a).

Type material

The type is presumed lost.

Material examined

SAMC_A072974 (2 fragments): Western margin, 168 km off Paternoster/173 km off Brak Estuary, 32°05'41.99"S, 16°19'47.99"E; 440 m. SAMC_A088910 (1 fragment): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAMC_A088911 (1 fragment): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAMC_A088912 (1 fragment): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAMC_A088914 (1 fragment): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAMC_A088915 (1 fragment): Western margin, 53 km from Saldanha/80 km off Berg River V Estuary, 33°06'35.77"S, 17°23'01.26"E; 375 m. SAM_A090136 (1 specimen): Western margin, 162 km from Hondeklipbaai/159 km off Spoeg Estuary, 31°10'25.80"S, 15°54'55.79"E; 434 m. SAM_H1605 (5 fragments): Western margin, 43 km off Melkbosstrand/7 km off Dwars (North) Estuary, 33°24'39.01"S, 18°10'11.80"E; depth unknown. SAM_H1608 (6 fragments): Western margin, 13 km from Cape Town/6 km off Diep Estuary, 33°52'59.66"S, 18°25'34.08"E; depth unknown. SAM_H3129 (8 fragments): Eastern margin, 30 km off Coffee Bay/20 km off Bulungulu Estuary, 32°15'00.00"S, 29°09'06.00"E; 500–520 m.

Description

Colony dendroid to bushy (branching pattern variable). Majority of budding intratentacular. Slender terminal branches bearing sympodially arranged corallites. Calicular size variable, reaching ≤ 15.6 × 9.0 mm in CD. Calice circular to elliptical (GCD:LCD = 1.0–1.3), with slightly serrated calicular margin. Peritheca finely granular, resulting in a smooth texture. Costae short and ridged, generally corresponding to primary septa. Corallum white.

Septa not arranged in regular systems nor cycles. Seven to nine slightly exsert primary septa extend deep into fossa. Secondary septa slightly less exsert and less wide than primaries, sometimes also extending deep into fossa. Tertiaries smaller than secondary septa, being rudimentary deep in fossa. Tertiaries sometimes missing in some systems. All septa with vertical and straight axial margins. Fossa deep and curved. Columella absent.

Distribution

Regional: Western to eastern margin of South Africa, extending from off Clanwilliam extending towards Coffee Bay; 350–520 m. Elsewhere: Cosmopolitan in temperate and tropical waters (Zibrowius and Gill 1990; Cairns 2000), being common in the Atlantic and rarely collected off the Indo-Pacific (Cairns 1999b); 60–2170 m.

Remarks

Recent molecular studies suggest that Lophelia is a synonym of Desmophyllum (Addamo et al. 2012, 2016). Such findings are demonstrated by the overwhelming genetic similarities between L. pertusa and D. dianthus, however, we recommend the sequencing of additional genes as a priority for future studies pertaining to this name change. Lophelia pertusa is one of the most well-known and studied azooxanthellate framework-building coral and was first reported in South African waters by Zibrowius and Gill (1990), who did not list locality data.

Goniocorella Yabe & Eguchi, 1932

Diagnosis

Colonial, extra-tentacular budding forming bushy colonies. Branch anastomosis common, the branches also united by slender, tubular coenosteal bridges. No pali nor columella. Tabular endothecal dissepiments common and widely spaced.

Type species

Pourtalosmilia dumosa Alcock, 1902c, by original designation.

Goniocorella dumosa (Alcock, 1902)

Fig. 3M, N

Pourtalosmilia dumosa Alcock, 1902c: 36–37, pl. 5, fig. 33.

Goniocorella dumosa. –Yabe and Eguchi 1932a: 389–390. –Eguchi 1965b: 291, 2 figs. –Cairns 1982: 31–34, pl. 9, figs 7–9, pl. 10, figs 1, 2. –Cairns and Keller 1993: 250. fig. 6E. –Cairns 1995: 80–81, pl. 22, figs E, F.

Type locality

Banda Sea, Indonesia (HMS ‘Siboga’ stns. 156 and 259: 0°29'02.00"S, 130°05'03.00"E and 5°29'02.00"S, 132°52'05.00"E, respectively); 469–487 m (Alcock 1902c).

Type material

The syntypes are deposited at the ZMA (Cairns 1994a).

Material examined

SAMC_A088913 (1 fragment): Southern margin, 110 km from Oubosstrand/off Tsitsikamma Estuary, 35°02'25.19"S, 23°59'33.60"E; 915 m. SAMC_A090137 (1 fragment): Southern margin, 203 km from Gouritsmond/214 km off Goukou Estuary, 36°08'21.59"S, 22°23'39.59"E; 997 m. SAMC_A090138 (1 fragment): Southern margin, 35 km from Knysna/33 km off Goukamma Estuary, 34°21'01.79"S, 22°51'01.19"E; 87 m. SAMC_A090139 (1 fragment): Southern margin, 25 km from Jeffreys Bay/20 km off Gamtoos Estuary, 34°04'46.20"S, 25°11'24.60"E; 69 m. SAM_H3185 (2 fragments): Eastern margin, 20 km from Cape Vidal/23 km off St Lucia Estuary, 27°59'30.00"S, 32°40'47.99"E; 550 m. SAM_H3186 (2 fragments): Southern margin, 31 km from Port Alfred/20 km off Kleinemond Estuary, 33°39'24.00"S, 27°11'42.00"E; 86 m. SAM_H3187 (1 fragment): Southern margin, 54 km from Port Edward/ km off Mdumbi Estuary, 33°00'00.00"S, 30°27'11.99"E; 900 m. SAM_H3188 (2 fragments): Eastern margin, 17 km from Margate/off Boboyi Estuary, 30°53'24.00"S, 30°31'41.99"E; 850 m. SAM_H3189 (16 fragments): Eastern margin, 36 km off Port Shepstone/49 km off Mtentu Estuary, 30°43'11.99"S, 30°48'47.99"E; 900 m. SAM_H3190 (14 fragment): Southern margin, 26 km from Kidds Beach/27 km off Ncera Estuary, 33°19'36.00"S, 27°52'23.99"E; 760 m.

Description

Corallum bushy. New branches formed from extra-tentacular budding at right angles from parent branch. Colonies reinforced by coenosteal bridges, which unite adjacent branches. Coenosteum with low and round granules. Branches cylindrical and straight, with circular to slightly elliptical (GCD:LCD = 1.0–1.4) corallites. Costae poorly developed, terminal corallites with slightly ridged C1–2.

Septa hexamerally arranged in three cycles according to the formula: S1 > S2 > S3. Upper region of all septa usually narrower than lower part. S1 slightly exsert, with straight and vertical axial margins. S2 1/3 the width of S1, also with straight and vertical margins. S3 rudimentary and bearing dentate axial margins. Fossa deep, usually filled with tabular endothecal dissepiments. Columella absent.

Distribution

Regional: Southern to eastern margin of South Africa, Port Alfred extending towards Cape Vidal; 86–997 m. Elsewhere: Indonesia (Cairns and Zibrowius 1997); Japan (Yabe and Eguchi 1932a; Cairns 1994a); New Zealand (Cairns 1995); Antarctic and Sub-Antarctic regions (Eguchi 1965b; Cairns 1982); 100–760 m.

Remarks

Goniocorella dumosa is known to contribute to the three-dimensional habitat structures in deep waters (Cairns 1982, 1995; Le Goff-Vitry et al. 2004). It is distinctive from the other framework-building caryophylliids (e.g., Solenosmilia variabilis and Lophelia pertusa) in asexually reproducing by extra-tentacular budding, having coenosteal bridges that reinforce the colony, and also by having prominent ridges on terminal corallites. The species was first reported from South Africa by Cairns and Keller (1993), a sub-sample from their South African record was also examined (SAM_H3190) in the present study.

Heterocyathus Milne-Edwards & Haime, 1848

Diagnosis

Corallum free and usually encapsulating a gastropod or scaphopod shell inhabited by a sipunculan worm. Costae at lateral theca distinct and either equal or unequal in thickness. At base costae transform into granulations. Lower part of corallum shows a relatively large worm opening (occasionally two) and several small pores.

Type species

Heterocyathus aequicostatus Milne-Edwards & Haime, 1848, by subsequent designation (Milne-Edwards & Haime, 1850b).

Heterocyathus aequicostatus Milne-Edwards & Haime, 1848

Fig. 3O, P

Heterocyathus aequicostatus Milne-Edwards & Haime, 1848a: 324, pl. 10, fig. 8. –Milne-Edwards and Haime 1857: 51. –Alcock 1893: 141. –Gardiner 1904: 105–112, 125 (in part), pl. 3, figs 1–11, 22–32, 39–43. –Gardiner 1905: 955. –Bourne 1905: 193–194, 213–226, pl. 3, pl. 4, figs 12–21. –Harrison and Poole 1909a: 898–899, pl. 85, fig. lA–F. –Harrison and Poole 1909b: 913. –Harrison 1911: 1026, pl. 58, fig. 12. –Faustino 1927: 83–87, pl. 8, figs 1–7. –Yabe and Eguchi 1932b: 443. –Gardiner and Waugh 1938: 186–187. –Umbgrove 1938: 265. –Eguchi 1941: 417. –Yabe and Eguchi 1941b: 213, fig. 6A, B. –Yabe and Eguchi 1941c: 270, figs 3–4. –Crossland 1952: 102–103. –Durham and Barnard 1952: 87–88, pl. 11, fig. 49A–D. –Scheer and Pillai 1983: 158, pl. 36, fig. 9. –Wells 1984: 310, fig. 4.1. –Zibrowius and Grygier 1985: 121. –Pillai 1986: 188. –Veron 1986: 558–559 (in part). –Hu 1988: 146, 147, pl. 3, figs 9, 12, 13, 16, 17. –Hoeksema and Best 1991: 226–230, figs 1–11. –Hodgson and Carpenter 1995: 243. –Cairns 1998: 382–384, fig. 3A, B. –Cairns et al. 1999: 22. –Veron 2000: 412–413, figs 1–4. –Stolarski et al. 2001: 324, figs 6A–D. –Cairns 2004a: 281. –Cairns 2009: 12. –Kitahara and Cairns 2021: 528–529, 531, figs 290, 291A–G.

Heterocyathus roussaeanus Milne-Edwards & Haime, 1848a: 324–325, pl. 10 fig. 9, 9A. –Vaughan and Wells 1943: pl. 41 figs 16, 16A.

Heterocyathus cochlea. –Gray 1849: 77, pl. 2, figs l, 2A. –Gray 1850: 410.

Psammoseris hemispherica. –Milne-Edwards and Haime 1851: 127. –Veron 1986: 610.

Stephanoseris rousseaui. –Milne-Edwards and Haime 1851: 127.

Stephanoseris lamellosa Verrill, 1865: 149.

Heterocyathus philippinensis Semper, 1872: 254 (in part: pl. 20 figs 13, 14). –Eguchi 1941a: 414.

Heterocyathus japonicus. –Yabe and Eguchi 1942b: 127–128, pl. 11 fig. 6 A, B.

Type locality

Unknown.

Type material

The type material was never traceable (Cairns 2004a).

Material examined

SAMC_A073100 (5 specimens): Eastern margin, 414 km south of Ponta Do Ouro/41 km off Groot Berg Estuary, 27°13'30.00"S, 32°49'30.00"E; 78 m. SAMC_A073106 (1 specimen): Eastern margin, 66 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°33'11.88"S, 32°43'00.12"E; 140 m. SAMC_A073186 (1 specimen): Eastern margin, 20 km from Durban/13 km off Mbokodweni Estuary, 30°01'05.87"S, 31°03'11.88"E; 150 m. USNM 90840 (1 specimen): 26 km from Port St. Johns/off Bulolo Estuary, 29°34'47.99"S, 31°41'59.99"E; 138 m.

Description

Corallum free, squat and always encapsulating a gastropod shell colonised by a sipunculid worm. Sipunculid efferent pore < 2 mm in diameter, located aborally. Base flat, but area with pore more prominent, giving a pear-shaped appearance. Calice slightly elliptical (GCD:LCD = 1.1). Largest specimen examined (SAMC_A073186) 8.2 × 7.8 mm in CD, and 4.6 mm in H. Costae equidistant, separated by narrow intercostal striae. Calicular margin lanceted. Corallum creamy.

Septa crowded and hexamerally arranged in four cycles according to the formula: S1 > S2 > S4 > S3 (48 septa). S1 most exsert, with rounded upper and oblique axial margins. S1 extend towards columella and bear a paliform lobe. S2 less exsert, ¾ the width of S1, and bearing multiple paliform lobes, which extend towards columella. S3 least exsert and smallest septa, but bear ≤ five paliform lobes. S4 almost as exsert as S2, bearing a paliform lobe (P4) that fuses to adjacent P3. All paliform lobes (P1–4) inclined towards columella. All septa have straight axial margins. Septal and palar faces granulated. Fossa shallow, containing a papillose columella composed of 10–25 cylindrical elements which are indistinguishable from pali.

Distribution

Regional: Eastern margin of South Africa, 19 km from Durban extending towards Groot Berg Estuary (414 km south of Ponta Do Ouro: Mozambique); 78–145 m. Elsewhere: Zanzibar (Gardiner and Waugh 1938); Australia (Cairns 1998); Japan (Verrill 1866); China; Thailand (Gray 1849); Philippines; and Indonesia (Hoeksema and Best 1991); New Caledonia (Kitahara and Cairns 2021); 0–268 m.

Remarks

Heterocyathus aequicostatus resembles H. alternatus in septal formula (S1 > S2 > S3 > S4), septa having straight axial margins, and in S3 bearing ≤ five pali, but can be distinguished in S1 bearing only one palus as compared with bearing two to three paliform lobes as in H. alternatus. Differences in costae among the two species may also be a distinguishing feature: H. aequicostatus has same sized costae as compared with those of H. alternatus sensu Hoeksema and Best 1991 that vary in size (with C1–2 sometimes being slightly narrower than other costae). Heterocyathus aequicostatus was first reported off South Africa by Gardiner (1904), who detailed the intraspecific variation by comparing the eastern margin to the western margin representatives. The examined eastern margin South African representatives add no new taxonomic knowledge to what is already known apart from extending the depth range reported by Hoeksema and Best (1991). Additionally, specimens examined include zooxanthellate representatives (> 40 m) (Hoeksema and Best 2015) and should therefore not be considered in biodiversity assessments focusing on azooxanthellate forms.

Figure 4. 

A, B Heterocyathus alternatus (ORI_DIIIe1_1, locality data unknown) A calicular view B lateral view C, D Heterocyathus monileseptatum sp. nov. (SAM_H1431, off Durban Harbour, 99 m) C calicular view D lateral view E, F Heterocyathus sulcatus (SAMC_A073123, off Shaka’s Rock, 100–105 m) E calicular view F lateral view G, H Labyrinthocyathus delicatus (SAM_H2836, off East London, 146–238 m) G calicular view H lateral view I, K Monohedotrochus capensis comb.nov. I, J (SAMC_A088924, off Kidds Beach, 247–147m) I calicular view J lateral view K (SAM_H3210, off Scottburgh, 690 m) calicular view L, M Polycyathus sp. (USNM 91677, off Port Dunford, 69 m) L calicular view M full view N, O Rhizosmilia robusta (USNM 91689, off Kosi Bay Estuary, 74 m) N calicular view O lateral view P Solenosmilia variabilis (SAM_H3158, off Cintsa, 630 m) P full view. Scale bars: 10 mm (A–G, I–O); 100 mm (H, P).

Heterocyathus alternatus Verrill, 1865

Fig. 4A, B

Heterocyathus alternata Verrill, 1865: 149

Heterocyathus alternatus . –Folkeson 1919: 10–11, pl. 1, figs 10–11. –Hoeksema and Best 1991: 230–231, figs 12–18. –Cairns 1998: 384, fig. 3D–E. –Cairns 1999a: 99–100, fig. 14E–F. –Cairns 2004a: 281.

Heterocyathus aequicostatus. –Boshoff 1981: 37 (in part).

Type locality

Gaspar Straits, between the Bangka and Belitung Islands, Sumatra, (Indonesia); depth unknown (Hoeksema and Best 1991).

Type material

The holotype is deposited at the YPM (Hoeksema and Best 1991).

Material examined

SAMC_A073105 (1 specimen): Eastern margin, 36 km from Cape Vidal/32 km off Mgobezeleni Estuary, 27°48'54.00"S, 32°38'24.00"E; 52 m. SAMC_A073126 (5 specimens): Eastern margin, 25 km from Cape Vidal/23 km off St Lucia Estuary, 27°54'42.11"S, 32°36'42.11"E; 42–50 m. SAMC_A073214 (1 specimen): Eastern margin, 37 km from Cape Vidal/32 km off Mgobezeleni Estuary, 27°48'47.88"S, 32°38'53.87"E; 50 m. SAM_H2808 (10 specimens): Eastern margin, 20 km from Cape Vidal/22 km off St Lucia Estuary, 27°59'04.99"S, 32°40'08.00"E; 550 m. SAM_H3118 (19 specimens): Eastern margin, 2 km from Durban/8 km off Umgeni Estuary, 29°52'00.00"S, 31°00'00.00"E; 99 m. SAM_H3119 (1 specimen): Southern margin, 3 km from Plettenberg Bay/2 km off Piesang Estuary, 34°03'06.80"S, 23°22'48.65"E; 30–35 m. ORI_DIIIe1_1 (1 specimen): Locality data unknown.

Description

Corallum squat, free, and with aboral or lateral efferent pores smaller than 1.5 mm in diameter. Base flat, but a prominent pore result in an asymmetric corallum. Calice circular to elliptical (GCD:LCD = 1.0–1.1). Largest specimen examined (SAMC_A073126) 9.5 × 8.6 mm in CD, and 6.4 mm in H. Costae well-defined, finely granulated, and usually unequal in width. C1–2 sometimes slightly narrower than other costae. Intercostal striae relatively deep but disappear towards base. Base finely granulated. Central part of corallum darker than costae and associated septa.

Septa loosely packed and having a star-like appearance, hexamerally arranged in four complete cycles according to the formula: S1 > S2 > S4 > S3 (48 septa). S1 highly exsert, each bearing two or three paliform lobes. S2 slightly less exsert than S1. S3 smallest septa, not as exsert as S1–2, and bearing one or two paliform lobes. S4 wider than S3 and bears four or five paliform lobes, which are distinguishable from columellar elements. At calicular margin, S4 fuses to adjacent S1–2 forming prominent lancets. All septa appear to have straight axial margins, with granules arranged perpendicular to septal faces. All paliform lobes cylindrical, rising slightly above columellar elements, with traces of small and sparsely arranged granules. Fossa relatively deep, containing a papillose columella composed of 5–20 cylindrical elements.

Distribution

Regional: Southern to eastern margin of South Africa, from Plettenberg Bay extending towards Cape Vidal; 30–150 m. Elsewhere: Off Indonesia (Hoeksema and Best 1991; Cairns 2004a); Vanuatu; and Wallis and Futuna Islands (Cairns 1999a); 0–319 m.

Remarks

Examined specimens closely resemble specimens reported by Hoeksema and Best (1991) as Heterocyathus alternatus. However, some of the examined specimens (e.g., SAMC_A073126) have a darker centre and costal pigmentation, characters which Hoeksema and Best (1991) suggested to be representative of H. sulcatus (see key from Hoeksema and Best 1991 [page 222]). Cairns (1999) also noted specimens of Heterocyathus that are conspecific to H. sulcatus, but lacked pigmentation, and other specimens of Heterocyathus that display colouration but are distinctively different from H. sulcatus. Based on this, we suggest that colouration is of low taxonomic importance. Nonetheless, H. alternatus closely resembles H. aequicostatus among the South African congers. Apart from the differences in septa and costae profile highlighted in the H. aequicostatus account (see remarks section), the less crowded septa and star-like appearance (a result of S4 inclination towards S3) of H. alternatus distinguishes it from H. aequicostatus. Furthermore, the pali of H. alternatus are distinguishable from columellar elements in comparison to that of H. aequicostatus, which are indistinguishable. Heterocyathus alternatus was first reported in South Africa by Boshoff (1991), who identified it as H. aequicostatus (ORI_DIIIe1). Additionally, specimens examined include zooxanthellate representatives (> 40 m) (Hoeksema and Best 2015) and should therefore not be considered in biodiversity assessments focusing on azooxanthellate forms.

Heterocyathus monileseptatum Filander & Kitahara, sp. nov.

Fig. 4C, D

Type locality

Off Durban Harbour, South Africa, 99 m.

Type material/

Holotype. SAM_H1431A: eastern margin, 5 km from Durban/7 km off Umgeni Estuary, (RV ‘Pieter Faure’: 29°52'00.00"S, 31°03'00.00"E); 99 m. Paratypes. SAM_H1431B (4 specimens): eastern margin, 5 km from Durban/7 km off Umgeni Estuary, (RV ‘Pieter Faure’: 29°52'00.00"S, 31°03'00.00"E); 99 m. SAM_H1246 (8 specimens): Locality data unknown.

Etymology

The species name monileseptatum (derived from Latin monile meaning “string of beads” and septum meaning “enclosure, wall, fence”) which alludes to the beaded septal margins.

Description

Corallum unattached and tall, with lateral efferent pores ranging in diameter from 1.00 ≤ 2.00 mm. Base flat, but a prominent basal pore results in an asymmetric corallum with an irregularly shaped base (BD = 1.1–1.6). Calice circular to elliptical (GCD:LCD = 1.0–1.1), with serrate calicular margin. Holotype (SAM_H1431) 11.0 × 9.7 mm in CD, 11.1 × 6.9 mm in BD, and 10.6 mm in H. Paratypes having one or two aboral pores randomly positioned. Costae equidistant and progressively decreasing in width towards base. All costae finely granulated. C4 bears distinctive low spine-like granules. At base each costae become a row of granules. Intercostal striae equal in width and depth. Corallum white, with theca and columella with blackish pigment.

Septa thin, spaced out, delicate, and hexamerally arranged in four complete cycles (which follow a Pourtalès plan) according to the formula: S1 ≥ S2 > S4 > S3 (48 septa). S1 most exsert and extend to columella with straight and smooth axial margins. S2 only slight less exsert and ca. as wide as S1. S2 axial margins slightly sinuous. S3 least exsert septa and 2/3 the size of S2. In each half-system, a pair of S4 joins common S3 deep in fossa, and extends towards columella as one septum. S3–4 junctions beaded. S4 more exsert than S3, and also with sinuous axial margin. S4 dimorphic in size: those adjacent to S1 being wider and more exsert than those adjacent to S2. At calicular margin, S4 fuses to adjacent S1–2 forming high rectangular lancets. Septal faces granular. No pali. Fossa deep, containing a poorly developed papillose columella composed of ≤ seven sparsely arranged rods.

Distribution

Regional: Eastern margin of South Africa, off Durban; 99 m.

Remarks

Amongst the six extant Heterocyathus species (H. aequicostatus, H. alternatus, H. antoniae Reyes, Santodomingo & Cairns, 2009, H. hemisphaericus Gray 1849, H. japanicus (Verrill, 1866), and H. sulcatus) (Hoeksema and Cairns 2021), the specimens reported herein are distinctive in the lack of pali, height of corallum, and by having a beaded axial margin at the S3–4 junctions. However, there are intermediate similarities with each of these species, for example Heterocyathus monileseptatum sp. nov. resembles H. aequicostatus, H. alternatus, and H. hemisphaericus in having septa arranged in four cycles (S1 > S2 > S4 > S3 = 48 septa), but differs from the first two species in having no pali rather than P1–4 (in case of H. aequicostatus) or P1, 3, 4 (in case of H. alternatus). On the contrary, the lack of pali in Heterocyathus monileseptatum sp. nov. is a similarity shared with H. hemisphaericus but this species can be distinguished by their septa profile: all septa are thin (max 0.6 mm) and solid in Heterocyathus monileseptatum sp. nov. compared with being thick (max 0.9 mm) and porous in H. hemisphaericus (Cairns 1998). The maximum height observed in Heterocyathus monileseptatum sp. nov. (10.9 mm) is higher than that reported in the Atlantic H. antoniae (9.0 mm) and both species also have septa arranged in a Pourtalès plan fashion. Nonetheless, differences between Heterocyathus sp. nov. and H. antoniae include (i) septa being arranged according to S1 > S2 > S4 > S3 and S1 = S2 = S4 >> S3, respectively; (ii) lack of pali in Heterocyathus monileseptatum sp. nov.; (iii) columella of Heterocyathus monileseptatum sp. nov. being papillose and composed of sparsely arranged rods versus a spongy columella composed of crispate elements in H. antoniae. Heterocyathus japanicus is also reported to have septa arranged in a Pourtalès plan (Zibrowuis 1997). However, Heterocyathus monileseptatum sp. nov. can be differentiated by beaded S3–4 axial margins, a feature that is unique in relation to all Heterocyathus.

Heterocyathus sulcatus (Verrill, 1866)

Fig. 4E, F

Stephanoseris sulcata Verrill, 1866: 48. –Vaughan 1905: 416.

Psammoseris cyclicioides Tenison-Woods, 1879 (in part): 10–11, pl. 1, figs 1–5. –Tenison-Woods 1880: 299–300.

Heterocyathus pulchellus Rehberg, 1892: 8–9, pl. 1, fig. 7A–B.

Homophyllia incrustans Dennant, 1906: 161, pl. 6, fig. 3A–B.

Heterocyathus aequicostatus . –Folkeson 1919: 8–10 (in part), pl. 1, figs 4–7. –Boshoff 1981: 37 (in part).

Heterocyathus cyclicioides. –Wells 1964: 109.

Heterocyathus sulcatus . –Hoeksema and Best 1991: 231–233, figs 19–23. –Cairns 1998: 384. –Cairns 1999a: 98–99, figs A–D. –Cairns et al. 1999: 22. –Stolarski et al. 2001: 320. –Randall 2003: 135. –Cairns 2004a: 281–282, fig. 3K. –Cairns 2009: 13. –Kitahara and Cairns 2021: 531–533, figs 291H–J, 292.

Type locality

Off Ceylon, Sri Lanka, depth unknown (Verrill 1866).

Type material

The holotype is deposited at the YPM (Verrill 1866).

Material examined

SAMC_A073054 (1 specimen): Eastern margin, 33 km from Richards Bay/39 km off Mlalazi Estuary, 29°04'00.00"S, 32°10'00.00"E; 50 m. SAMC_A073071 (1 specimen): Eastern margin, 9 km from Shaka’s Rock/13 km off Tongati Estuary, 29°34'23.87"S, 31°17'53.88"E; 60 m. SAMC_A073089 (1 specimen): Eastern margin, 67 km south of Ponta Do Ouro/14 km off Mgobezeleni Estuary, 27°26'12.11"S, 32°44'12.11"E; 55–60 m. SAMC_A073105 (1 specimen): Eastern margin, 36 km from Cape Vidal/32 km off Mgobezeleni Estuary, 27°48'54.00"S, 32°38'24.00"E; 52 m. SAMC_A073108 (1 specimen): Eastern margin, 42 km south of Ponta Do Ouro/27 km off Kosi Bay Estuary, 27°13'36.12"S, 32°49'18.11"E; 75 m. SAMC_A073123 (24 specimens): Eastern margin, 51 km from Shaka’s Rock/41 km off Zinkwasi Estuary, 29°30'17.99"S, 31°45'44.99"E; 100–105 m. SAMC_A073144 (1 specimen): Eastern margin, 35 km off Cape Vidal/32 km off St Lucia Estuary, 27°49'41.87"S, 32°38'12.11"E; 54 m. SAMC_A073156 (1 specimen): Eastern margin, 35 km from Cape Vidal/32 km off St Lucia Estuary, 27°49'41.87"S, 32°38'12.11"E; 54 m. SAMC_A073161 (1 specimen): Eastern margin, 26 km from Port St. Johns/off Bulolo Estuary, 31°49'59.99"S, 29°39'59.99"E; 140–145 m. SAM_H1245 (15 specimens), SAM_H1430 (1 specimen): Locality data unknown. SAM_H1472 (2 specimens): Eastern margin, 2 km from Durban/8 km off Umgeni Estuary, 29°52'00.00"S, 31°00'00.00"E; 99 m. SAM_H1512 (2 specimens): Eastern margin, locality data unknown; 55–165 m. SAM_H3112 (7 specimens): Eastern margin, 9 km off Shaka’s Rock/2 km off Tongati Estuary, 29°34'18.96"S, 31°11'05.25"E; 66 m. ORI_DIIIe1_2 (1 specimen): Locality data unknown.

Description

Corallum unattached and variable in shape. All specimens examined encapsulate a gastropod shell. Shape of corallum correlates with size and shape of gastropod shell. Aboral efferent pore not exceeding 2.0 mm in diameter. Calice circular to elliptical (GCD:LCD = 1.0–1.1). Largest specimen examined (SAMC_A073161) 9.8 × 9.0 mm in CD, and 6.4 mm in H. Costae granulated, unequal in size with C3–4 wider than C1–2, and progressively diminishing in size towards base. Base smooth. Upper parts of corallum, columella, and S1–2 darker than other corallum elements.

Septa hexamerally arranged in four complete cycles according to the formula: S1 ≥ S2 > S3 > S4 (48 septa). S1 most exsert, extend to columella with sinuous axial margins and bear a well-developed pali (P1). S2 slightly less exsert and may be equal or less wide than S1. S2 also have sinuous axial margins bordered by a smaller pali. S3 least exsert, also with sinuous axial margins bordered by variable sized pali. S4 dimorphic in development: those adjacent to S1 being wider and more exsert than those adjacent to S2. Approximately 1/2 distance to columella each S4 fuses to adjacent S1–2 forming a V-shaped pattern. Pali cylindrical and bear meniane-like ridges. Fossa shallow, containing a papillose columella.

Distribution

Regional: Eastern margin of South Africa, off Port St. Johns extending towards Kosi Bay Estuary (42 km south of Ponta Do Ouro: Mozambique); 50–164 m. Elsewhere: Indonesia (Hoeksema and Best 1991; Cairns 2004a); Australia (Cairns 1998); Vanuatu; Wallis and Futuna Islands (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); 11–351 m.

Remarks

Heterocyathus sulcatus differs from the other three South African congeners in having S3 > S4 as compared with S4 > S3 as in H. aequicostatus, H. alternatus, and Heterocyathus monileseptatum sp. nov. The presence of meniane-like structures on the palar faces of H. sulcatus further differentiates it from the other South African representatives. Part of the specimens reported herein were identified by Boshoff (1981) as H. aequicostatus, thus this account serves as a first record for the species in South African territory.

Labyrinthocyathus Cairns, 1979

Diagnosis

Corallum solitary, ceratoid to subcylindrical, and firmly attached. Costae poorly defined or composed of transverse epithecal ridges. Pali absent. Columella well developed and composed of an interconnected maze of lamellar plates.

Type species

Labyrinthocyathus langae Cairns, 1979, by original designation.

Labyrinthocyathus delicatus (von Marenzeller, 1904)

Fig. 4G, H

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

Cyathoceras cornu. –Gardiner 1904: 121–122.

Labyrinthocyathus sp. –Cairns 1979: 70, pl. 11, figs 10–11.

Paracyathus indicus. –Boshoff 1981: 38.

Labyrinthocyathus delicatus. –Zibrowuis and Gili 1990: 44. –Cairns and Keller 1993: 244.

Type locality

Off the Agulhas Bank, South Africa (SS ‘Valdivia’ stn. 104: 35°16'00"S, 22°26'00"E); 155 m (von Marenzeller 1904a).

Type material

Unknown.

Material examined

DEFF_AI2–INV 135 (2 specimens): Eastern margin, 37 km from Cintsa/21 km off Cwili Estuary, 32°49'59.99"S, 28°30'00.00"E; 228 m. SAMC_A073158 (2 specimens): Eastern margin, 10 km from Port Edward/24 km off Bilanhlolo Estuary, 31°05'48.00"S, 30°18'47.99"E; 140 m. SAMC_A073173 (1 specimen): Eastern margin, 24 km from Coffee Bay/17 km off Mdumbi Estuary, 31°58'00.00"S, 29°22'59.99"E; 200 m. SAMC_A073180 (13 specimens): Southern margin, 33 km from Mazeppa Bay/24 km off Cwili Estuary, 32°45'47.88"S, 28°36'24.12"E; 240–250 m. SAM_H1482 (4 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H2805 (1 specimen): Eastern margin, locality data unknown; 550 m. SAM_H2828 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H2832 (1 specimen): Southern margin, 3 km from Kei Mouth/off De Mond-Heuningnes Estuary, 32°42'31.81"S, 28°21'54.38"E; 159 m. SAM_H2834 (3 specimens): Southern margin, 25 km from Gonubie/24 km off Gqunube Estuary, 33°06'17.99"S, 28°10'59.99"E; 155 m. SAM_H2835 (20 specimens): Southern margin, 29 km from Cintsa/3 km off Morgan Estuary, 32°42'30.47"S, 28°22'07.88"E; 159 m. SAM_H2836 (7 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H2837 (2 specimens): Southern margin, 11 km from East London/3 km off Buffalo Estuary, 33°00'53.67"S, 27°55'50.67"E; 128 m. SAM_H2845 (2 specimens): Eastern margin, 17 km from Margate/off Boboyi Estuary, 30°53'24.00"S, 30°31'41.99"E; 850 m. SAM_H3131 (2 specimens): Southern margin, 40 km from Cintsa/29 km off Cwili Estuary, 32°55'00.00"S, 28°31'00.00"E; 630 m. SAM_H3132 (1 specimen): Eastern margin, 28 km from Coffee Bay/19 km off Bulungulu Estuary, 32°14'53.99"S, 29°10'23.99"E; 620–560 m. SAM_H3133 (7 specimens): Southern margin, 32 km off Mazeppa Bay/24 km off Kobole Estuary, 32°28'36.00"S, 28°58'48.00"E; 710–775 m. SAM_H3134 (1 specimen): Southern margin, 28 km from Mazeppa Bay/27 km off Kobole Estuary, 32°32'59.99"S, 28°55'00.00"E; 775–790 m. SAM_H4242 (1 specimen): Eastern margin, 11 km from Port St. Johns/15 km off Bulolo Estuary, 31°39'42.00"S, 29°38'59.99"E; 300–540 m. ORI_DIIIg1 (1 specimen): Eastern margin, other locality unknown.

Imagery data

AB_357 E (1 specimen): Eastern margin, 34 km from Port Dunford/38 km off Mlalazi Estuary, 29°10'00.00"S, 32°04'59.99"E; 168 m. BMNH 1973.2.20.26 (2 specimens): Eastern margin, 22 km from Port Shepstone/21 km off Damba Estuary, 30°46'59.99"S, 30°40'00.00"E; 457 m.

Description

Corallum solitary, ceratoid to trochoid, attached to substrate through a pedicel (PD:GCD = 0.3–0.5) that expands into an encrusting base. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), calicular margin thick and slightly serrated. Largest specimen examined (SAM_H2836) 11.6 × 11.4 mm in CD, 4.8 mm in PD, and 17.8 mm in H. Theca bears thin transverse ridges composed of fine granules. Costae poorly developed, granular, separated by faint and narrow intercostal striae, and more prominent near calicular margin. Corallum white.

Septa hexamerally arranged in four cycles according to the formula: S1–2 > S3 > S4 (48 septa). S1–2 largest septa and equal to only slightly more exsert than S2, almost reaching columella. S3 4/5 the size of S1–2. Higher cycle septa (S3–4) progressively less exsert. S4 ½ to ¾ the width of S3, with the least sinuous axial margin. All septal faces bear low rounded granules. Fossa moderately deep, containing a well-defined labyrinthiform columella composed of interconnected lamellar plates.

Distribution

Regional: Southern to eastern margin, off East London extending towards Port Dunford; 128–790 m. Elsewhere: off south-eastern Mozambique (Cairns and Keller 1993); 155–1000 m.

Remarks

Labyrinthocyathus delicatus was described by von Marenzeller (1904a) based on specimens sampled from off the Agulhas Bank, South Africa. Subsequently, Gardiner (1904) also reported on specimens collected off the Agulhas region but as Cyathoceras cornu. Zibrowius and Gili (1991) noted that Gardiner’s (1904) specimens of C. cornu were representatives of L. delicatus. As Gardiner’s (1904) specimens were not examined during the course of the present study, their identification remains tentative but in agreement with Zibrowius and Gili (1990) and Cairns and Keller (1993) as L. delicatus. Boshoff (1981) further reported the species in the region, but as Paracyathus indicus Duncan, 1889.

Monohedotrochus Kitahara & Cairns, 2005

Diagnosis

Corallum solitary, attached, straight, and elongate-conical to trochoid. Base monocyclic. Septotheca costate. Pedicel and base thick. Pali may be present, indistinguishable from columella. Columella papillose.

Type species

Monohedotrochus capitolii Kitahara & Cairns, 2005, by original designation.

Monohedotrochus capensis (Gardiner, 1904), comb. nov.

Fig. 4I–K

Caryophyllia capensis Gardiner, 1904: 113–114, pl. 1, fig. 4A–D. –Boshoff 1981: 36. –Zibrowius and Gill 1990: 44.

Desmophyllum cristagalli. –Boshoff 1981: 37.

Balanophyllia capensis. –Boshoff 1981: 40 (in part).

Paraconotrochus capensis . –Cairns and Parker 1992: 21.

Type locality

Off East London, South Africa (33°03'00"S, 27°57'00"E); 59 m (Gardiner 1904).

Type material

The holotype is lodged at the NMNH.

Material examined

SAMC_A072992 (1 specimen): Southern margin, other locality data unknown. SAMC_A073228 (1 specimen): Southern margin, False Bay; depth unknown. SAMC_A073233 (1 specimen): Southern margin, 14 km from Cape Point/10 km off Buffels Wes Estuary, 34°13'59.99"S, 18°30'00.00"E; 42 m. SAM_A073245 (2 specimens): Eastern margin, 34 km off Coffee Bay/7 km off Ntlonyane Estuary, 32°15'11.99"S, 28°57'42.00"E; 47 m. SAMC_A088924 (1 specimen): Southern margin, 23 km from Kidds Beach/24 km off Ncera Estuary, 33°18'01.37"S, 27°51'30.58"E; 247–147m. SAMC_A088927 (1 specimen): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAM_A090072 (1 specimen): Western margin, 13 km from Pringle Bay/off Buffels Oos Estuary, 34°18'36.00"S, 18°42'53.99"E; 71 m. SAMC_A090073 (1 specimen): Western margin, 23 km from Pringle Bay/20 km off Buffels Oos Estuary, 34°10'05.99"S, 18°47'03.00"E; 36 m. SAMC_A090074 (2 specimens): Eastern margin, 34 km from Coffee Bay/7 km off Ntlonyane Estuary, 32°15'11.99"S, 28°57'42.00"E; 47 m. SAMC_A090075 (1 specimen): Western margin, 49 km from Cape Point/51 km off Buffels Wes Estuary, 34°43'18.00"S, 18°12'29.99"E; 360–365 m. SAMC_A090076 (1 specimen): Southern margin, 16 km from Cape Point/off Buffels Wes Estuary, 34°15'00.00"S, 18°36'00.00"E; 51 m. SAMC_A090078 (1 specimen): Southern margin, 14 km from Cape Point/10 km off Buffels Wes Estuary, 34°13'59.99"S, 18°30'00.00"E; 42 m. SAMC_A090081 (1 specimen): Southern margin, 12 km from Pringle Bay/11 km off Buffels Oos Estuary, 34°16'59.99"S, 18°45'00.00"E; 58 m. SAMC_A090140 (1 specimen): Southern margin, 92 km from Oubosstrand/89 km off Tsitsikamma Estuary, 34°53'21.93"S, 24°06'56.47"E; 355 m. SAM_H1365 (2 specimens): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 40 m. SAM_H1366 (1 specimen): Southern margin, 5 km from East London/3 km off Blind Estuary, 33°00'13.79"S, 27°56'59.99"E; 59 m. SAM_H1374 (7 specimens): Southern margin, 241 km from Agulhas/247 km off De Mond-Heuningnes Estuary, 36°40'00.00"S, 21°25'59.99"E; 200 m. SAM_H1406 (2 specimens): Western margin, off Somerset West; depth unknown. SAM_H1439 (1 specimen): Southern margin, 25 km from Gonubie/24 km off Gqunube Estuary, 33°06'17.99"S, 28°10'59.99"E; 155 m. SAM_H1474 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3060 (1 specimen): Southern margin, 28 km from Gonubie/27 km off Buffalo Estuary, 33°09'29.99"S, 28°03'06.00"E; 86 m. SAM_H3061 (4 specimens): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 93 m. SAM_H3210 (4 specimens): Eastern margin, 30 km from Scottburgh/20 km off Fafa Estuary, 30°33'24.00"S, 30°48'35.99"E; 690 m. SAM_H3211 (1 specimen): Eastern margin, 17 km from Margate/off Boboyi Estuary, 30°53'24.00"S, 30°31'41.99"E; 850 m. SAM_H3398 (2 specimens): Southern margin, 11 km from Gansbaai/38 km off Bot River Lagoon, 34°39'27.93"S, 19°17'03.00"E; 36 m. ORI_DIIIa2 (2 specimens), ORI_DIIId1 (2 specimens), ORI_EIa4_2 (5 specimens): Eastern margin, other locality data unknown. USNM 100854 (1 specimen): Southern margin, 5 km from Port Elizabeth/3 km off Bakens River Estuary, 33°56'52.31"S, 25°37'20.70"E; depth unknown.

Imagery data

BMNH 1939.7.20.37–39 (1 specimen), BMNH 1950.1.11.63 (1 specimen): South Africa, other locality data unknown. SAM_H1377 (2 specimens): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 93 m.

Description

Corallum ceratoid to trochoid, firmly attached to substrate by a slender pedicel (PD:GCD = 0.3–0.7) that expands into a monocyclic encrusting base. Calice slightly elliptical (GCD:LCD = 1.1–1.3). Largest specimen examined (SAMC_A090140) 26.2 × 23.3 mm in CD, 9.6 mm in PD, and 48.1 mm in H. Costae flat to slightly convex, being prominent at calicular margin. Costae bear irregular and small granules. Intercostal striae thin and shallow. Both costae and intercostal striae become progressively faint towards base. Corallum white with theca being slightly light brown.

Septa hexamerally arranged in five cycles, sometimes the last cycle being incomplete, according to the formula: S1–3 > S3 > S4 > S5 (96 septa). S1–2 highly exsert and equal in width, extending to columella with straight axial margins. S3 less exsert, 1/3 less the width of S1–2, but in larger specimens meet columella deeper in fossa. However, in smaller specimens (< 13.0 mm in CD), S3 do not join columella and has a slightly dentate axial margin. S4 less exsert and ½ the width of S3, with dentate axial margin. S5 not exsert, rudimentary, with the most dentate axial margin. Fossa of moderate depth, containing a papillose columella composed of a group of 15–20 well-defined rods.

Distribution

Regional: Western to eastern margin of South Africa, off Gansbaai extending towards Scottburgh; 36–365 m. Elsewhere: No other distributional records are known.

Remarks

The species was first reported in South African boundaries by Gardiner (1904) as Caryophyllia capensis, and later tentatively placed in the genus Paraconotrochus (Cairns and Parker 1992). However, among the three genera (Caryophyllia, Paraconotrochus, and Monohedotrochus) only the latter lacks paliform lobes before S3. In addition, after examining Gardiner’s (1904: [fig. 4C]) illustrations, no pali were observed. Furthermore, the coralla of specimens examined display well-defined granular costae, whereas those of Paraconotrochus are generally smooth and costae absent or poorly defined. Based on these differences, we propose the new combination Monohedotrochus capensis (Gardiner, 1904). Within the examined material some are represented by small specimens (i.e., SAM_H3210 [Fig. 4K] and SAM_H3211), which resemble Gardineria Vaughan, 1907 in having transversely wrinkled epitheca and outer septal margins being separated from calicular margin by a notch. However, juvenile Monohedotrochus capensis and Gardineria species are difficult to distinguish and thus these two samples (SAM_H3210 and SAM_H3211) are added here with caution.

Polycyathus Duncan, 1876

Diagnosis

Corallum colonial. Corallites cylindrical to slightly conical bud from a common coenosteum or from stolons. Septotheca costate. Three to four cycles of septa. Pali present before all but last septal cycle. Columella papillose.

Type species

Polycyathus atlanticus Duncan, 1876, by monotypy.

Polycyathus sp.

Fig. 4L, M

Material examined

USNM 91677 (1 colony): Eastern margin, 33 km from Port Dunford/35 km off Mlalazi Estuary, 29°10'59.99"S, 32°01'59.99"E; 69 m.

Description

The colony consists of 12 ceratoid to cylindrical corallites that reach ≤ 4.9 mm in H, and bud from a common coenosteum. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), calicular margin thin and serrated. Theca glistening and covered by low-profile granules. Costae poorly developed. Corallum predominantly beige; but pali, columella, and base white.

Septa hexamerally arranged in four cycles, the last cycle being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 (≤ 34 septa). S1 most exsert, and as wide or only slightly wider than S2. Both S1–2 with sinuous axial margin, bearing pali which are as thick as septa. S3 not exsert, 3/4 the width of S1–2, and have vertical axial margin. S4 rudimentary. Septal faces granulated and slightly sinuous. Pali present and distinct in all but last septal cycle (14–16 pali). Half-systems with S4, P3 sometimes join neighbouring P2 resulting in a V-shaped appearance. All paliform lobes terminate at same level and form a crown encircling columella. Paliform lobes sometimes indistinguishable from columellar elements. Fossa of moderate depth containing a papillose columella composed of a group of fairly spaced rods.

Distribution

Regional: Eastern margin, Port Dunford; 69 m.

Remarks

Although corallites of the examined colony are mostly damaged, features that characterise the genus are still distinguishable. However, more specimens are required to enable a thorough comparison to the other Polycyathus representatives. This colony represents a new record of the genus in South Africa.

Rhizosmilia Cairns, 1978

Diagnosis

Colonies formed by extra-tentacular budding from a common basal coenosteum. Corallite base increase in diameter by adding exothecal dissepiments over raised costae producing concentric rings of partitioned chambers that resemble polycyclic development. Septotheca costate and granular. Septal axial margins smooth. Paliform lobes occur before penultimate septal cycle. Columella papillose or lamellar. Vesicular endotheca present.

Type species

Rhizosmilia gerdae Cairns, 1978, by original designation.

Rhizosmilia robusta Cairns in Cairns & Keller, 1993

Fig. 4N, O

Rhizosmilia robusta Cairns in Cairns & Keller, 1993: 250–253, pl. 6, figs F–I. –Cairns and Zibrowius 1997: 133–134. –Cairns 1999a: 107. –Cairns et al. 1999: 24. –Cairns 2009: 14. –Kitahara et al. 2010a 108, figs 88–89. –Kitahara et al. 2010b: 9. –Kitahara and Cairns 2021: 554–556, figs 305D–I, 306.

Type locality

Off Inhaca Island, Mozambique (RV ‘Anton Bruun’ stn. 373B: 26°00'00"S, 33°05'00"E); 135 m (Cairns and Keller 1993).

Type material

The holotype is deposited at the NMNH (Cairns and Keller 1993).

Material examined

USNM 91689 (3 specimens): Eastern margin, 42 km south of Ponta Do Ouro/27 km off Kosi Bay Estuary, 27°13'30.00"S, 32°49'30.00"E; 74 m.

Description

Adapted from Cairns and Keller 1993: Colony phaceloid. Corallites trochoid, firmly attached through a massive pedicle (PD:GCD = 0.03), that expands into a thin encrusting base. Lower pedicel reinforced by concentric rings of hollow chambers. Calice elliptical (GCD:LCD = 1.18), with a jagged calicular margin. Holotype (USNM 91681) 31 × 26.20 mm in CD, 16.80 mm in PD, and 25 mm in H. Costae equal and bearing low and rounded granules. Intercostal striae shallow. Corallum white.

Septa arranged in five cycles according to the formula: S1 > S2 > S3 > S4 > S5. S1 moderately exsert, and extend to columella with vertical to straight axial margins. Higher cycle septa (S2–5) progressively less exsert and less wide, except for those S5 adjacent to S1 which are more exsert than S4. Axial margins of S2 also straight, but those of S3–4 slightly sinuous. S5 rudimentary and irregular in profile (i.e., may be straight to slightly sinuous or sinuous to dentate). All septal faces smooth, bearing only sparsely spaced low-profile granules. Small pali present before penultimate cycle, usually indistinguishable from columellar elements. Fossa deep and bearing endothecal dissepiments. Columella trabecular, composed of an irregular group of intertwined elements.

Distribution

Regional: Eastern margin of South Africa, off Kosi Bay Estuary (41 km south of Ponta Do Ouro: Mozambique); 75 m. Elsewhere: Mozambique; Madagascar (Cairns and Keller 1993); Philippines (Cairns and Zibrowius 1997); New Caledonia (Kitahara and Cairns 2021); 66–996 m.

Remarks

As no new samples have been examined, the description above is adapted from Cairns and Keller (1993), who noted that although the holotype is a phaceloid colony with four corallites, paratypes (specimen examined and imaged herein) represent individual corallites that were broken apart from a larger colony or have not yet formed a colony. Variation in septal cycles correlates to corallum size: the fourth septal cycle is complete at a GCD between 8 and 9 mm, whereas the fifth is completed when the corallite is > 19–21 mm in GCD. The largest corallite (Philippines, see Cairns and Zibrowius 1997 [page 134]), measures 29.8 × 36.7 mm in CD, 18.1 mm in PD, 41.6 mm in H, have eight pairs of S6, and one pair of S7 (total of 116 septa). Notable differences between Rhizosmilia robusta from the Pacific and southwestern Indian Ocean are the position of paliform lobes, which are present before the penultimate cycle (P4, if S5 is present; and P3 if S5 is absent) in the latter and occur before S4 (irrespective of the presence of S6–7) in the former. This is the only representative of the genus in South African waters.

Solenosmilia Duncan, 1873

Diagnosis

Corallum firmly attached, colonial. Colony dendroid or sub-phaceloid formed by intratentacular budding. Stereome granular, costae sometimes correspond to first septal cycle. Tabular endothecal dissepiments. Columella small.

Type species

Solenosmilia variabilis Duncan, 1873, by monotypy.

Solenosmilia variabilis Duncan, 1873

Fig. 4P

Solenosmilia variabilis Duncan, 1873: 328, pl. 42, figs 11–18. –Pourtalès 1878: 206. pl. 1, figs 1–3. –Pourtalès 1880: 108. –Moseley 1881: 181, pl. 9, figs 1–5. –von Marenzeller 1904: 310–311, pl. 15, figs 4, 4A. –Zibrowius 1974a: 768–769. –Gravier 1920: 94–96, pl. 9, figs 153–156. –Hoffmeister 1933: 14, pl. 4, fig. 7. –Gardiner and Waugh 1939: 229–230. –Cairns 1979: 136–138, pl. 26, figs 2–4. –Zibrowius 1980: 143–145, pl. 75, figs A–N.– Scheer and Pallai 1983: 160. –Cairns and Parker 1992: 29–30, pl. 8, figs D, E.– Cairns and Keller 1993: 250, fig. 6D. –Cairns 2004a: 284.

Solenosmilia jeffreyi Alcock, 1898: 27–28, pl. 3, fig. 3, 3A, B.

Type locality

Off Spain (HMS ‘Porcupine’ stns. 17 and 32: 39°42'00"N, 9°43'00"W and 35°41'00"N, 7°08'00"W, respectively); 1190–2003 m (Duncan 1873).

Type material

The syntype is deposited at the NHMUK (Cairns 1979, 2004a).

Material examined

DSCS–INV 122 (1 fragment): Southern margin, 240 km from Agulhas/247 km off De Mond-Heuningnes Estuary, 36°45'34.13"S, 21°12'46.61"E; 513 m. SAMC_A088916 (1 fragment): Southern margin, 280 km from Cape St. Francis/287 km off Slang Estuary, 36°43'40.13"S, 25°08'53.47"E; 622 m. SAMC_A090142 (1 fragment): Southern margin, 240 km from Agulhas/247 km off De Mond-Heuningnes Estuary, 36°45'34.13"S, 21°12'46.61"E; 513 m. SAMC_A090143 (1 fragment): Southern margin, 287 km from Cape St. Francis/291 km off Slang Estuary, 36°47'35.77"S, 24°38'35.69"E; 520 m. SAM_H1397 (1 fragment): Southern margin, 3 km from East London/1 km off Buffalo Estuary, 33°01'29.99"S, 27°55'00.00"E; 566–928 m. SAM_H2807 (1fragment): Eastern margin, 20 km from Cape Vidal/22 km off St Lucia Estuary, 27°59'04.99"S, 32°40'08.00"E; 550 m. SAM_H2840 (14 fragments): Eastern margin, 19 km from Cape Vidal/22 km off St Lucia Estuary, 27°59'30.00"S, 32°40'00.00"E; 550 m. SAM_H3034 (13 fragments): Eastern margin, 36 km off Port Shepstone/49 km off Mtentu Estuary, 30°43'11.99"S, 30°48'47.99"E; 900 m. SAM_H3035 (10 fragments): Eastern margin, 16 km from Port Shepstone/off Boboyi Estuary, 30°49'05.99"S, 30°34'59.99"E; 930 m. SAM_H3036 (21 fragments): Eastern margin, 36 km from Port Shepstone/28 km off Mhlabatshane Estuary, 30°43'00.00"S, 30°48'47.99"E; 780 m. SAM_H3037 (1 fragment): Eastern margin, 17 km from Margate/off Boboyi Estuary, 30°53'24.00"S, 30°31'41.99"E; 850 m. SAM_H3140 (17 fragments): Southern margin, 26 km from Kidds Beach/27 km off Ncera Estuary, 33°19'36.00"S, 27°52'23.99"E; 760 m. SAM_H3141 (13 fragments): Southern margin, 32 km off Mazeppa Bay/24 km off Kobole Estuary, 32°28'36.00"S, 28°58'48.00"E; 710–775 m. SAM_H3142 (3 fragments): Southern margin, 22 km from Gonubie/21 km off Gqunube Estuary, 33°06'00.00"S, 28°08'17.99"E; 700–650 m. SAM_H3158 (38 fragments): Southern margin, 40 km from Cintsa/29 km off Cwili Estuary, 32°55'00.00"S, 28°31'00.00"E; 630 m. SAM_H3179 (1 fragment): Southern margin, 31 km from Port Alfred/20 km off Kleinemond (Oos) Estuary, 33°39'24.00"S, 27°11'42.00"E; 86 m.

Description

Corallum bushy achieved by intra-tentacular budding. Budding dichotomous. Calice rarely exceeds 6 mm in CD. Coenosteum completely smooth or granular. Costae ridged, with granules arranged in a longitudinal manner. Corallum white and sometimes light brown.

Septa hexamerally arranged in three or four cycles according to the formula: S1 > S2 > S3 > S4. S1 highly exsert and extend deep into fossa with straight axial margins. S2 less exsert and 1/3 the width of S1, but otherwise similar in profile. S3 not exsert and forms in upper fossa, appearing rudimentary. S4 usually absent, but when present has dentate or laciniate axial margin. Septal faces bear tall and slender granules. Tabular endothecal dissepiments common and widespread. Columella usually absent, but some corallites have a small spongy columella deep in fossa.

Distribution

Regional: Southern to eastern margin of South Africa, off Cape St. Francis extending towards Cape Vidal; 86–930 m. Elsewhere: Cosmopolitan, except from the continental Antarctica; 220–2165 m.

Remarks

Together with Madrepora oculata, Lophelia pertusa, and Goniocorella dumosa, Solenosmilia variabilis is one of the most studied framework-building coral species. The equal distomedial budding in S. variabilis makes it easily distinguishable from the other deep-water framework-building species. Solenosmilia variabilis was first reported from South Africa off the Agulhas region (von Marenzeller 1904a). Subsequently, Cairns and Keller (1993) documented its northward regional distribution to off Durban. Part of the examined material includes sub-samples (SAM_H3037, SAM_H3141, and SAM_H3158) of Cairns and Keller’s (1993) records, all of which are broken fragments. Therefore, details on the size of the base and end branches are not included in the description. Nevertheless, Cairns (1979) provides a more detailed description of these features.

Stephanocyathus Seguenza, 1864

Diagnosis

Solitary, patellate to bowl-shaped, and free. Costae usually well developed, some of which are sometimes highly spinose. Paliform lobes usually present on all septa. Columella trabecular, papillose, or a solid fusion of axial septal margins.

Type species

Stephanocyathus elegans Seguenza, 1864, by subsequent designation (Wells 1936).

Stephanocyathus Acinocyathus Wells, 1984

Diagnosis

Stephanocyathus with six elongate spines corresponding to C1.

Type species

Stephanotrochus spiniger von Marenzeller, 1888, by original designation.

Stephanocyathus (Acinocyathus) explanans (von Marenzeller, 1904)

Fig. 5A–F

Stephanotrochus explanans von Marenzeller, 1940a: 304–307, pl. 8, fig. 19A, B. –Gardiner and Waugh 1938: 192.

Stephanocyathus nobilis. –Boshoff 1981: 39.

Stephanocyathus (Acinocyathus) spiniger. –Cairns and Keller 1993: 243–244.

Stephanocyathus (Acinocyathus) explanans. –Cairns and Zibrowius 1997: 119, fig. 14E. –Cairns 1998: 38. –Cairns 2004a: 285.

Type locality

Off Sumatra, Zanzibar Island and Pemba (SS ‘Valdivia’ stns. 194, 243 and 245: 0°15'02"N, 98°08'08"E, 6°39'01"S, 39°30'08"E, and 05°27'09"S, 39°18'08"E, respectively); 245–614 m (von Marenzeller 1904a).

Type material

Ten syntypes are deposited at the ZMB (Cairns 2004a).

Material examined

SAMC_A090144 (2 specimens): Eastern margin, 8 km from Port St. Johns/11 km off Bulolo Estuary, 31°39'43.19"S, 29°36'38.16"E; 96–98 m. ORI_DIIIk2 (I specimen): Eastern margin, 33 km from Durban/31 km off Beachwood Mangroves, 29°55'00.00"S, 31°19'59.99"E; 442 m. USNM 62500 (4 specimens): Eastern margin, 11 km from Durban/4 km off Beachwood Mangroves, 29°46'29.16"S, 31°04'18.42"E; 183–220 m.

Imagery data

RV ‘Galathea’ stn. 202 (1 specimen): Eastern margin, Natal.

Description

Corallum bowl-shaped, free, with a slightly rounded base. BD smaller than CD; base has a basal attachment scar. All specimens bear short, straight, and slender C1 costal spines, and are usually longer (in relation to CD) in juveniles. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), with a serrated calicular margin. Largest specimen examined (USNM 62500) 30.4 × 27.6 mm in CD, 26.6 in BD, and 11.2 mm in H. Costae slightly convex and granular, extending from calicular margin and disappearing towards base. Costae absent at base epicentre, which is usually eroded. Corallum white.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 > S5 (≤ 72 septa). S1 highly exsert and extend to columella with straight axial margin. A deep notch separates P1 from S1. P1 fuse to columella. S2 as exsert and as large to only slightly smaller than S1. P2 also fuse to columella, but is slightly smaller than P1. S3 ~ 1 mm less exsert and ¾ the width of S1–2. P3 broad and separated from S3 by a narrow notch. S4 flanked by S5 as wide and exsert as S3, but those unflanked S4 are the least exsert septa, and only ½ the width of S3. S4 become rudimentary deeper in fossa. P4 fuse to P3. S5 closely resembling unflanked S4. All septal margins straight. Paliform lobes also straight, and appear to be arranged in a single palar crown. Fossa relatively shallow, containing a papillose columella, composed of ≤ 20 granular and closely-packed rods.

Distribution

Regional: Eastern margin of South Africa, off Port St. Johns extending towards Durban; 96–695 m. Elsewhere: Indonesia (Cairns and Zibrowius 1997); Tanzania; Madagascar (Cairns and Keller 1993); 180–1016 m.

Remarks

Stephanocyathus (A.) explanans differs from the only other subgeneric extant species [S. spiniger (von Marenzeller, 1888)], in its calicular margin, number of septa, and costal spines. Cairns and Zibrowius (1997) elaborated on these taxonomic differences, highlighting that S. explanans may be distinguished in having a serrated calicular margin (lanceted in S. spiniger), 48–72 septa (instead of 96 septa), and 6–11 marginal circular spines (consistently 6 elongated and basally compressed in S. spiniger). Stephanocyathus explanans was first reported from South Africa by Gardiner and Waugh (1938), however no further locality information apart from noting that specimens were collected “off South Africa” was provided. Subsequently, Boshoff (1981) incorrectly identified the species as Stephanocyathus (O.) nobilis (von Marenzeller, 1904a) (ORI_D111k2), a record later corrected as S. explanans by Cairns and Keller (1993). The South African representatives add no new morphological information apart from S1 being slightly wider than S2 in some of the examined specimens.

Figure 5. 

A, F Stephanocyathus (Acinocyathus) explanans: A, C (USNM 62500, off Durban, 183–220 m) Calicular view B basal view C lateral view D, F (SAMC_A090144, off Port St Johns, 96–98 m) D calicular view E basal view F lateral view G, I Stephanocyathus (Odontocyathus) campaniformis (SAM_H3178, off Coffee Bay, 1420 m) G calicular view H basal view I lateral view J, L Stephanocyathus (Odontocyathus) nobilis (SAM_H1697, off Groot Berg Estuary, 1050 m) J calicular view K basal view L lateral view M, N Tethocyathus virgatus (SAMC_A073180, off Mazeppa Bay, 240–250 m) M calicular view N lateral view O, P Trochocyathus (Trochocyathus) sp. 1 (SAM_H1244, off East London, 59 m) O calicular view P lateral view. Scale bars: 10 mm.

Stephanocyathus (Odontocyathus) Moseley, 1881

Diagnosis

Stephanocyathus with 12–18 short basal spines or tubercles (C1–2, sometimes C3), sometimes fusing into a basal rim.

Type species

Platytrochus coronatus Pourtalès, 1867, by monotypy.

Stephanocyathus (Odontocyathus) campaniformis (von Marenzeller, 1904)

Fig. 5G–I

Stephanotrochus campaniformis von Marenzeller, 1904a: 302, pl. 18, figs 20, 20A.

Stephanocyathus campaniformis . –Gardiner and Waugh 1938: 189, 191. –Zibrowius 1980: 103. –Zibrowius and Gili 1990: 32–35, pl. 4, figs A–F, pl. 5, figs E–J

Stephanocyathus (Odontocyathus) campaniformis. –Cairns and Keller 1993: 243, fig. 5A, B.

Type locality

Off Walvis Ridge, Namibia (SS ‘Valdivia’ stn. 83: 25°25'03"S, 6°12'04"E); 981 m (von Marenzeller 1904a).

Type material

Two syntypes are deposited at the ZMB (Zibrowius and Gili 1990).

Material examined

SAM_H3178 (1 specimen): Eastern margin, 39 km from Coffee Bay/31 km off Hluleka Estuary, 32°00'42.00"S, 29°33'00.00"E; 1420 m.

Description

Corallum highly variable in shape, ranging from flat, cylindrical, bulbous, to bowl-shaped (as in the examined specimen) with rounded base. BD smaller than CD, and having a basal scar indicating point of attachment. Calice slightly elliptical (GCD:LCD = 1.1), with serrated calicular margin. Only specimen examined 29.8 × 27.5 mm in CD, 24.3 in BD, and 12.6 mm in H. Lower costae unevenly sized and spaced, with C1–2 prominent and extending from calicular margin to base, where they bear basal tubercles. C3–5 equal, separated by fine striae, and less distinct towards base. All costae covered with fine granules. Corallum white.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to the formula: S1–2 > S3 > S4 > S5 (≤ 80 septa). S1–2 highly exsert, and extend to columella with straight axial margins. Each S1–2 bears a palus separated from its respective septum by a deep notch. P1 fuse to columella. S3 less exsert and ¾ the width of S1–2, with a straight axial margin, and separated from respective P3 by a notch. When flanked by S4, S3 and S4 join deeper in fossa, meeting columella as one septum. S4 less exsert, ½ the width of S3, with straight to slightly dentate axial margins. S5 equally exsert as surrounding S4, but rudimentary in development. Septal faces covered with fine granules. Pali variable in size and shape. Fossa relatively shallow, but reported to be deep in larger specimens. An under-developed papillose columella present, usually a result of pali fusing.

Distribution

Regional: Eastern margin of South Africa, off Coffee Bay; 1420 m. Elsewhere: Madagascar Plateau (Cairns and Keller 1993); Walvis Ridge (off Namibia) (Zibrowius and Gili 1990); 882–1610 m.

Remarks

Among the Atlantic extant species of Stephanocyathus (Odontocyathus), S. campaniformis closely resembles S. nobilis (Moseley, 1873), but differs in having a considerably smaller adult size (7.5–11.5 mm) compared with that reported for S. nobilis (10.0–15.0) (Cairns and Keller 1993; Zibrowius and Gili 1990), and a higher H:GCD ratio (0.4–1.3) unlike that of S. nobilis (0.4–0.8 mm) (Cairns and Keller 1993; Zibrowius and Gili 1990). The resemblance of the Indian and Atlantic Ocean S. campaniformis representatives has been historically noted (Cairns and Keller 1993), and specimens reported herein add no taxonomic knowledge to this discussion. Specimens examined herein represent a new South African record, thus extending the Indian Ocean records from the Madagascar Plateau further southwards.

Stephanocyathus (Odontocyathus) nobilis (Moseley, 1873)

Fig. 5J–L

Ceratrochus nobilis Moseley in Thomson, 1873: 402, fig. 3. –Moseley in Thomson 1876: 554.

Stephanotrochus nobilis. –Moseley 1881: 155, pl. 3, fig. 3A–B. –Jourdan 1895: 20.

Stephanotrochus nitens Alcock in Wood-Mason & Alcock, 1891: 7–8.

Stephanotrochus platypus . –Jourdan 1895: 19, pl. 2, figs 14–16.

Stephanotrochus diadema var. nobilis. –Gravier 1920: 47–51, pl. 5, figs 80–86, pl. 6, figs 87–89, pl. 14, figs 205, 206.

Sabinotrochus opulens Gravier, 1920: 54, pl. 6, figs 101–103.

Stephanocyathus nobilis . –Gardiner and Waugh 1938: 189–192, pl. 6, figs 13, 15. –Pillai and Scheer 1976: 16. –Zibrowius 1980: 101–108, pl. 51, figs A–K.

Stephanocyathus (Odontocyathus) nobilis. –Cairns 1979: 110–111, pl. 20, figs 7, 10. –Cairns and Keller 1993: 242, fig. 5D, E.

Type locality

South of Flores, Azores (HMS ‘Challenger’ stn. 38°30'00"N, 31°14'00"W); 1830 m (Moseley 1876).

Type material

The holotype is deposited at the NHMUK (Zibrowius 1980).

Material examined

SAM_H1697 (1 specimen): Eastern margin, 58 km south of Ponta Do Ouro/46 km off Mgobezeleni Estuary, 27°21'18.00"S, 33°03'53.99"E; 1050 m.

Description

Corallum bowl-shaped with rounded base. BD smaller than CD. Base eroded but usually having a basal scar indicating point of previous attachment. Prominent basal tubercles corresponding to C1–2. Calice slightly elliptical (GCD:LCD = 1.03), with serrated calicular margin. Only specimen examined 41.3 × 40.0 mm in CD, 29.3 in BD, and 24.9 mm in H. Lower costae unevenly sized and spaced. C1–2 prominent and extending from calicular margin to base. Theca mostly smooth and glossy, with traces of costae, which change to 2–3 tubercles at basal inflection point, and progressively becoming less prominent at basal centre. C3–5 subequal, with thin and deep striae at calicular margin, becoming faint and slowly disappearing towards base. All costae covered by fine granules. Corallum white.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 > S5 (≤ 76 septa). S1–2 highly exsert and extend to columella with straight axial margins. P1–2 separated from their respective septum by a shallow notch. P1 reaches/fuses to columella. S3 less exsert and 3/4 the width of S1–2. S3 axial margins straight to slightly sinuous and bear a palus (P3). P3 sometimes joined by adjacent S4 deeper in fossa. S4 less exsert and ½ the width of S3, with straight axial margins. S5 rudimentary but as exsert as S4. All septal faces covered with fine granules. Pali variable in size, shape, and height. P1–3 form an inner crown encircling the well-developed papillose columella composed of closely packed rods. Fossa relatively deep, but reported to be shallow in smaller specimens.

Distribution

Regional: Eastern margin of South Africa, off Mgobezeleni Estuary (58 km south of Ponta Do Ouro: Mozambique); 1050 m. Elsewhere: Mozambique (Cairns and Keller 1993); off Zanzibar; off Kenya (Gardiner and Waugh 1938); Madagascar (Gardiner and Waugh 1938); Arabian Sea (Wood-Mason and Alcock 1891a); Maldives (Gardiner and Waugh 1938); off England; Azores; Gulf of Guinea (Zibrowius 1980); 609–2200 m.

Remarks

As mentioned in the remarks section of Stephanocyathus (O.) campaniformis, S. nobilis may be confused with S. campaniformis but differs in corallum size and height in relation to GCD. Both species have overlapping distributional patterns and are reported to occur in the Indian and Atlantic Ocean basins, this observation leading up to historical discussions around the validity of the Indian Ocean representatives as S. nobilis. However, Cairns and Keller (1993) agreed with Zou (1988) and confirmed that the Indian and Atlantic Ocean representatives are indeed one species of S. nobilis. The current Indian Ocean record represents a southern range extension from Mozambique and is a new record for South Africa.

Tethocyathus Kühn, 1933

Diagnosis

Corallum solitary, turbinate to trochoid, fixed or free. Septotheca covered by thick epitheca. Paliform lobes before all but last septal cycle in two distinct crowns. Columella papillose at top.

Type species

Thecocyathus microphyllus Reuss, 1871, by original designation.

Tethocyathus virgatus (Alcock, 1902)

Fig. 5M, N

Trochocyathus (Thecocyathus) virgatus Alcock, 1902a: 98–99. –Alcock 1902c: 16–17, pl. 2, fig. 13.

Tethocyathus virgatus. –Cairns 1995: 65–66, pl. 16C–F. –Cairns and Zibrowius 1997: 114–115. –Cairns 1999a: 86. –Cairns 2004a: 286. –Cairns 2009: 9. –Kitahara et al. 2010a: 115. –Kitahara and Cairns 2021: 570–571, 573, figs 316, 317A–C.

Type locality

Off Sulu Archipelago (HMS ‘Siboga’ stns. 96 and 105: South–east of Pearl Bank and 6°08'00"S, 121°19'00"E, respectively); 15–275 m (Alcock 1902c).

Type material

Two syntypes are deposited at the ZMA (Cairns 2004a).

Material examined

SAMC_A073083 (1 specimen): Eastern margin, 28 km from Richards Bay/40 km off Mlalazi Estuary, 29°00'54.00"S, 32°12'06.12"E; 215 m. SAMC_A073180 (2 specimens): Southern margin, 33 km from Mazeppa Bay/24 km off Cwili Estuary, 32°45'47.88"S, 28°36'24.12"E; 240–250 m. USNM 91674 (1 specimen): Eastern margin, 32 km from Port Dunford/37 km off Mlalazi Estuary, 29°05'10.79"S, 32 08'10.79"E; 95 m.

Description

Corallum solitary, ceratoid to subcylindrical, and attached to substrate by a thick pedicel (PD:GCD = 0.2–0.7) that expands into a polycyclic encrusting base. Calice elliptical (GCD:LCD = 1.1–1.2), with a jagged calicular margin. Largest specimen examined (USNM 91674) 13.4 × 11.8 mm in CD, and 26.3 mm in H. Costae unequal, flat, granular, separated by shallow intercostal striae, and becoming less prominent towards base. Epitheca may be thick and penetrated by lenticular pores of acrothoracid barnacles, or thin with no incrustation. Corallum pigmented purple-black, that fades into a white colouration towards base.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to the formula: S1 > S2 > S3 ≥ S4 > S5 (< 68 septa). Septa closely packed. S1–2 exsert, independent, thickest, and most swollen. Higher cycle septa (S3–4) progressively less exsert and less thick. S3 1/2 the width of S1–2. S4 dimorphic in size: those adjacent to S1 equal to or slightly wider than S3; and those adjacent to S2 usually less wide than S3. S5 same width as S4 adjacent to S1, if present. All septa have straight axial margins except for S3, which have slightly sinuous ones. Septal faces granular. Pali large, closely packed, and present before all septal cycles but last, forming a high paliform crown encircling a papillose columella composed of a group of closely-packed and low-profile rods.

Distribution

Regional: Southern to eastern margin of South Africa, from Kei Mouth extending towards Port Dunford; 95–250 m. Elsewhere: Philippines; Indonesia (Cairns and Zibrowius 1997); Vanuatu; Wallis and Futuna Islands (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); New Zealand (Cairns 1995); Australia (Cairns 2004a); 95–1200 m.

Remarks

Tethocyathus virgatus is distinguished from the other Recent congeners in having a pigmented CS1, S1–2 being the thickest septa, well-developed lamellar pali, and by attaining the largest corallum size. The examined specimens represent a new record for the region. The South African representative differs from the Australian in having septa of the fifth cycle (< 68 septa) as compared with only four (48 septa), and in having S3 that bear a slightly sinuous rather than a straight axial margin. Nonetheless, all the other taxonomic features agree with what is known to be characteristic of the species.

Trochocyathus Milne-Edwards & Haime, 1848

Diagnosis

Corallum solitary, turbinate to ceratoid, or bowl-shaped, fixed or free. Transverse division may be present. Septotheca costate, sometimes covered with a thin epitheca. Pali before all but last cycle of septa. Columella papillose.

Type species

Turbinolia mitrata Goldfuss, 1827, by subsequent designation (Milne-Edwards and Haime 1850b).

Trochocyathus (Trochocyathus) Milne-Edwards & Haime, 1848

Diagnosis

Trochocyathus lacking basal costal spines and with other than discoidal coralla.

Type species

Turbinolia mitrata Goldfuss, 1827, by subsequent designation (Milne-Edwards and Haime 1850b).

Trochocyathus (Trochocyathus) sp. 1

Figs 5O, P, 6A, B

Trochocyathus rawsonii. –Gardiner 1904: 100–103, 124, pl. 1, fig. 2A, B, pl. 2, fig. D, E, G, H, J, K . –Cairns and Keller 1993: 241.

Caryophyllia gigas. –Boshoff 1981: 36.

Endopachys grayi. –Boshoff 1981: 42 (in part).

Material examined

SAMC_A073233 (2 specimens): Southern margin, 14 km from Cape Point/10 km off Buffels Wes Estuary, 34°13'59.99"S, 18°30'00.00"E; 42 m. SAM_H1244 (1 specimen): Southern margin, 7 km from East London/5 km off Buffalo Estuary, 33°02'59.99"S, 27°56'59.99"E; 59 m. SAM_H1449 (1 specimen): Southern margin, 14 km from Mazeppa Bay/20 km off Great KeiEstuary, 32°34'00.00"S, 28°33'00.00"E; 174 m. SAM_H3115 (1 specimen): Southern margin, 2 km off Mosselbaai/10 km off Hartenbos Estuary, 34°10'37.57"S, 22°09'19.14"E; 55 m. SAM_H3117 (1 specimen): Southern margin, 246 km from Mazeppa Bay/243 km off Mendu Estuary, 33°43'11.99"S, 30°48'47.99"E; 780 m. SAM_H3177 (2 specimens): Southern margin, 15 km from Port Alfred/11 km off Riet Estuary, 33°39'18.00"S, 27°11'35.99"E; 90 m. SAM_H3833 (1 specimen): Eastern margin, 34 km from Coffee Bay/7 km off Ntlonyane Estuary, 32°15'11.99"S, 28°57'42.00"E; 47 m. SAM_H3834 (1 specimen): Southern margin, 23 km from Port Elizabeth/22 km off Bakens River Estuary, 33°50'41.99"S, 25°47'30.00"E; 36 m. ORI_DIIIa4_3 (1 specimen): Locality data unknown, 300 m. ORI_EId1 (2 specimens): Eastern margin, other locality data unknown. USNM 77220 (3 specimens): Eastern margin, 28 km from Coffee Bay/19 km off Bulungulu Estuary, 32°14'53.99"S, 29°10'23.99"E; 620–560 m.

Imagery data

BMNH 1950.03.22.17 (1 specimen), BMNH 1950.01.10.112 (1 specimen), NHMUK 1970.01.26.11–20 (2 specimens): South Africa, other locality data unknown. Mortensen Java Expedition (12 specimens): Eastern margin, off Durban, 128 m.

Description

Corallum solitary, ceratoid to trochoid, mostly attached through a variably sized pedicel (PD:GCD = 0.3–0.7). Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), calicular margin slightly serrated. Largest specimen examined (ORI_DIIIa4_3) 11.1 × 10.1 mm in CD, 12.0 mm in H, and 6.6 mm in PD. Costae prominent from calicular margin to base, similar in width to associated septa, equidistant, low, bearing small granules, and separated by thin intercostal furrows. Corallum white.

Septa hexamerally arranged in four cycles according to the formula: S1 ≥ S2 > S3 > S4 (48 septa); sometimes a pair of S5 present in a half-system (e.g., ORI_EId1). S1 most exsert, equal or slightly wider than S2, with straight or slightly sinuous axial margin. S2 ¼ wider than S3 with moderately sinuous axial margin. S3 as sinuous as S2. S4 rudimentary and bearing dentate axial margin. S3–4 progressively less exsert than S2. Pali present before all but last septal cycle. P2–3 joining deeper in fossa, forming thick chevrons before S2. Pali tall, being distinctively higher than columellar elements. All septal and palar faces covered with small granules. Fossa moderately deep, containing a papillose columella composed of a group of 6–18 loosely-packed, low-profile rods.

Distribution

Regional: Southern to eastern margin of South Africa, from off Cape Point towards Coffee Bay; 36–780 m.

Remarks

The validity of Gardiner’s (1904) specimens as Trochocyathus (T.) rawsonii has long been questioned due to their well-developed costae, slender pedicel, and rarely bearing S5 (Cairns 1979; Zibrowius and Gili 1990; Cairns and Keller 1993). The examined material conforms to these characteristics and closely resembles Gardiner’s (1904) illustrations. Nevertheless, the specimens reported herein, together with Gardiner’s (1904), might represent a new species. However, as Trochocyathus have great intraspecific variation, a formal description is postponed. Nonetheless, these new records extend the known distribution of this taxon towards the eastern margin of South Africa and also expand its known depth range (both upper and lower).

Trochocyathus (Trochocyathus) sp. 2

Fig. 6C–F

Material examined

DEFF/SAEON_A33997 (1 specimen): Southern margin, 39 km from Cape Padrone/40 km off Boknes Estuary, 34°03'53.52"S, 26°42'11.58"E; 100 m. DSCS–INV 529 (1 specimen): Southern margin, 35 km from Cape St. Francis/70 km off Slang Estuary, 34°47'05.0"S, 24°45'42.3"E; 392–418 m SAMC_A073141 (3 specimens): Eastern margin, 33 km from Coffee Bay/18 m off Ntlonyane Estuary, 32°16'41.88"S, 29°06'00.00"E; 300 m. SAMC_A073150 (3 specimens): Southern margin, 26 km from Mazeppa Bay/33 km off Great Kei Estuary, 32°41'12.12"S, 28°43'54.12"E; 480–490 m. SAMC_A073166 (1 specimen): Eastern margin, 19 km off Port St. Johns/21 km off Bulolo Estuary, 31°45'00.00"S, 29°40'47.99"E; 125 m. SAMC_A073175 (1 specimen): Eastern margin, 28 km from Coffee Bay/16 km off Hluleka Estuary, 31°55'58.79"S, 29°25'12.00"E; 300 m. SAMC_A073211 (3 specimens): Eastern margin, 5 km from Cape Vidal/16 km off St Lucia Estuary, 28°08'24.00"S, 32°36'24.00"E; 165 m. SAMC_A073235 (2 specimens): Southern margin, False Bay; depth unknown. SAMC_A088920 (1 specimen): Southern margin, 38 km from Cintsa/22 km off Cwili Estuary, 32°50'30.72"S, 28°30'41.33"E; 250–226 m. SAMC_A088921 (3 specimens): Southern margin, 140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A088925 (1 specimen): Southern margin, 140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A088926 (8 specimens): Southern margin, 140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAMC_A090146 (5 specimens): Southern margin, 68 km from Cape St. Francis/70 km off Slang Estuary, 34°47'35.77"S, 24°38'35.69"E; 520 m. SAMC_A090147 (2 specimens): Southern margin, 140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. SAM_H1243 (3 specimens): Southern margin, 11 km from Infanta/12 km off Duiwenhoks Estuary, 34°27'00.00"S, 20°58'00.00"E; 47 m. SAM_H1370 (1 specimen): Southern margin, 28 km from Gonubie/27 km off Buffalo Estuary, 33°09'29.99"S, 28°03'06.00"E; 86 m. SAM_H1384 (1 specimen): Southern margin, 2 km from Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 95 m. SAM_H1388 (14 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H1391 (1 specimen): Southern margin, 24 km from Cape Padrone/35 km off Boknes Estuary, 33°58'00.00"S, 26°21'00.00"E; 46 m. SAM_H1400 (1 specimen): Southern margin, 74 km off Cape St. Francis/83 km off Slang Estuary, 34°52'00.00"S, 24°56'59.99"E; 137 m. SAM_H1407 (1 specimen): Southern margin, 2 km from Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 192 m. SAM_H1408 (1 specimen): Southern margin, 14 km from Mazeppa Bay/20 km off Great Kei Estuary, 32°34'00.00"S, 28°33'00.00"E; 174 m. SAM_H1419 (1 specimen): Southern margin, 4 km from Plettenberg Bay/7 km off Piesang Estuary, 34°06'00.00"S, 23°23'59.99"E; 146 m. SAM_H1443 (26 specimens): Southern margin, 241 km from Agulhas/247 km off De Mond-Heuningnes Estuary, 36°40'00.00"S, 21°25'59.99"E; 200 m. SAM_H1487 (13 specimens): Southern margin, 2 km from Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 229 m. SAM_H3103 (2 specimens): Southern margin, 2 km from Mosselbaai/11 km off Hartenbos Estuary, 34°11'10.12"S, 22°09'40.59"E; 165–183 m. SAM_H3106 (9 specimens): Southern margin, 846 km from Port St. Johns/842 km off Mkweni Estuary, 36°40'00.00"S, 21°25'59.99"E; 366 m. SAM_H3111 (1 specimen): Eastern margin, 18 km from Cape Vidal/27 km off Mfolozi Estuary, 28°16'18.00"S, 32°38'48.00"E; 670 m. SAM_H3114 (2 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3116 (9 specimens): Eastern margin, 22 km from Cape Vidal/28 km off Mfolozi Estuary, 28°17'23.99"S, 32°40'47.99"E; 550 m. SAM_H3170 (15 specimens): Eastern margin, 28 km from Coffee Bay/19 km off Bulungulu Estuary, 32°14'53.99"S, 29°10'23.99"E; 620–560 m. SAM_H3172 (3 specimens): Southern margin, 29 km from Mazeppa Bay/25 km off Kobole Estuary, 32°29'30.00"S, 28°57'06.00"E; 650–700 m. SAM_H3173 (1 specimen): Southern margin, 47 km from Port Alfred/14 km off Mgwalana Estuary, 33°30'18.00"S, 27°22'05.99"E; 80 m. SAM_H3175 (1 specimen): Eastern margin, 30 km from Coffee Bay/20 km off Bulungulu Estuary, 32°15'00.00"S, 29°09'06.00"E; 520–500 m. SAM_H3176 (3 specimens): Eastern margin, 30 km from Scottburgh/20 km off Fafa Estuary, 30°33'24.00"S, 30°48'35.99"E; 690 m. SAM_H3830 (2 specimens): Western margin, 49 km from Cape Point/51 km off Buffels Wes Estuary, 34°43'18.00"S, 18°12'29.99"E; 360–365 m. SAM_H3832 (3 specimens): Southern margin, 14 km from Cape Point/10 km off Buffels Wes Estuary, 34°13'59.99"S, 18°30'00.00"E; 42 m.

Description

Corallum ceratoid to trochoid, and attached to a variably sized pedicel (PD:GCD = 0.2–0.7). Calice circular to slightly elliptical (GCD:LCD = 1.0–1.2), with serrate calicular margin. Largest specimen examined (SAM_H1400) 17.4 × 17.2 mm in CD, 6.0 mm in PD, and 39.6 mm in H. Costae prominent from calicular margin to base, ca. the same width as associated septa, equidistant, low, bearing small granules, and separated by thin intercostal furrows. Corallum white.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to formula S1 > S2 ≥ S3 > S4 > S5 > S6 (≤ 60 septa). S1 most exsert, independent, slightly wider than S2, and bearing straight to slightly sinuous axial margin. S2 slightly less exsert than S1, and extending slightly less to centre of calice with moderately sinuous axial margin. S3 equal or slightly smaller than S2, but have axial margin equally sinuous. S4 predominantly restricted to upper calice in specimens with four septal cycles, but dimorphic and variable in development half-systems bearing S5 or S5–6 (e.g., SAMC_A090146 = 6:6:12:24:10:2, SAM_H1407 = 6:6:12:24:12). In the latter, S4 closer to S1 have the same rudimentary profile as S5 and S4 flanked by S5 having a similar profile as S3 or slightly less wide. S4 axial margin coarsely dentated. If present, S6 restricted to calicular margin. S3–5 progressively less exsert than S1–2. Pali present before all but last septal cycle. P2 most recessed. P2–3 form a chevron-like pattern before S2, a pattern also taken by S4 flanked by S5 which sometimes bear pali. All septal and palar faces granulated. Fossa moderately deep, containing a papillose columella composed of a group of 17–30 rod elements which mostly are as high as pali elements.

Distribution

Regional: Western to eastern margin of South Africa, off Cape Point extending towards Cape Vidal; 42–700 m.

Remarks

Although specimens examined closely resemble Trochocyathus (T.) sp. 1 cf. T. rawsonii sensu Gardiner (1904) in having well-developed costae extending from the calicular margin to base, an often curved coralla, and the presence of S5, Trochocyathus sp. 2 occasionally has a pair of S6, even though the fifth cycle is incomplete (SAMC_A090146 = 6:6:12:24:10:2), and an independent S1 which extends further to the columella. Apart from that, the specimens reported herein differ from Trochocyathus (T.) sp. 1 cf. T. rawsonii sensu Gardiner (1904) in having a systematic development of septal cycles (irrespective of an increase mm in CD). While this material may represent a new species of Trochocyathus, a name will not be introduced until the intraspecific variation is better understood and compared with that from congeners.

Figure 6. 

A, B Trochocyathus (Trochocyathus) sp. 1 (SAM_H1449, off Mazeppa Bay, 174 m) A calicular view B lateral view C, F Trochocyathus (Trochocyathus) sp. 2 C, D (SAM_H1388, East London, 90 m) C calicular view C lateral view E, F (SAM_H1407, off Mosselbaai, 192 m) E calicular view F lateral view G, L Trochocyathus (Trochocyathus) sp. cf. rawsonii sensu Cairns & Keller, 1993 G, J (SAM_H1440, off East London, 90 m) G calicular view H lateral view I calicular view J lateral view. 1993 K, L (SAMC_A090156, off Sedgefield, 74 m) K calicular view L lateral view M, P Trochocyathus (Trochocyathus) sp. 3 M, N (SAM_H3124, off East London, 90 m) O calicular view P lateral view O, P (BMNH 1939.7.20.47, off Richards Bay, 165 m) O calicular view P lateral view. Lateral view. Scale bars: 10 mm.

Trochocyathus Trochocyathus sp. cf. rawsonii sensu Cairns in Cairns & Keller, 1993

Fig. 6G–L

Trochocyathus rawsonii sensu Cairns 1979: 77, fig. 6, pl. 13. –Cairns and Keller 1993: 241–242, figs 4E, H.

Type locality

Off Barbados, eastern Caribbean Island (USCSS ‘Hassler’); 183 m (Cairns 1979).

Type material

Three syntypes are deposited at the MCZ (Cairns 1979).

Material examined

DEFF/SAEON_A32823 (1 specimen): Southern margin, 131 km from Gouritsmond/off Goukamma Estuary, 35°14'57.1"S, 22°50'48.1"E; 511 m. SAMC_A073015 (1 specimen): Southern margin, 32 km from Mazeppa Bay/19 km off Mendu Estuary, 32°25'00.11"S, 28°58'18.11"E; 330–340 m. SAMC_A073206 (2 specimens): Eastern margin, 36 km from Richards Bay/49 km off Nhlabane Estuary, 29°02'12.11"S, 32°19'36.12"E; 760–800 m. SAMC_A073220 (1 specimen): Southern margin, 13 km from Port St. Johns/12 km off Bulolo Estuary, 31°44'48.12"S, 29°33'00.00"E; 370 m. SAMC_A090091 (1 specimen): Eastern margin, 152 km from Margate/off Bilanhlolo Estuary, 31°39'07.00"S, 29°39'42.00"E; 300–540 m. SAMC_A090156 (3 specimens): Southern margin, 14 km from Sedgefield/off Cunge Estuary, 34°09'17.46"S, 22°48'37.32"E; 74 m. SAM_H1360 (1 specimen): Southern margin, 11 km from East London/3 km off Buffalo Estuary, 33°00'53.67"S, 27°55'50.67"E; 128 m. SAM_H1415 (1 specimen): Southern margin, 8 km from Pringle Bay/6 km off Buffels Oos Estuary, 34°17'55.37"S, 18°49'10.85"E; 33 m. SAM_H1440 (3 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3110 (26 specimens): Eastern margin, 18 km from Cape Vidal/27 km off Mfolozi Estuary, 28°16'18.00"S, 32°38'48.00"E; 670 m. SAM_H3113 (2 specimens): Southern margin, 11 km from East London/3 km off Buffalo Estuary, 33°00'53.67"S, 27°55'50.67"E; 128 m. SAM_H3171 (8 specimens): Southern margin, 40 km from Cintsa Mouth/29 km off Cwili Estuary, 32°55'00.00"S, 28°31'00.00"E; 630 m. SAM_H3174 (1 specimen): Southern margin, 22 km from Gonubie/21 km off Gqunube Estuary, 33°06'00.00"S, 28°08'17.99"E; 700–650 m. SAM_H3829 (1 specimen): Southern margin, 98 km from Gansbaai/103 km off Buffels Oos Estuary, 35°15'18.00"S, 18°39'18.00"E; 547 m.

Description

Corallum ceratoid to trochoid, attached to a narrow pedicel (PD:GCD = 0.3–0.5). Calice circular to slightly elliptical (GCD:LCD = 1.0–1.3), calicular margin slightly serrate. Largest specimen examined (SAM_H3113) 12.2 × 11.2 mm in CD, 4.9 mm in PD, and 17.5 mm in H. Costae prominent and bearing small granules near calicular margin, progressively becoming narrower towards base. Costae separated by deep furrows. Corallum white, but sometimes purplish with white columella.

Septa hexamerally arranged in four cycles according to the formula: S1 ≥ S2 > S3 > S4 (48 septa). S1–2 usually equally exsert, but sometimes S2 may be slightly less exsert than S1. S1 meets columella with straight to slightly sinuous axial margins. S1 slightly wider to equal in width to S2. S2 axial margins straight to slightly sinuous. Higher cycle septa (S3–4) progressively less exsert. S3 ~ ¼ less wide than S2, with moderately sinuous axial margins. S4 ½ the width of S3, and bear extremely sinuous and sometimes dentate axial margins. Pali present before all but last septal cycle, being taller than columellar elements, slightly sinuous, and thick, with some half-systems having P2–3 joining in front of S2 before extending towards columella. All septal and palar faces being granulated. Fossa moderately deep, containing a papillose columella composed of a group of low-profile and compact intertwined rods.

Distribution

Regional: Southern to eastern margin of South Africa, from off Pringle Bay extending towards Cape Vidal; 33–800 m. Elsewhere: north-western Madagascar and off Walter Shoal (Cairns and Keller 1993); 760–780 m.

Remarks

Although specimens resemble Gardiner’s (1904) Trochocyathus (T.) sp. in having four cycles with the following septal formula S1 ≥ S2 > S3 > S4, they differ in S4 consistently bearing sinuous to dentate axial margins unlike the exclusively dentate appearance observed in Gardiner’s (1904) representatives. The level of granulation on septal and palar faces may also distinguish the two forms, of which Trochocyathus (T.) sp. cf. T. rawsonii sensu Cairns and Keller (1993) has low coverage appearance (with large granules prominent along the septal faces) as compared with the fine and dense granules throughout both the pali and septa (giving specimens a rough appearance) in Gardiner’s (1904) Trochocyathus forms. Furthermore, the pali are compact in Trochocyathus (T.) sp. cf. T. rawsonii sensu Cairns and Keller (1993) rather than being loosely packed (Trochocyathus (T.) sp. 1 cf. T. rawsonii sensu Gardiner (1904)). However, some specimens of Trochocyathus (T.) sp. cf. T. rawsonii sensu Cairns and Keller (1993) diverge from the typical compact pali, thus bearing a tall and thin pali (SAM_H1415 and SAMC_A090156).

Trochocyathus (Trochocyathus) sp. 3

Fig. 6M–P

Caryophyllia berteriana. –Gardiner 1904: 112–113.

Trochocyathus (Trochocyathus) sp. –Zibrowius 1982: 114, pl. 1, figs 1, 2.

Trochocyathus (Trochocyathus) sp. A. –Cairns and Keller 1993: 240, fig. 4C, D.

Material examined

SAMC_A073157 (1 specimen): Eastern margin, 10 km from Port Edward/24 km off Bilanhlolo Estuary, 31°05'48.11"S, 30°18'47.88"E; 140 m. SAMC_A073213 (1 specimen): Eastern margin, 29 km from Durban/14 km off Mbokodweni Estuary, 30°06'24.12"S, 31°00'47.88"E; 160–170 m. SAM_H3124 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. USNM 91530 (3 specimens): Eastern margin, 37 km south of Ponta Do Ouro/23 km off Kosi Bay Estuary, 27°11'03.59"S, 32°50'32.39"E; 100 m. USNM 91566 (1 specimen): Eastern margin, 42 km south of Ponta Do Ouro/27 km off Kosi Bay Estuary, 27°13'30.00"S, 32°49'30.00"E; 74 m.

Imagery data

BMNH 1939.7.20.47 (1 specimen): Eastern margin, 20 km from Richards Bay/8 km off Nhlabane Estuary, 28°40'00.00"S, 32°15'00.00"E; 165 m.

Description

Corallum ceratoid, mostly attached to substrate through a narrow pedicel (PD:GCD = 0.4–0.6) and a thin encrusting base. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), calicular margin slightly serrate. Largest specimen examined (USNM 91566) 8.2 × 7.3 mm in CD, and 9.0 mm in H. Costae prominent and bearing small granules near calicular margin, and progressively disappear towards pedicel. Theca becomes transversely ridged beyond 3 mm from calicular margin. Corallum white, and sometimes light brown around calicular margin.

Septa hexamerally arranged in four cycles according to the formula: S1 ≥ S2 > S3 ≥ S4 (48 septa). However, largest specimen (USNM 91566) displays S1–2 > S4–3. S1 most exsert and extend slightly further into centre of calice than S2. S1 axial margins straight to slightly sinuous. S2 equally wide or ¼ less wide than S1, with moderately sinuous axial margins. Higher cycle septa (S3–4) progressively less exsert (if at all). S3 with most sinuous axial margins. S4 rudimentary and bear straight to slightly sinuous axial margins. Pali present before all but last septal cycles. P2–3 joining deeper in fossa, forming a chevron-like structure before S2. Septal and palar faces finely granulated. Fossa moderately deep, pali indistinguishable from the closely-packed rods of the papillose columella.

Distribution

Regional: Southern to eastern margin of South Africa, from East London extending towards Kosi Bay Estuary (37 km south of Ponta Do Ouro: Mozambique); 74–170 m. Elsewhere: Mozambique, Madagascar, and Saya de Malha Bank; 74–315 m (Cairns and Keller 1993).

Remarks

The examined specimens closely resemble Trochocyathus (T.) sp. A reported by Cairns and Keller (1993) in all features, except for its septa all having sinuous axial margins instead of only S3 having sinuous axial margins. Zibrowius (1982) also noted a Trochocyathus specimen (BMNH 1939.7.20.47) that resembles Cairns and Keller’s (1993) specimens in all septa being straight except S3 and in septa being arranged according to S1–2 > S4–3. The representative images of BMNH 1939.7.20.47 unfortunately do not show the tertiary septa’s axial margin, and for that reason this Trochocyathus specimen is added to the records of species accounted for here. Nevertheless, a new name will not be introduced until the variation within Trochocyathus (T.) rawsonii is better understood.

Vaughanella Gravier, 1915

Diagnosis

Corallum solitary, patellate to trochoid, and usually firmly attached by a robust pedicel. Septotheca costate. Paliform lobes present on all but last septal cycle. Columella papillose.

Type species

Caryophyllia margaritata Jourdan, 1895, by monotypy.

Vaughanella concinna Gravier, 1915

Fig. 7A–D

Vaughanella concinna Gravier, 1915: 10. –Gravier 1920: 63, pl. 9, fig. 138–143. –Zibrowius 1980: 104–105, pl. 52, figs A–K, pl. 53, figs A–L. –Cairns 1999a: 90–91, fig. 11G, H. –Cairns et al. 1999: 25. –Cairns 2009: 12. –Kitahara and Cairns 2021: 439–440, figs 237D–F, 238.

Vaughanella oreophila. –Cairns 1995: 70, pl. 18, figs D, E.

Type locality

South of São Jorge, Azores (‘Prince de Monaco Expedition’ stn. 1349: 38°35'30"N, 28°05'45"W); 1250 m (Gravier 1920),

Type material

The lectotype and 6 paralectotypes are deposited at the MOM (Zibrowius 1980).

Material examined

SAMC_A072973 (1 specimen): Locality data unknown; 517 m.

Imagery data

MN_SM134 (1 specimen): Eastern margin, 36 km off Port Shepstone/49 km off Mtentu Estuary, 30°43'11.99"S, 30°48'47.99"E; 900 m.

Description

Corallum trochoid, robust, and firmly attached to substrate through a thick and reinforced pedicel (PD:GCD = 0.40–0.5) and encrusting base. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), sometimes slightly flared, with serrated calicular margin. Largest imaged specimen (MN_SM134) 20.0 × 18.0 mm in CD, 8.0 mm in PD, and 19.0 mm in H. Costae well-developed, but unequal in width. C1–2 wider than remaining costae. C3–5 progressively narrower. Intercostal striae, broad. All costae finely granular. Corallum white.

Septa hexamerally arranged in five cycles, the last cycle being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 > S5 (only one pair of S5 present in the specimens examined = 50 septa). S1–2 highly exsert, extending towards columella with straight axial margins. Each S1–2 bears a pointed and tall palus. P1 fuses to columellar elements. S3 less exsert and ¾ the width of S2, bearing tall and well-defined pali, which are separated from septa by a wide notch. S4 less exsert and dimorphic in size: generally, ¼ the width of S3 (S4 rudimentary in SAMC_A072973) but same size as S3 when flanked by S5. S5 restricted to calicular margin. All septa have straight axial margins, except for S3 which have straight to slightly sinuous axial margins. Septal faces slightly granular. Pali smooth, indistinguishable from columellar elements, except for P3 which are thicker than P1–2 and rise higher in fossa. Fossa moderately deep, containing a papillose columella composed of a compact group of rods.

Distribution

Regional: Eastern margin of South Africa, off Port Edward; 517–900 m. Elsewhere: Wallis and Futuna (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); New Zealand (Cairns 1995); Celtic Sea, Azores, and Madeira Archipelago (Zibrowius 1980); 500–3018 m.

Remarks

Both specimens reported (one of which is represented by imagery data) meet the known taxonomic distinction of Vaughanella, particularly V. concinna in having all higher cycle septa (S1–3) bearing pali (P1–3). However, they differ in costae extending from calicular margin towards base, as compared with theca being smooth and porcelaneous (Cairns 1995). The presence of P3 and one palus per septa distinguishes this species from the other two Pacific congeners, one of which lacks P3 (V. oreophila Keller, 1981) and the other has multiple lobes per septa (V. multipalifera Cairns, 1995). The South African representative (SAMC_A072973) displays convergence of S4 towards S3, a feature reported in the Atlantic forms, but differs in pali being smooth as compared with being granulated (Zibrowius 1980). Nonetheless, regional distribution is based on MN_SM134 (imagery data) as SAMC_A072973 lacks locality information.

Figure 7. 

A, D Vaughanella concinna A, B (MN_SM134, off Port Edward, 900 m) A calicular view B lateral view C, D (SAMC_A072973, locality data unknown, 517 m) C calicular view D lateral view E, F Deltocyathus rotulus (USNM 91550, off Scottburgh, 1360 m) E calicular view F lateral view G, H Atlantia denticulata sp. nov. (SAMC_A090157, off Gouritsmond, 170 m) G close-up of corallites H full view I, J Balanophyllia (Balanophyllia) bonaespei (USNM 1423303, off the Agulhas, 32 m) I calicular view J lateral view K, N Balanophyllia (Balanophyllia) capensis K, M (SAM_H3048, off Cape Point, depth unknown) calicular view L, N (BMNH_ 1939.7.20.479-500, locality data unknown) lateral view O, P Balanophyllia (Balanophyllia) diademata (SAMC_A073016, off Richards Bay, 500 m) O calicular view P Lateral view. Scale bars: 10 mm (A–H, K–P); 100 mm (I, J).

Family Deltocyathidae Kitahara, Cairns, Stolarski & Miller, 2012

Deltocyathus Milne-Edwards & Haime, 1848

Diagnosis

Solitary, discoidal to patellate, usually free. Septotheca costate. Septa arranged in 4–5 cycles, only S1 being independent. Pali before all but last cycle. Axial margins of higher cycle pali join to faces of adjacent septa (deltas). Columella papillose.

Type species

Turbinolia italica Michelotti, 1838, by monotypy. 181

Deltocyathus italicus (Michelotti, 1838)

Turbinolia italica Michelotti, 1838: 51, pl. 1, fig. 8.

Deltocyathus agassizii . –Moseley 1876: 546, 551 (in part).

Deltocyathus italicus . –Pourtalès 1880: 101 (in part), pl. 1, figs 2, 3. –Moseley 1881: 145–147 (in part). –von Marenzeller 1904a: 281 (in part). –Gravier 1920: 34–36 (in part). –Keller 1975: 177, pl. 2, figs 1–4B. –Kitahara 2007: 502–503. –Kitahara and Cairns 2009: 236.

Deltocyathus sp. cf. D. italicus. –Cairns 1979: 95–97, pl. 17, figs 1–3. –Reyes et al. 2009: 16, fig. 3E, F.

Deltocyathus conicus Zibrowius, 1980: 83–85, pl. 39, figs A–L. –Zibrowius and Gili 1990: 35, pl. 2, figs M–O.

Type locality

Tortona, Italy (Miocene) (Cairns 1979).

Type material

The holotype is lost (Cairns 1979).

Material examined

None.

Distribution

Regional: Western margin of South Africa, off Alexander Bay and Port Nolloth; 882–1412 m (Zibrowius and Gili 1991). Elsewhere: widespread in the Caribbean and Gulf of Mexico (Cairns 1978, 1979; Cairns et al. 2009); off Rio de Janeiro in Brazil; Bermuda (Cairns 1979; Kitahara 2007); Colombia (Reyes et al. 2009); Gulf of Gasco; Azores; Morocco; Miocene of Italy; Gulf of Guinea; Angola (Zibrowius 1980); 403–2634 m.

Remarks

This species is well described by Cairns (1979) and Zibrowius (1980), who both noted that the corallum is patellate and free, with a prominent conical base bearing no basal scar. The costae are ridged, unequal in size, and with a dentate appearance. The regional occurrence of Deltocyathus italicus is based on the Zibrowius and Gili (1991) record as no other specimens have been examined. Apart from the inter-species variation in the H:D ratio and columella development, highlighted by Zibrowius and Gili (1991), D. italicus may be differentiated from the other Atlantic species of Deltocyathus in having a strongly conical base (Cairns 1979b; Zibrowius 1980).

Deltocyathus rotulus (Alcock, 1898)

Fig. 7E, F

Trochocyathus rotulus Alcock, 1898: 16, pl. 2, figs 1, 1A.

Deltocyathus fragilis Alcock, 1902a: 99, 100. –Alcock 1902c: 21, pl. 1, figs 15, 15A.

Deltocyathus rotulus. –van der Horst 1931: 6. –Gardiner and Waugh 1938: 196. –Yabe and Eguchi 1937: 129. –Keller 1982: 50. –Cairns and Keller 1993: 245, pl. 5, fig. I. –Cairns 1994: 55, 56, pl. 24, figs J, K. –Cairns and Zibrowius 1997: 125, 126, fig. 16A–C. –Cairns 1999a: 91–92. –Cairns 2004a: 280. –Kitahara and Cairns 2009: 238, fig. 1B. –Kitahara and Cairns 2021: 399, 401, figs 204, 216A–D.

Type locality

North of Maldive Atoll, Flores Seas (HMS ‘Investigator’ stn. 216: 7°24'S, 118°15.2'E); 794 m (Alcock 1898; Kitahara and Cairns 2009).

Type material

The holotype is presumed to be deposited at the IM (Cairns 2004a; Kitahara and Cairns 2009).

Material examined

USNM 91550 (1 specimen): Eastern margin, 28 km from Scottburgh/21 km off Mkomazi, 30°11'59.99"S, 32°01'00.00"E; 1360 m.

Description

Corallum discoidal, unattached, with a flat to slightly bowl-shaped base. Calice slightly elliptical (GCD:LCD = 1.2), with a lanceted and scalloped calicular margin. Specimen examined 30.4 × 25.5 mm in CD. All costae serrated and prominent at calicular margin. Intercostal spaces deep and wider at calicular margin, becoming progressively narrower towards base. Costae granular, resulting in a serrated costal margin. Corallum white, being light reddish brown around columella.

Septa hexamerally arranged in five cycles according to the formula: S1–2 > S3 > S4 > S5 (total of 96 septa). S1–2 appearing independent septa, but fusing to neighbouring septa through porous connections. S1–2 moderately exsert, bear pali, each of which is separated from its septum by a notch. S3 ¾ the width of S1–2, less exsert, and sometimes each bearing a small palus. S4 less wide than S3, but equally exsert, and each bearing a tall palus, which form a distinctive crown around columella. S5 rudimentary. All septa fuse to adjacent septa, but position of fusion varies: S1–4 join neighbouring septa near columella whilst S5 join S4 near calicular margin. P1–3 small, with P1–2 being ¼ or ½ the size of P4, and P3 consistently half the size of P1–2, positioned at porous fusions. P4 positioned further away from columella, alluding to a crown of 24 pali. Upper margins of all septa smooth. Fossa shallow, containing a papillose columella composed of irregularly shaped papillae.

Distribution

Regional: Eastern margin of South Africa, off Scottburgh; 1360 m. Elsewhere: Mozambique; Zanzibar (Cairns and Keller 1993); Maldives (Alcock 1898; Gardiner and Waugh 1938); Gulf of Aden (Gardiner and Waugh 1938); Sri Lanka (van der Horst 1931); Tanzania (Gardiner and Waugh 1938); Japan (Cairns 1994); Philippines; Indonesia; Malaysia (Alcock 1902c; Cairns and Zibrowius 1997); Vanuatu; Wallis, and Futuna region (Cairns 1999a); Australia (Cairns 2004a); New Caledonia (Kitahara and Cairns 2009, 2021); 210–2340 m.

Remarks

Among the Deltocyathus s from South Africa, the Indo-Pacific D. rotulus differs from the Atlantic D. italicus in having a large and discodial corallum, a total of five cycles of septa (the highest cycle sometimes incomplete), costae giving a serrated appearance, and the reddish brown pigmentation around the columella.

Family Dendrophylliidae Gray, 1847

Atlantia López & Capel, 2020

Diagnosis

Colonies bushy, phaceloid to dendroid, achieved by extra-tentacular budding (frequently from theca of a parent corallite at an acute angle). No epitheca. Septa normally arranged and granular. Columella poorly to moderately developed. Gender: feminine.

Type species

Atlantia caboverdiana (Ocaña & Brito, 2015), by subsequent designation (Capel et al. 2020).

Remarks

Although Atlantia resembles Cladopsammia Lacaze-Duthiers, 1897; Astroides Quoy & Gaimard, 1827; Enallopsammia Sismonda, 1871; Tubastraea Lesson, 1829; and Dendrophyllia de Blainville, 1830 in having new corallites budding from the common basal coenosteum of colony or from the edge zone of corallites, it differs by: (i) always being attached (ii) having normally arranged septa, (iii) poorly developed, and (iv) porosity of corallum consistently uniform (Capel et al. 2020).

Atlantia denticulata Filander & Kitahara, sp. nov.

Fig. 7G, H

Type locality

Off Gouritsmond, Southern margin of South Africa (FV ‘Compass Challenger’, 35°31'52.31"S, 22°07'25.28"E); 170 m.

Material

Holotype : SAMC_A090157, Southern margin, 132 km from Gouritsmond/143 km off Goukou Estuary, 35°31'52.31"S, 22°07'25.28"E; 170 m. Paratypes: DEFF/SAEON_CCH009 (15 specimens): Southern margin, 132 km from Gouritsmond/133 km off Gourits Estuary, 35°31'52.31"S, 22°07'25.28"E; 170 m. DEFF/SAEON_A32786 (3 specimens): Southern margin, 145 km from Gouritsmond/152 km off Goukou Estuary, 35°39'19.79"S, 22°02'10.68"E; 175 m. UCTES_SST91P (4 specimens): Southern margin, other locality data unknown.

Etymology

The species name denticulata (derived from the Latin dens for small tooth + suffix –ātus) refers to the axial margins of S3 being dentate.

Description

Corallum phaceloid to dendroid. Budding extra-tentacular from base or theca of parent corallite. Holotype consists of 84 corallites, and is 84.6 mm in H. Calice circular to elliptical (GCD:LCD = 1.0–1.1). Epitheca absent. Theca porous, especially near calicular margin. Costae thick, equal in width, granular, and extend to base. Intercostal striae deep. Corallum white.

Septa hexamerally arranged in a normal fashion, with four complete cycles according to the formula: S1 > S2 > S3 > S4 (48 septa). S1 almost meet opposite septa with vertical axial margins. S2 1/3 to ½ width of S1, also with vertical axial margins. S3 1/3 the width of S2 and have slightly laciniate to dentate axial margins. S4 rudimentary, bearing dentate to laciniate axial margins. Septa not exsert. Septal faces bear granules sparsely arranged. Fossa deep, containing a rudimentary columella.

Distribution

Regional: Southern margin, off Gouritsmond; 170 m. Elsewhere: Only known from the type locality.

Remarks

The genus Atlantia was recently described by Capel et al. (2020), who outlined the significance of the normally arranged septa (i.e., Pourtalès Plan absent), poorly developed columella, and uniform corallum porosity in distinguishing the genus from other dendrophylliids. The colonies examined herein represents a new species in the genus. Atlantia denticulata sp. nov. differs from the type species (A. caboverdiana) in its primary septa extending further into columella (almost meeting with other primaries), tertiary septa never fusing to neighbouring primaries, the rudimentary nature of the columella, and the axial margins of S3 consistently being dentate to laciniate.

Balanophyllia Wood, 1844

Diagnosis

Corallum solitary, turbinate to trochoid, fixed or free. Costae usually well developed. Synapticulotheca especially well developed near calice. Septa arranged in Pourtalès plan. Pali may or may not be present. Columella spongy.

Balanophyllia (Balanophyllia) Wood, 1844

Diagnosis

Having a conical corallum firmly attached through a polycyclic base.

Type species

Balanophyllia calyculus Wood, 1844, by monotypy.

Balanophyllia (Balanophyllia) bonaespei van der Horst, 1938

Fig. 7I, J

Balanophyllia bonaespei van der Horst, 1938: 142–145, pl. 5, figs 2–5.

Type locality

Oudekraal, Cape Peninsula, South Africa, depth unknown (van der Horst 1938).

Type material

Unknown.

Material examined

USNM 1423303 (neotype: 1 specimen): Southern margin, 11 km off Agulhas/9 km off De Mond-Heuningnes Estuary, 34°47'12.12"S, 20°08'35.87"E; 32 m.

Description

Corallum trochoid and fixed to substrate by a thin encrusting base. Calice circular to elliptical (GCD:LCD = 1.2). Only specimen examined (USNM 1423303) 9.0 × 7.8 mm in CD and 9.3 mm in H. Epitheca thin from middle corallum to encrusting base. Epitheca bear transverse ridges. Costae conspicuous near calicular margin, being equal in width, granular, and porous. Intercostal striae deep and as porous as costae. Corallum white.

Septa hexamerally arranged in a prominent Pourtalès plan, with five cycles, the last cycle being incomplete, according to the formula: S1 > S2 ≥ S4 > S3 = S5 (≤ 55 septa). S1–2 independent and not reaching columella. S1 most exsert, and have straight to dentate axial margins. S2 slightly less exsert, and 1/3 smaller than S1, also with straight to dentate axial margins. S3–4 progressively less exsert. S3 1/3 the width of S2, and displaying the most dentate axial margins. S4 dimorphic in size: in half-systems without S5, S4 are as wide as S2. Whilst in half-systems with S5, S4 neighbouring S1 as wide as S3 and S4 neighbouring S2 as wide to slightly less wide than S2. Furthermore, half-systems without S5, S4 fuse before S2 and extend to columella as one septum, but in complete half-systems, S5 joins in front of S2 and extend to columella as one septum. S5, if present, similar in size as S3. Fossa of moderate depth, containing a slender and elongate spongy columella aligned with GCD.

Distribution

Regional: Western (van der Horst 1938) and Southern margin of South Africa, extending from off Oudekraal (van der Horst 1938) towards the Agulhas; 32 m. Elsewhere: Only known from South Africa.

Remarks

Only one specimen of Balanophyllia (B.) bonaespei has been collected subsequent to its original description. Since the type is untraceable, we therefore assign this examined specimen as a neotype here. The examined specimen agrees with van der Horst’s (1938) description and illustration in that S5 (when present in half-system) are arranged in a Pourtalès plan, fusing with S3 in front of S4, and then merging with each other before meandering towards S2. This new record further extends the distribution of B. bonaespei to the Southern margin of South Africa. Balanophyllia bonaespei closely resembles B. capensis, and distinguishing features are outlined in the remarks section of that species.

Balanophyllia (Balanophyllia) capensis Verrill, 1865

Fig. 7K, N

Balanophyllia capensis Verrill, 1865: 149. –van der Horst 1938: 140–142. pl. 6, figs 1–6. –Boshoff 1981: 40. –Cairns 1999b: 25.

Balanophyllia bonaespeii. –Boshoff 1981: 40.

Balanophyllia cummingii. –Boshoff 1981: 41.

Balanophyllia (Balanophyllia) capensis. –Cairns 2001: 16.

Type locality

Off Simonstown, South Africa; 567 m (Verrill 1865; van der Horst 1938).

Type material

The holotype is deposited at the YPM (Gall 2020).

Material examined

DEFF/SEAON_A33997 (2 specimens): Southern margin, 39 km from Cape Padrone/40 km off Boknes Estuary, 34°03'53.52"S, 26°42'11.58"E; 100 m. DEFF_BD17.INV02B (2 specimens): Southern margin, 27 km off Cintsa Mouth/25 km off Bulura Estuary, 33°01'22.58"S, 28°17'18.05"E; 122 m. DEFF_NANSEN–INV 16 (1 specimen): Eastern margin, 16 km from Scottburgh/off Mahlongwana Estuary, 30°18'01.19"S, 30°54'47.40"E; 226 m. ORI_EIa3 (28 specimens), ORI_EIa4 (1 specimen), ORI_EIa5 (1 specimen): Eastern margin, other locality data unknown. SAMC_A073003 (2 specimens): Southern margin, other locality data unknown. SAMC_A073020 (8 specimens): Southern margin, Wavecrest Rocks; depth unknown. SAMC_A073032 (1 specimen): Eastern margin, 9 km from Shaka’s Rock/12 km off Mhlali Estuary, 29°32'06.00"S, 31°19'47.99"E; 50 m. SAMC_A073034 (12 specimens): Eastern margin, 53 km from Shaka’s Rock/46 km off Zinkwasi Estuary, 29°33'29.87"S, 31°46'59.88"E; 180 m. SAMC_A073080 (3 specimens): Eastern margin, 49 km from Mtunzini/42 km off Nyoni Estuary, 29°21'24.12"S, 31°56'12.11"E; 180 m. SAMC_A073111 (1 specimen): Southern margin, off Buffalo Bay; 10 m. SAMC_A073121 (2 specimens): Eastern margin, 19 km from Shaka’s Rock/off Mhlali Estuary, 29°32'53.88"S, 31°25'30.00"E; 65 m. SAMC_A073122 (2 specimens): Eastern margin, 39 km from Port St. Johns/13 km off Mkweni Estuary, 31°30'06.11"S, 29°55'12.00"E; 200 m. SAMC_A073163 (2 specimens): Eastern margin, 18 km from Coffee Bay/16 km off Mdumbi Estuary, 32°01'59.87"S, 29°19'05.88"E; 200–210 m. SAMC_A073174 (8 specimens): Eastern margin, 14 km from Coffee Bay/9 km off Mdumbi Estuary, 31°58'48.00"S, 29°16'48.00"E; 90 m. SAMC_A073188 (1 specimen): Western margin, 31 km from Cape Point/13 km off Krom Estuary, 34°09'24.00"S, 18°16'29.99"E; 75 m. SAMC_A073194 (1 specimen): Eastern margin, 46 km from Gonubie/5 km off Berg River V Estuary, 32°57'11.87"S, 28°02'48.12"E; 30 m. SAMC_A073201 (2 specimens): Eastern margin, 10 km from Shaka’s Rock/12 km off Mhlali Estuary, 29°31'59.88"S, 31°19'59.88"E; 51 m. SAMC_A073224 (1 specimen): Southern margin, Cape Point; 13 m. SAMC_A073227 (5 specimens): Southern margin, 18 km from Pringle Bay/16 km off Buffels Oos Estuary, 34°12'36.00"S, 18°46'54.00"E; 40 m. SAMC_A073229 (2 specimens): Southern margin, False Bay; depth unknown. SAMC_A073230 (2 specimens): Southern margin, False Bay; depth unknown. SAMC_A073231 (12 specimens): Southern margin, False Bay; depth unknown. SAMC_A073234 (1 specimen): Southern margin, False Bay; depth unknown. SAMC_A073244 (2 specimens): Southern margin, other locality data unknown. SAMC_A073246 (1 specimen): Eastern margin, 34 km from Coffee Bay/7 km off Ntlonyane Estuary, 32°15'11.99"S, 28°57'42.00"E; 47 m. SAMC_A090104 (1 specimen): Southern margin, 24 km from Pringle Bay/21 km off Buffels Oos Estuary, 34°09'18.00"S, 18°49'36.00"E; 18 m. SAMC_A090106 (1 specimen): Western margin, off Paternoster; depth unknown. SAMC_A090107 (1 specimen): Southern margin, 15 km off Arniston/21 km off De Mond-Heuningnes Estuary, 34°46'59.99"S, 20°19'00.00"E; 80 m. SAMC_A090108 (3 specimens): Southern margin, False Bay; depth unknown. SAMC_A090109 (1 specimen): Southern margin, 8 km from Port Elizabeth/3 km off Bakens River Estuary, 33°58'05.99"S, 25°38'53.99"E; 9 m. SAMC_A090110 (1 specimen): Southern margin, 11 km off Pringle Bay/8 km off Buffels Oos Estuary, 34°16'29.99"S, 18°49'29.99"E; 14–17 m. SAMC_A090111 (6 specimen): Southern margin, 19 km from Pringle Bay/16 km off Buffels Oos Estuary, 34°12'36.00"S, 18°46'41.99"E; 40 m. SAMC_A090120 (2 specimens): Western margin, 16 km from Cape Town/off Diep Estuary, 33°58'59.99"S, 18°21'00.00"E; 17 m. SAM_H1368 (9 specimens): Southern margin, 29 km from Kenton On Sea/off Boesmans Estuary, 33°53'39.99"S, 26°51'00.00"E; 121 m. SAM_H1371 (1 specimen): Southern margin, 3 km from East London/2 km off Buffalo Estuary, 33°00'48.00"S, 27°55'18.73"E; intertidal. SAM_H1377 (1 specimen): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 93 m. SAM_H1383 (1 specimen): Southern margin, 18 km from Cape Padrone/30 km off Boknes Estuary, 33°49'00.00"S, 26°16'59.99"E; 65 m. SAM_H1402 (2 specimens): Southern margin, 16 km from Cape Padrone/28 km off Boknes Estuary, 33°54'15.00"S, 26°22'59.99"E; depth unknown. SAM_H1423 (1 specimen): Southern margin, off Great Fish Point Lighthouse, 183 m. SAM_H1438 (1 specimen): Southern margin, 8 km from Pringle Bay/6 km off Buffels Oos Estuary, 34°17'55.37"S, 18°49'10.85"E; 33 m. SAM_H1471 (3 specimens): Eastern margin, 27 km from Mtunzini/25 km off Matigulu Estuary, 29°10'36.00"S, 31°51'00.00"E; 115 m. SAM_H1479 (3 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°04'59.99"S, 27°49'29.99"E; 146–283 m. SAM_H1483 (9 specimens): Eastern margin, 27 km from Mtunzini/25 km off Matigulu Estuary, 29°10'36.00"S, 31°51'00.00"E; 115 m. SAM_H1484 (2 specimens): Western margin, 14 km from Cape Town/8 km off Diep Estuary, 33°53'59.99"S, 18°23'59.99"E; 40 m. SAM_H1501 (1 specimen): Eastern margin, 207 km from Coffee Bay/off Mdumbi Estuary, 33°12'00.00"S, 30°49'00.00"E; 73 m. SAM_H3043 (6 specimens): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 93 m. SAM_H3044 (9 specimens): Southern margin, 17 km from Cape Point/7 km off Elsies Estuary, 34°12'36.38"S, 18°27'45.71"E; 31 m. SAM_H3045 (1 specimen): Southern margin, 29 km from Kenton On Sea/off Boesmans Estuary, 33°53'39.99"S, 26°51'00.00"E; 121 m. SAM_H3046 (3 specimens): Southern margin, 6 km from Kenton On Sea/5 km off Boknes Estuary, 33°43'07.59"S, 26°37'37.95"E; 90 m. SAM_H3047 (6 specimens): Southern margin, 2 km from Mosselbaai/10 km off Hartenbos Estuary, 34°10'37.57"S, 22°09'19.14"E; 54 m. SAM_H3048 (1 specimen): Southern margin, 20 km from Cape Point/4 km off Elsies Estuary, 34°11'30.71"S, 18°25'55.55"E; depth unknown. SAM_H3143 (1 specimen): Southern margin, 177 km from Gonubie/18 km off Berg River V Estuary, 33°04'36.00"S, 28°06'35.99"E; 90 m. SAM_H3144 (19 specimens): Southern margin, 47 km from Port Alfred/14 km off Mgwalana Estuary, 33°30'18.00"S, 27°22'05.99"E; 80 m. SAM_H3145 (19 specimens): Southern margin, 15 km from Port Alfred/11 km off Riet Estuary, 33°39'18.00"S, 27°01'36.00"E; 90 m. SAM_H3146 (1 specimen): Southern margin, 46 km from Port Alfred/12 km off Mgwalana Estuary, 33°29'24.00"S, 27°21'11.99"E; 80 m. SAM_H3367 (2 specimens): Southern margin, 15 km from Port Elizabeth/16 km off Bakens River Estuary, 33°49'59.99"S, 25°40'00.00"E; depth unknown. SAM_H3835 (1 specimen): Southern margin, 21 km from East London/18 km off Buffalo Estuary, 33°08'59.99"S, 28°01'59.99"E; 84 m. USNM 91776 (9 specimens): Eastern margin, 26 km from Durban/22 km off Beachwood Mangroves, 29°48'00.00"S, 31°16'00.00"E; 232 m.

Imagery data

BMNH 1939.7.20.479–500 (2 specimens), SS ‘Valdivia’ (1 specimen): South Africa, other locality data unknown. MN_SM179 (1 specimen): Southern margin, 47 km from Port Alfred/14 km off Mgwalana Estuary, 33°30'18.00"S, 27°22'05.99"E; 80 m. YPM 6827 (holotype): Western margin, off SIMONSTOWN; 567 m.

Description

Corallum ceratoid to trochoid, straight to slightly curved, and attached through a robust pedicel (PD:GCD = 0.4–0.7) that expands into a thin encrusting base. Calice circular to elliptical (GCD:LCD = 1.1–1.3), calicular margin finely serrate. Largest specimen examined (SAM_H3146) 16.1 × 13.0 mm in CD, and 31.9 mm in H. Thick epitheca covering most of corallum, with transverse ridges. Costae conspicuous near calicular margin, equal in width, slightly convex, separated by porous and thin intercostal striae, and covered with fine pointed granules. Intercostal striae slightly sinuous. Epitheca slightly more solid than upper theca. Corallum white.

Septa hexamerally arranged in five cycles, the last being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 > S5 or S1 ≥ S2 > S3 > S5 > S4 (≤ 74 septa). S1–2 independent but not reaching columella. S1 most exsert, with straight to dentate axial margins. S2 slightly less exsert, sometimes being slightly less wide, also having straight to dentate axial margins. S3–4 progressively less exsert. S3 half width of S2, with most dentate axial margins. S4 commonly arranged in a Pourtalès plan: in a half-system a pair of S4 fuses before S3, and then fuses before S2, before merging to columella. S4 axial margins laciniate. Sometimes S5 merge before S3, before meeting columella as one septum. In other cases, S5 restricted to calicular margin. Fossa of moderately depth, containing a varied slender and elongate spongy columella.

Distribution

Regional: Western to southern margin of South Africa, extending from off Cape Town towards Mtunzini; 9–232 m. Elsewhere: Only known from South Africa.

Remarks

Balanophyllia (B.) capensis has overlapping distribution and common taxonomic characteristics (such as independent S1–2 that never meet columella, S4 arranged in a Pourtalès plan, and presence of epitheca) with B. bonaespei, but may be distinguished by its larger corallum (B. capensis with GCD = 16.1 mm vs. B. bonaespei GCD = 9.0 mm), and thicker epitheca that does not have transverse ridges.

Balanophyllia (Balanophyllia) diademata van der Horst, 1927

Fig. 7O, P

Balanophyllia diademata van der Horst, 1927: 4–5, pl. 2, figs 8, 9. –Cairns and Keller 1993: 220.

Balanophyllia (Balanophyllia) diademata. –Cairns 2001: 16.

Type locality

Off the north-west of Port Dunford; 165 m (van der Horst 1927).

Type material

Unknown.

Material examined

DEFF_FHolon–INV 1 (1 specimen): Eastern margin, off Sodwana; 120 m. SAMC_A073016 (1 specimen): Eastern margin, 31 km from Richards Bay/46 km Mlalazi Estuary, 29°00'54.00"S, 32°15'35.99"E; 500 m.

Description

Corallum ceratoid, straight to slightly curved, and attached by a robust pedicel (PD:GCD = 0.5) that expands into a thin encrusting base. Calice slightly elliptical (GCD:LCD = 1.4–1.5). Largest examined specimen (SAMC_A073016) 12.7 × 8.5 mm in CD, 6 mm in PD, and 24.8 mm in H. Examined specimens lack epitheca. Costae corresponding to septa size (with C3–5 being narrow), porous, densely covered with granules, and separated by narrow and deep intercostal furrows.

Septa hexamerally arranged in five incomplete cycles according to the formula: S1 > S2 > S3 > S5 > S4 (≤ 82 septa). S1–2 most exsert, independent, and reach columella with straight and laciniate axial margins. Higher cycle septa (S3–5) progressively less exsert. S3 dimorphic in size: those in half-systems with S5 ~ ½ size of S2; but ¼ the size of S2 in half-systems without S5. S3 have slightly dentate to laciniate axial margins. S4 also dimorphic in size: ¼ width of S3 in half-systems with S5; but double the width of S3 in half-systems that lack S5. A pair of S4 merges in front of flanked S3 before joining columella as one septum. S4 axial margins slightly more dentate than S3. S5 (when present) has double the width of S4, fusing towards flanked S4 before extending to columella as one septum. S5 bear the most dentate axial margins. All septal faces covered with granules. Septa appear slightly crowded. Fossa of moderate depth, containing a slender, low-profile, and elongated spongy columella.

Distribution

Regional: Eastern margin of South Africa, off Richards Bay extending towards Sodwana; 120–500 m. Elsewhere: Only known from South Africa.

Remarks

The examined material complements van der Horst’s (1927) description of Balanophyllia (B.) diademata, particularly in the irregularity of the number of septal cycles per half-system. However, the examined specimens lack epitheca, whilst van der Horst’s (1927) figured specimen show epitheca reaching ≤ 2.0 mm from calicular margin.

Balanophyllia (Balanophyllia) diffusa Harrison & Poole, 1909

Fig. 8A, B

Balanophyllia diffusa Harrison & Poole, 1909a: 906, pl. 85, fig. 4A, B. –Gardiner and Waugh 1939: 239–240. –Cairns and Keller 1993: 275, fig. 13 A–D.

Balanophyllia (Balanophyllia) diffusa. –Cairns 2001: 16.

Type locality

Off Mergui Archipelago, Myanmar; 5–37 m (Harrison and Poole 1909).

Type material

Unknown.

Material examined

USNM 91780 (1 specimen): Eastern margin, 27 km south of Ponta Do Ouro/18 km off Kosi Bay Estuary, 27°06'00.00"S, 32°52'58.80"E; 74 m. USNM 91782 (1 specimen): Eastern margin, 26 km from Port St. Johns/off Bulolo Estuary, 29°34'47.99"S, 31°41'59.99"E; 138 m.

Description

Corallum ceratoid to subcylindrical, straight to slightly curved, attached. Calice elliptical (GCD:LCD = 1.1–1.3), with slightly lanceted calicular margin. Largest specimen examined (USNM 91782) 11.9 × 9.2 mm in CD, and 34.0 mm in H. Epitheca absent. Costae well developed throughout corallum, broad, separated by porous and thin intercostal striae, and covered with low-profile granules. Corallum white.

Septa hexamerally arranged in five cycles, last cycle being incomplete, according to the formula: S1 ≥ S2 > S4 > S3; or if S5 present S1–2 > S3 > S5 > S4 (≤ 81 septa). S1–2 most exsert, both independent, sometimes reaching columella with straight axial margins. Higher cycle septa (S3–5) progressively less exsert, if at all. S3 ½ size (USNM 91780), or only slightly smaller than S2 (USNM 91782). S3 axial margins straight and vertical. S4 dimorphic in size: those neighbouring S1 ~ as wide as S2; but those neighbouring S2 being the same size as S3, with dentate axial margins (USNM 91780). In half-systems with S5, S4 ~ ½ the width of S3, whereas in half systems without S5, S4 similar in size to S1. S4 bear straight and slightly dentate axial margins (USNM 91782). In most cases a pair of S4 merge before S3 and extend to columella as one septum but sometimes the S5 is arranged in a Pourtalès Plan. Septal face granulation perpendicularly arranged. Fossa shallow, containing a spongy columella that sometimes is slender and aligned in GCD plane (USNM 91782), or relatively small and restricted to fossa (USNM 91780).

Distribution

Regional: Southern to eastern margin of South Africa, from off Shaka’s Rock extending towards Kosi Bay Estuary (27 km south of Ponta Do Ouro: Mozambique); 74–138 m. Elsewhere: Mozambique (Cairns and Keller 1993); Pemba (Gardiner and Waugh 1939); Tanzania (Cairns and Keller 1993); northern banks of Kenya (Cairns and Keller 1993); Maldives (Gardiner and Waugh 1939); Red Sea (Gardiner and Waugh 1939); 6–274 m.

Remarks

Records of Balanophyllia (B.) diffusa listed herein are those reported by Cairns and Keller (1993), who noted that the quasi-colonial nature of Gardiner and Waugh’s (1939 [page 239])) specimens is a result of specimens settling closer to one another. The largest examined specimen (USNM 91782) has closely packed septa, and a relatively complex and elongated columella, as compared with the other specimen (USNM 91782), which exhibits loosely packed septa and a fairly simple and short columella.

Figure 8. 

A, B Balanophyllia (Balanophyllia) diffusa A (USNM 91782, off Port St. Johns, 138 m) calicular view B (USNM 91780, off Kosi Bay Estuary, 74 m) calicular view C, D Balanophyllia (Balanophyllia) sp. cf. malouinensis C (SAM_H3069, off East London, 146–238 m) calicular view D (SAM_H3068, off Mazeppa Bay, 174 m) lateral view E, F Balanophyllia (Eupsammia) stimpsonii (SAM_H3831, off Cape Point, 97–99 m) E calicular view F lateral view G, H Dendrophyllia arbuscula (SAMC_A073119, off Cape Vidal, 65–70 m) G calicular view H lateral view I, J Dendrophyllia cladonia I (SAM_H1445, off Plettenberg Bay, 146 m) calicular view J (SAM_ H2833, off Gonubie, 155 m) lateral view K, L Dendrophyllia cornigera K (USNM 91827, off Durban, 232 m) calicular view L (SAM_ H3841, off Pringle Bay, depth unknown) lateral view M, N Dendrophyllia dilatata (SAMC_A073016, off Richards Bay, 500 m) M calicular view N lateral view O, P Dendrophyllia ijimai O (USNM 91844, off Shaka’s Rock, 68–70 m) calicular view P (SAMC_A090121, off Port Dunford, 85 m) lateral view. Scale bars: 10 mm (AI, K, M, O); 100 mm (J, L, N, P).

Balanophyllia Balanophyllia sp. cf. malouinensis Squires, 1961

Fig. 8C, D

Balanophyllia malouinensisSquires, 1961: 15, 39–40, 46, fig. 5. –Squires 1969: 17–18. pl. 6, map 2. –Cairns 1979: 206. –Cairns 1982: 52–53, pl. 16, figs 4–7, pl. 17, figs 1–3, pl. 18, fig. 7. –Cairns 2005: 43.

Balanophyllia (Balanophyllia) malouinensis. –Cairns 2001: 17.

Type locality

South of East Falkland Island, Atlantic archipelago (52°32'00"S, 61°15'00"W); 358 m (Cairns 1982).

Type material

The holotype is deposited at the AMNH (Cairns 1982).

Material examined

DEFF_BD17–INV02.B (2 specimens): Southern margin, 27 km from Cintsa Mouth/25 km off Bulura Estuary, 33°01'22.58"S, 28°17'18.05"E; 122 m. SAMC_A073031 (3 specimens): Eastern margin, 12 km from Port St. Johns/11 km off Bulolo Estuary, 31°44'17.87"S, 29°32'42.00"E; 300 m. SAMC_A073085 (1 specimen): Eastern margin, 7 km south of Ponta Do Ouro/8 km off Kosi-Kumpungwini (Sifungwe) Estuary, 26°54'40.32"S, 32°55'05.88"E; 47 m. SAMC_A073183 (3 specimens): Eastern margin, 27 km from Richards Bay/40 km off Mlalazi Estuary, 29°00'24.11"S, 32°12'00.00"E; 152 m. SAMC_A073184 (2 specimens): Eastern margin, 39 km from Scottburgh/36 km off Fafa Estuary, 30°34'00.00"S, 31°00'00.00"E; 900–625 m. SAMC_A073186 (1 specimen): Eastern margin, 20 km from Durban/13 km off Mbokodweni Estuary, 30°01'05.87"S, 31°03'11.88"E; 150 m. SAM_H833 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3065 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3066 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3067 (15 specimens): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3068 (3 specimens): Southern margin, 14 km from Mazeppa Bay/20 km off Great Kei Estuary, 32°34'00.00"S, 28°33'00.00"E; 174 m. SAM_H3069 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3070 (1 specimen): Southern margin, 25 km from Gonubie/24 km off Gqunube Estuary, 33°06'17.99"S, 28°10'59.99"E; 155 m.

Description

Corallum ceratoid to sub-cylindrical, straight to slightly curved, attached to substrate by a robust pedicel (PD:GCD = 0.30–0.6) that expands into a thin encrusting base. Calice circular to elliptical (GCD:LCD = 1.0–1.1), with serrate calicular margin. Largest specimen examined (SAM_H3069) 14.5 × 13.4 mm in CD, 4.0 mm in PD, and 24.0 mm in H. Synapticulotheca well developed and porous, visible at calicular margin. Costae poorly developed, separated by porous and thin intercostal striae, and covered with fine pointed granules. Epitheca thin, irregularly banded. Corallum white.

Septa hexamerally arranged in five cycles, last cycle being incomplete, according to the formula: S1–2 > S4 > S3 > S5 (≤ 52 septa). S1–2 most exsert, independent, and reach columella deeper in fossa with straight axial margins. Higher cycle septa (S3–4) progressively less exsert. S3 1/4 the width of S2, with straight to slightly sinuous axial margins. S4 as wide as S2 and arranged in a Pourtalès plan: in each half-system a pair of S4 fuses before S3 and extend to columella as one septum with laciniate axial margins. S5, if present, sometimes restricted to calicular margin, or arranged in Pourtalès plan before common S4. Septal faces bear granules sparsely arranged. Fossa of moderate depth, containing a thick and elongate spongy columella.

Distribution

Regional: Southern to eastern margin of South Africa, extending from off East London extending towards Kosi-Kumpungwini (Sifungwe) Estuary (7 km south of Ponta Do Ouro: Mozambique); 47–900 m. Elsewhere: Sub-Antarctic and Antarctica (Cairns 1982); Chile (Cairns et al. 2005); 75–1137 m.

Remarks

Although the examined specimens closely resemble Balanophyllia (B.) malouinensis in their: (i) spongy columella, (ii) synapticulotheca being restricted to the calicular margin, (iii) thin epitheca, and (iv) poorly developed costae, variation in the septal development exists. The examined material consistently has exsert septa (particularly the primaries) unlike those examined by Cairns (1982) which are not exsert. Furthermore, the examined specimens have the septal formula S1–2 > S4 > S3 > S5, whereas that of B. malouinensis is S1–2 > S5 > S3 > S4. Nonetheless, dendrophylliids are known to exhibit plasticity, making it difficult to evaluate intra-specific variations.

Balanophyllia (Balanophyllia) vanderhorsti Cairns, 2001

Balanophyllia ponderosa van der Horst, 1926: 49–50, pl. 3, figs 6, 7. –Cairns and Keller 1993: 274. –Cairns 1994: 83, pl. 3A, B.

Balanophyllia (Balanophyllia) vanderhorsti Cairns, 2001: 16 (replacement name for junior homonym). –Tachikawa 2005: 11, Pl. 4, figs I, J.

Type locality

Off the Maldives; 53 m (van der Horst 1926).

Type material

The holotype is deposited at the NHMUK (Cairns and Keller 1993).

Material examined

None.

Distribution

Regional: Eastern margin of South Africa, off Richards Bay; 51 m. Elsewhere: Seychelles (van der Horst 1926); Maldives (van der Horst 1926); off Sri Lanka (Alcock 1893); Japan (Eguchi 1968; Cairns 1994; Tachikawa 2005); 14–59 m.

Remarks

No new records of Balanophyllia (B.) vanderhorsti were examined during the present study. Therefore, the occurrence of this species in South Africa is based solely on previous published records. Boshoff’s (1981) record of B. vanderhorsti (reported as B. ponderosa (which is a junior homonym)) has basal stolons and therefore represents Rhizopsammia Verrill, 1870. To reiterate, Cairns and Keller (1993) observed that this South African species closely resembles B. regalis (Alcock, 1893) in lacking an epitheca.

Balanophyllia (Eupsammia) Milne-Edwards & Haime, 1848

Diagnosis

Corallum solitary, conical, and usually free, having a monocyclic base; corallum often curved or bent. Asexual budding may occur from margin zone. Epitheca present or absent (present in type species); synapticulotheca always costate. Costae bear short, hispid spines. Pourtalès plan present; fifth cycle of septa often present. Columella elongate, spongy. Endotheca absent.

Type species

Madrepora trochiformis Pallas, 1766, by subsequent designation (Milne-Edwards and Haime 1850b).

Balanophyllia (Eupsammia) stimpsonii (Verrill, 1865)

Fig. 8E, F

Eupsammia stimpsonii Verrill, 1865: 150.

Eupsammia stimpsoniana.– Verrill 1866: 29, pl. 2, figs 3, 3a.

Rhodopsammia socialis Semper, 1872: 260–261, pl. 20, figs 1–4. –Faustino 1927: 229, pl. 75, figs 9–12.– Alcock 1893: 147.

Rhodopsammia affinis Semper, 1872: 261–262, pl. 19, fig. 7A, B.

Rhodopsammia incerta Semper, 1872: 264, pl. 19, fig. 8A, B.– Faustino 1927: 231, pl. 75, figs 3, 4.

Leptopsammia conica van der Horst, 1922: 68–69, pl. 8, figs 14, 15.

Balanophyllia affinis . –Faustino 1927: 228–232, pl. 75, figs 1, 2. –van der Horst 1922: 62. –van der Horst 1931:10. –Gardiner and Waugh 1939: 240.

Balanophyllia stimpsonii . –Zibrowius 1985: 234–235, figs 1–14. –Zibrowius and Grygier 1985: 126–127. –Cairns and Keller 1993: 274. –Cairns and Zibrowius 1997: 176–177. –Cairns 2004a: 313.

Balanophyllia (Eupsammia) stimpsoni. –Cairns 2001: 19. –Kitahara and Cairns 2021: 289–290, figs 142, 152H–J.

Type locality

North China Sea; depth unknown (Verrill 1865).

Type material

Two syntypes are deposited at the YPM (Zibrowius 1985; Cairns 2004a).

Material examined

SAMC_A073157 (2 specimens): Eastern margin, 10 km from Port Edward/24 km off Bilanhlolo Estuary, 31°05'48.11"S, 30°18'47.88"E; 140 m. SAM_H3196 (1 specimen): Eastern margin, 9 km from Shaka’s Rock/2 km off Tongati Estuary, 29°34'00.00"S, 31°10'59.99"E; 66 m. SAM_H3831 (1 specimen): Southern margin, 16 km from Cape Point/18 km off Buffels Oos Estuary, 34°23'17.99"S, 18°39'24.00"E; 97–99 m.

Description

Corallum ceratoid, slightly curved, unattached. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1), with serrate calicular margin. Largest specimen examined (SAM_H3831) 10.2 × 9.4 mm in CD, and 21.5 mm in H. Theca well developed. Upper theca porous, and lower theca more solid (epitheca). Costae poorly developed near calicular margin, becoming distinctively visible towards base, and covered with fine pointed granules. Epitheca irregularly banded. Intercostal striae thin and porous. Corallum creamy.

Septa hexamerally arranged in four complete cycles according to the formula: S1 > S2 > S4 > S3 (48 septa). S1–2 most exsert, with straight axial margins. S1 independent and extend towards columella. S2 slightly less wide than S1. Higher cycle septa (S3–4) progressively less exsert, and bear laciniate axial margins. S3 1/3 the width of S2. In complete half-systems a pair of S4 fuses before S3 and extend to columella as one septum. All septa eventually join columella deep in fossa. Septal faces granular, with granules arranged perpendicular to septal margin. Fossa of moderate depth, containing a low, slender, and elongated spongy columella.

Distribution

Regional: Southern to eastern margins of South Africa, off Cape Point extending towards Shaka’s Rock; 66–99 m. Elsewhere: Philippines; Indonesia (Cairns and Zibrowius 1997); Australia (Cairns 2004a); New Caledonia (Kitahara and Cairns 2021); Chesterfield Islands (Cairns and Zibrowius 1997); Somalia; Mozambique (Zibrowius and Grygier 1985); Seychelles; Reunion (Zibrowius 1985); Gulf of Oman (Zibrowius and Grygier 1985); Sri Lanka (van der Horst 1926; Gardiner and Waugh 1939); Gulf of Manaar (Bourne 1905); Maldives (Gardiner and Waugh 1939); Andaman Sea (Alcock 1893); Mergui Archipelago; Myanmar (Harisson and Poole 1909); 18–95 m.

Remarks

Balanophyllia (E.) stimpsonii resembles B. carinata (Semper, 1872) amongst the other four Recent unattached congeners (B. caribbeana Cairns, 1977a, B. imperialis Kent, 1871, B. pittieri Vaughan, 1919, and B. regalis (Alcock, 1983) in its septal arrangement in four cycles (in contrast to five incomplete ones). As stated by Cairns and Zibrowius (1997), Balanophyllia (E.) stimpsonii differs from B. carinata in having a less compressed corallum, lack of keeled thecal edges, and S1 > S2. Balanophyllia (E.) stimpsonii is also known to be a host for an ascothoracidan barnacle species (Zibrowius and Grygier 1985). Balanophyllia (E.) stimpsonii is common shallow water Indo-Pacific species, which was previously reported off False Bay and Natal (Zibrowius 1985).

Dendrophyllia Blainville, 1830

Diagnosis

Colonies formed by extra-tentacular budding, resulting in three general forms: arborescent colonies with axial corallites; small bushy colonies with sparse branching; or dendroid colonies with sympodial branching. All forms originate from a single basal stem. Synapticulothecate. Costae usually clearly defined. Septa arranged in Pourtalès plan. Pali may be present. Columella spongy. Tabular endothecal dissepiments may be present.

Type species

Madrepora ramea Linnaeus, 1758, by subsequent designation (Milne-Edwards and Haime 1850b).

Dendrophyllia arbuscula van der Horst, 1922

Fig. 8G, H

Dendrophyllia arbuscula van der Horst, 1922: 53, pl. 8, fig. 6 (in part: ‘Siboga’ Stn. 277, pl. 8, fig. 6). –Yabe and Eguchi 1942b: 162, 166–167. –Crossland 1952: 92. –Eguchi 1968: C55–C56, pl. C21, figs 5, 13. –Wells 1964: 108. –Pillai and Scheer 1974: 462, fig. 7A. –Pillai 1983: 89. –Veron 1986: 578. –Cairns 1994: 90–91, pl. 38, figs I–L. –Cairns 1995: 125–126, pl. 43, figs E, F. –Ogawa and Takahashi 1995: 17, pl. 1, figs 1–7. –Cairns and Zibrowius 1997: 192–193, fig. 29A–C. –Cairns 1998: 408–409. –Cairns 1999a: 133–134. –Cairns et al. 1999: 26. –Cairns 2001: 34. –Cairns 2004a: 267, 315. –Lam et al. 2009, 732–733, fig. 1E–I.

Dendrophyllia micranthus. –van der Horst 1922: 50.

Dendrophyllia subcornigera Eguchi, 1968: C64, pl. C32, figs 3, 4.

Dendrophyllia horsti Gardiner & Waugh, 1939: 237–238, pl. 2, figs 5, 6. –Fricke and Schuhmacher 1983: 184, fig. 14D.

Dendrophyllia sp. cf. D. horsti. –Cairns and Keller 1993: 278, pl. 13, figs F, I.

Type locality

Off Banda Sea, Indonesia (HMS ‘Siboga’ stns. 260 and 277); 45–90 m (van der Horst 1922).

Type material

Three syntypes are deposited at the ZMA (Cairns 1994).

Material examined

SAMC_A073119 (1 colony): Eastern margin, 39 km from Cape Vidal/29 km off Mgobezeleni Estuary, 27°47'23.99"S, 32°38'53.87"E; 65–70 m. SAM_H3064 (1 colony): Southern margin, 18 km from Cape Padrone/30 km off Boknes Estuary, 33°49'00.00"S, 26°16'59.99"E; 65 m. SAM_H5104 (1 colony): Eastern margin, 59 km from Cape Vidal/9 km off Mgobezeleni Estuary, 27°36'38.45"S, 32°40'02.99"E; 59 m. USNM 91815 (2 colonies): Eastern margin, 90 km from Shaka’s Rock/9 km off Boesmans Estuary, 29°32'12.11"S, 31°19'47.99"E; 50 m. USNM 91816 (3 colonies): Eastern margin, 17 km south of Ponta Do Ouro/10 km off Kosi-Kumpungwini (Sifungwe) Estuary, 27°00'11.87"S, 32°54'18.00"E; 68 m. USNM 91817 (1 colony): Eastern margin, 17 km south of Ponta Do Ouro/11 km off Kosi-Kumpungwini (Sifungwe) Estuary, 27°00'24.11"S, 32°55'12.00"E; 66 m. USNM 91818 (2 colonies): Eastern margin, 19 km south of Ponta Do Ouro/12 km off Kosi-Kumpungwini (Sifungwe) Estuary, 27°01'05.87"S, 32°55'05.88"E; 69–73 m. USNM 91819 (1 colony): Eastern margin, 29 km south of Ponta Do Ouro/19 km Kosi Bay Estuary, 27°06'47.87"S, 32°52'54.12"E; 74 m. USNM 91820 (1 colony): Eastern margin, 41 km south of Ponta Do Ouro/26 km off Kosi Bay Estuary, 27°13'00.11"S, 32°49'41.87"E; 72 m.

Description

Colony small, attached, and bear few corallites, of which one is axial; remaining budding from lower or upper corallum of axial corallite. Primary corallite elongate cylindrical, and firmly attached to substrate through a robust pedicel (PD:GCD = 0.9). Secondary corallites low (< 5 mm in H). Calice slightly elliptical (GCD:LCD = 1.1–1.4), calicular margin lanceted. Largest corallite examined (axial) 12.0 × 10.7 mm in CD, and 31.2 mm in H. Costae conspicuous, broad, equal in width, flat, slightly porous, and granular. Intercostal furrows shallow and quite porous. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1 ≥ S2 > S3 > S4 or S1 ≥ S2 > S4 > S3 (48 septa), with S5 sometimes present. At calicular margin, upper outer margin of S4 fuses to neighbouring septa (S1 or S2) forming small triangular lancets. S1 independent, slightly exsert, and extend towards columella with straight axial margins. S2 also independent, slightly less exsert, as wide to only slightly smaller than S1, and also bearing straight axial margins. Higher cycle septa (S3–4) becoming progressively less exsert, if at all. S3 dimorphic in size, being 1/3 the width of S2 or sometimes rudimentary, with dentate to laciniate axial margins. S4 arranged in a Pourtalès plan: a pair of S4 curves towards common S3 fusing before it deep in fossa, may be a 1/3 wider than S3, axial margins dentate to laciniate. Septal faces finely granular. Fossa deep, containing a massive columella usually swirled in a clockwise direction.

Distribution

Regional: Southern and Eastern margin of South Africa, off Cape Padrone extending towards Kosi-Kumpungwini (Sifungwe) Estuary (17 km south of Ponta Do Ouro: Mozambique); 50–73 m. Elsewhere: Japan, China Sea (Cairns 1994); Philippines, Indonesia (Cairns and Zibrowius 1997); Vanuatu (Cairns 1999a); New Caledonia (Kitahara and Cairns 2021); Australia (Cairns 1998); New Zealand (Cairns 1995); Red Sea (Fricke and Schuhmacher 1983); Maldives; Pemba; Tanzania (Gardiner and Waugh 1939; Cairns and Keller 1993); 2–386 m.

Remarks

Dendrophyllia arbuscula is historically known from the eastern margin of South Africa. One of the examined specimens (SAM_H3064) extends the regional distribution south towards Cape Padrone. Although Gardiner and Waugh (1939) noted the resemblance of D. horsti to D. arbuscula, an observation also made by Cairns and Keller (1993), it was only four years later that D. horsti was listed as a junior synonym of D. arbuscula (Cairns & Zibrowius, 1997). Nonetheless, based on growth form, D. arbuscula is within Cairns’(1994) second Dendrophyllia morphological group. Two other South African congers (D. cladonia van der Horst, 1927 and D. cornigera (Lamarck, 1816)) are known to form small and bushy colonies that originate from an axial corallite (with relatively few additional corallites to the primary), however, dissimilarities will be discussed in these species’ accounts.

Dendrophyllia cladonia van der Horst, 1927

Fig. 8I, J.

Dendrophyllia cladonia van der Horst, 1927: 3–4, pl. 1, figs 5, 6, pl. 2, fig. 7. –Zibrowius and Gili 1990: 44. –Cairns and Keller 1993: 279. –Cairns 2001: 34.

Type locality

Off Port Shepstone (?), South Africa (RV ‘Pieter Faure’ at ca. 30°44'15.3"S, 30°27'35.0"E); 457 m (van der Horst 1927).

Type material

Type specimen is possibly deposited at the NHMUK (GBIF 2020).

Material examine

DEFF_BD13–INV 03 (6 fragments): Southern margin, 25 km from Cintsa Mouth/off Bulura Estuary, 32°58'59.08"S, 28°19'10.82"E; 104 m. DEFF/SAEON_D00491 (1 fragment): Southern margin, 138 km from Agulhas/145 km off De Mond-Heuningnes Estuary, 35°54'16.19"S, 20°45'46.19"E; 135 m. DEFF/SAEON_D00584 (1 fragment): Southern margin, 56 km from Knysna/57 km off Goukamma Estuary, 34°34'32.40"S, 23°06'04.20"E; 111 m. DEFF/SEAON_D00829 (1 fragment): Southern margin, 153 from Agulhas/157 km off De Mond-Heuningnes Estuary, 35°51'57.60"S, 21°07'14.87"E; 122 m. DEFF/SAEON_D00832 (1 fragment): Southern margin, 171 km from Stilbaai/172 km off Goukou Estuary, 35°54'51.47"S, 21°42'48.59"E; 165 m. DEFF/SAEON_D00851 (1 fragment): Southern margin, 50 km from Agulhas/55 km off De Mond-Heuningnes Estuary, 35°08'14.27"S, 20°24'55.08"E; 113 m. SAMC_A073012 (1 fragment): Eastern margin, 10 km from Port Edward/27 km off Bilanhlolo Estuary, 31°06'46.79"S, 30°17'48.12"E; 120–125 m. SAMC_A073015 (2 fragments): Southern margin, 32 km from Mazeppa Bay/19 km off Mendu Estuary, 32°25'00.11"S, 28°58'18.11"E; 330–340 m. SAMC_A073028 (4 fragments): Eastern margin, 26 km from Cape Vidal/25 km off St Lucia Estuary, 27°54'18.00"S, 32°37'59.87"E; 105 m. SAMC_A073042 (1 fragments): Eastern margin, 53 km from Shaka’s Rock/46 km off Zinkwasi Estuary, 29°32'53.88"S, 31°47'12.11"E; 200 m. SAMC_A073046 (10 fragments): Eastern margin,9 km from Port Edward/9 km off Blinde Estuary, 31°05'23.99"S, 30°18'00.00"E; 125 m. SAMC_A073057 (1 fragment): Eastern margin, 6 km from Cape Vidal/17 km off St Lucia Estuary, 28°08'17.88"S, 32°36'54.00"E; 200 m. SAMC_A073076 (1 fragment): Locality unknown. SAMC_A073165 (1 fragment): Eastern margin, 28 km south of Ponta Do Ouro/17 km off Kosi Bay Estuary, 27°06'18.00"S, 32°52'00.12"E; 50 m. SAMC_A073212 (4 fragment): Eastern margin, 5 km from Cape Vidal/16 km off St Lucia Estuary, 28°07'05.88"S, 32°36'35.99"E; 145 m. SAM_H1362 (1 fragment): Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 90 m. SAM_H1412 (6 fragments): Eastern margin, 2 km from Port Shepstone/Mzimkulu Estuary, 30°44'14.23"S, 30°27'34.72"E; depth unknown. SAM_H1445 (1 fragment): Southern margin, 4 km from Plettenberg Bay/7 km off Piesang Estuary, 34°06'00.00"S, 23°23'59.99"E; 146 m. SAM_H1447 (1 fragment): Eastern margin, 38 km from St. Lucia/31 km off Nhlabane Estuary, 28°40'59.99"S, 32°34'00.00"E; 73 m. SAM_H1513 (1 fragment): Southern margin, 68 km from Kidds Beach/off Gouda Estuary, 33°36'00.00"S, 28°10'59.99"E; 174 m. SAM_H2833 (1 fragment): Southern margin, 25 km from Gonubie/24 km off Gqunube Estuary, 33°06'17.99"S, 28°10'59.99"E; 155 m. SAM_H3042 (1 fragment) Southern margin, 28 km from Port Alfred/3 km off Old Woman’s Estuary, 33°30'00.00"S, 27°08'59.99"E; 93 m. SAM_H3058 (1 fragment): Southern margin,9 km from Pringle Bay/7 km off Buffels Oos Estuary, 34°17'30.00"S, 18°48'00.00"E; 33 m. SAM_H3064 (1 fragment): Southern margin, 18 km from Cape Padrone/30 km off Boknes Estuary, 33°49'00.00"S, 26°16'59.99"E; 65 m. SAM_H3413 (1 fragment): Southern margin, 18 km from Gansbaai/38 km off Ratels Estuary, 34°44'12.00"S, 19°25'36.00"E; 70–50 m. SAM_H3838 (1 fragment): Eastern margin, 7 km from Port Shepstone/off Boboyi Estuary, 30°48'00.00"S, 30°29'05.99"E; depth unknown. USNM 91823 (48 fragments): Eastern margin, 26 km from Port St. Johns/off Bulolo Estuary, 29°34'47.99"S, 31°41'59.99"E; 138 m. USNM 91825 (7 fragments): Eastern margin, 35 km from Port Dunford/38 km off Mlalazi Estuary, 29°10'00.00"S, 32°04'59.99"E; 168 m.

Description

Colony with axial corallite reaching ≤ 93.5 mm. Primary corallite elongate, cylindrical, and firmly attached to substrate through a robust pedicel (PD:GCD = 1.2–1.5). Secondary corallites ceratoid/trochoid to cylindrical. Calice slightly circular to elliptical (GCD:LCD = 1.0–1.3); calicular margin lanceted. Largest corallite examined (axial) 11.1 × 10.7 mm in CD, 17.0 mm in PD, and 93.5 mm in H. Costae conspicuous throughout corallum, broad, equal in width, flat, slightly porous, and granular. Intercostal furrows shallow and quite porous. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1 ≥ S2 > S4 > S3 (48 septa). S1 independent and reach columella with vertical and straight axial margins. S2 also independent, and as wide to only slightly smaller than S1. S2 axial margins also vertical and straight. S3 smallest septa, and bear dentate to laciniate axial margins. S4 arranged in a Pourtalès plan: in each system the S4 neighbouring S1 fuses before S2, before meeting columella. S4 also have dentate to laciniate axial margins. Septal faces finely granular. Fossa deep, containing a massive spongy columella.

Distribution

Regional: Southern and Eastern margin of South Africa, off Gansbaai extending towards Kosi Bay Estuary (276 km south of Ponta Do Ouro: Mozambique); 33–340 m. Elsewhere: Mozambique (Cairns and Keller 1993); 49–457 m.

Remarks

As in Dendrophyllia arbuscula, D. cladonia belongs to the second Dendrophyllia group (Cairns 2001), which form small and bushy colonies originating from an axial corallite and with relatively few additional corallites to the primary. Dendrophyllia cladonia may be distinguished by its Pourtalès plan arrangement, whereby S4 adjacent to S1 are wider than S4 neighbouring S2, and merge towards S2 before joining the columella (Cairns and Keller 1993), and by its smaller axial corallite GCD (< 9 mm). Under-developed D. cladonia may be confused with Balanophyllia capensis but can be differentiated by having S4 > S3 (S3 > S4 in B. capensis). Apart from the septal arrangement, D. cladonia can further be distinguished by its S4 merging towards S2, and have a more robust pedicel. Although the calicular margin of D. cladonia is rarely intact, making it difficult to evaluate septa exsertness, the arrangement of septa assists in distinguishing species from both D. arbuscula and B. capensis (as highlighted above).

Dendrophyllia cornigera (Lamarck, 1816)

Fig. 8K, L

Madrepora ramea. –Linnaeus 1758: 797.

Caryophyllia cornigera Lamarck, 1816: 228.

Dendrophyllia cornigera. –Milne-Edwards and Haime 1848c: 100. –Lacaze-Duthiers 1879: 216, pl. 11, fig. 8. –von Marenzeller 1904a: 313, pl. 18, fig. 21. –Moseley 1881: 198. –Gravier 1920: 104, pl 12, figs 186–192. –Joubin 1922: 8, figs 4, 5. –van der Horst 1926: 44. –van der Horst 1928: 1, pl. 1, fig. 1. –Best 1970: 315, fig. 15. –Zibrowius et al. 1975: 97, pl. 5G. –Zibrowius 1979: 19, pl. 1, figs 1, 2. –Zibrowius 1980: 172–175, pl. 87, fig. 1A. –Blainville 1834: 354. –Ocaña et al. 2015: 44–43. –Cairns 2001: 33.

Type locality

Unknown, presumed to be off the Mediterranean (Zibrowius 1980).

Type material

Type specimen is presumably lost (Zibrowius 1980).

Material examined

DEFF/SAEON_D00817 (1 specimen): Southern margin, 23 km from Plettenberg Bay/25 km off Piesang Estuary, 34°16'15.60"S, 23°24'50.40"E; 95 m. SAMC_A072969 (1 specimen): Locality data unknown; 125 m. SAMC_A072982 (2 specimens): Southern margin, 14 km from Pringle Bay/16 km off Buffels Oos Estuary, 34°23'48.00"S, 18°41'05.99"E; 88 m. SAMC_A073003 (9 specimens): Locality data unknown. SAMC_A073010 (2 specimens): Eastern margin, 20 km from Durban/13 km off Mbokodweni Estuary, 30°01'05.87"S, 31°03'11.88"E; 150 m. SAMC_A073023 (3 specimens): Locality data unknown. SAMC_A073031 (3 specimens): Eastern margin, 12 km from Port St. Johns/11 km off Bulolo Estuary, 31°44'17.87"S, 29°32'42.00"E; 300 m. SAMC_A073087 (3 specimens): Eastern margin, 32 km from Port Dunford/33 km off Mlalazi Estuary, 29°11'24.00"S, 31°59'23.99"E; 50 m. SAMC_A073148 (3 specimens): Eastern margin, 26 km from Cape Vidal/25 km off St Lucia Estuary, 27°54'18.00"S, 32°37'59.87"E; 105 m. SAMC_A073174 (1 specimen): Eastern margin, 132 km from Coffee Bay/13 km off Bitou Rivier Estuary, 31°58'48.00"S, 29°16'48.00"E; 90 m. SAMC_A073272 (1 specimen): Locality data unknown; 101 m. SAMC_A090119 (2 specimens): Eastern margin, 28 km south of Ponta Do Ouro/18 km off Kosi Bay Estuary, 27°06'29.99"S, 32°52'54.00"E; 70 m. SAM_H1361 (2 specimens): Southern margin, off CAPE ST BLAIZE; 212 m. SAM_H3062 (1 specimen): Southern margin, 11 km from East London/5 km off Gouda Estuary, 33°05'03.24"S, 27°49'33.40"E; 146–238 m. SAM_H3063 (1 specimen): Southern margin, 28 km from Gonubie/27 km off Buffalo Estuary, 33°09'29.99"S, 28°03'06.00"E; 86 m. SAM_H3147 (1 specimen): Southern margin, 15 km from Port Alfred/11 km off Riet Estuary, 33°39'18.00"S, 27°01'36.00"E; 90 m. SAM_H3148 (1 specimen): Eastern margin, 32 km from Coffee Bay/9 m off Ntlonyane Estuary, 32°15'00.00"S, 29°00'47.99"E; 90 m. SAM_H3412 (1 specimen): Southern margin, 14 km from Gansbaai/31 km off Bot River Lagoon, 34°36'35.99"S, 19°12'36.00"E; 78 m. SAM_H3839 (1 specimen): Southern margin, 9 km from East London/6 km off Buffalo Estuary, 33°04'00.00"S, 27°56'59.99"E; depth unknown. SAM_H3840 (1 specimen): Southern margin, 15 km from Pringle Bay/14 km off Buffels Oos Estuary, 34°15'06.00"S, 18°44'48.00"E; 53 m. SAM_H3841 (1 specimen): Southern margin, 12 km from Pringle Bay/10 km off Buffels Oos Estuary, 34°17'35.99"S, 18°45'00.00"E; depth unknown. SAM_H3842 (1 specimen): Southern margin, 19 km from Pringle Bay/16 km off Buffels Oos Estuary, 34°12'36.00"S, 18°46'41.99"E; 40 m. SAM_H3843 (1 specimen): Southern margin, 12 km from Pringle Bay/10 km off Buffels Oos Estuary, 34°17'35.99"S, 18°45'00.00"E; depth unknown. SAM_H3844 (1 specimen): Southern margin, 12 km from Pringle Bay/11 km off Buffels Oos Estuary, 34°17'59.99"S, 18°44'30.00"E; 40 m. USNM 91827 (6 specimens): Eastern margin, 26 km from Durban/22 km off Beachwood Mangroves, 29°48'00.00"S, 31°16'00.00"E; 232 m.

Imagery data

BMNH 1939.7.20.8 (2 specimens): Eastern margin, off Umhloti River mouth; depth unknown. BMNH 1939.7.20.317 (2 specimens): Locality data unknown. ZMA 1194 (2 specimens): Southern margin; 95 m.

Description

Colony bushy, stemming from an elongate, cylindrical, and firmly attached axial corallite that has a robust pedicel (PD:GCD = 1.1). Secondary corallites ceratoid/trochoid to cylindrical. Calice circular to slightly elliptical (GCD:LCD = 1.0–1.1); calicular margin lanceted. Largest corallite examined (axial) 9.3 × 8.3 mm in CD, 9.9 mm in PD, and 55.0 mm in H. Costae conspicuous throughout corallum, broad, equal in width, flat, slightly porous, and granular. Intercostal furrows shallow and quite porous. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1 ≥ S2 > S4 > S3 (48 septa). S1 independent, not reaching columella, and bear dentate (occasionally smooth) axial margins. Remaining septa (S2–4) also bear dentate axial margins, of which that of S4 having conspicuously longer teeth. S2 also independent, as wide to only slightly smaller than S1. S3 least wide septa. S4 arranged in a Pourtalès plan: in each system a pair of S4 neighbouring S1 curves towards common S2, fusing with S4 closer to S2 deeper in fossa before reaching columella. Septal faces finely granular. Fossa moderately deep, containing a spongy columella.

Distribution

Regional: Western to eastern margins of South Africa, from off Pringle Bay extending towards off Kosi Bay Estuary (28 km south of Ponta Do Ouro: Mozambique); 40–300 m. Elsewhere: Mediterranean (Duncan 1873; Ocaña et al. 2015); Morocco (Zibrowius 1983); Celtic Sea (Joubin 1922); off France (Lacaze-Duthiers 1879; Moseley 1881); Bay of Biscay (Milne-Edwards and Haime 1848c); Portugal (Zibrowius 1980); Canary Islands (Gravier 1920); Maldives; Almirantes; and Providence (van der Horst 1926); 89–600 m.

Remarks

Dendrophyllia cornigera has been systematically recorded in the southwest Indian Ocean (van der Horst 1926), specifically off the southern Agulhas (von Marenzeller 1904a; van der Horst 1927) and KwaZulu-Natal region (van der Horst 1927; Cairns and Keller 1993). Nonetheless, the examined specimens are in accordance with the description of Mediterranean specimens (Zibrowius 1980). Dendrophyllia cornigera is distinguished from D. arbuscula and D. cladonia by S4 bearing a distinctively long dentate axial margins as compared with S4 bearing slightly dentate to laciniate axial margins (D. arbuscula and D. cladonia).

Dendrophyllia dilatata van der Horst, 1927

Fig. 8M, N

Dendrophyllia dilatata van der Horst, 1927: 2–3, figs 2–4. –Zibrowius and Gili 1990: 44. –Cairns and Keller 1993: 278–279. –Cairns 1994: 89. –Cairns 2001: 34.

Type locality

Off Richards Bay, South Africa (RV ‘Pieter Faure’ stn. 12103: ca. 28°47'00.0"S, 32°20'00.0"E); 97 m (van der Horst 1927).

Type material

The holotype is deposited at the ZMA (Creuwels 2020).

Material examined

SAMC_A073016 (2 specimens): Eastern margin, 31 km from Richards Bay/46 km Mlalazi Estuary, 29°00'54.00"S, 32°15'35.99"E; 500 m. SAMC_A073068 (1 specimen): Eastern margin, 47 km from Cape Vidal/28 km off Mgobezeleni Estuary, 27°43'11.99"S, 32°40'36.11"E; 100 m.

Description

Colony unattached and arborescent, with sympodial branching formed by extra-tentacular budding. Corallum ceratoid to sub-cylindrical, straight to slightly curved. Largest specimen examined (SAMC_A073016) 105.5 mm in H, with ≤ 30 corallites. Calice elliptical (GCD:LCD = 0.9–1.2), with serrated calicular margin. Epitheca absent. Costae well developed at upper corallum, rounded, granular, non-perforate, and separated by thin, porous, intercostal striae. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1–2 > S4 > S3 (48 septa). S1–2 both independent, reaching the columella with straight axial margins, which may also be finely serrated. S1 most exsert, followed by S2. S3 ~ 1/3 the width of S2. S4 slightly wider than flanked S3, arranged in Pourtalès plan: in each half-system a pair of S4 fuses before S3 with jaggedly dentate axial margins. Fossa deep, containing a large, and slightly raised spongy columella.

Distribution

Regional: Eastern margin of South Africa, off Richards Bay towards Cape Vidal; 97–500 m. Elsewhere: Mozambique (Cairns and Keller 1993); 97–132 m.

Remarks

Dendrophyllia dilatata resembles Dendrophyllia sp. 1 in its growth form: i.e., large, dendroid colonies with fairly regular sympodial arranged corallites (Cairns 1994). However, it can be distinguished by having four cycles of septa as compared with three (Dendrophyllia sp. 1), a well-developed Pourtalès plan development (poorly developed in Dendrophyllia sp. 1), lacking P2 which is present in Dendrophyllia sp. 1, and a larger columella (smaller in Dendrophyllia sp. 1).

Dendrophyllia ijimai Yabe & Eguchi, 1934

Fig. 8O, P

Dendrophyllia ijimai Yabe & Eguchi, 1934: 2026. –Eguchi 1965a: 294, 2 figs. –Eguchi 1968: C65 (in part: pl. C16, figs 1, 2, pl. C22, fig. 1). –Kikuchi 1968: 9, pl. 15, fig. 2. –Eguchi and Miyawaki 1975: 54. –Cairns and Keller 1993: 280, fig. 13G. –Cairns 1994: 89, pl. 38C, F. –Cairns 1999a: 133. –Cairns et al. 1999: 26. –Cairns 2001: 34. –Cairns 2004a: 267, 315. –Kitahara and Cairns 2021: 310, 312, figs 164, 165A–C.

Dendrophyllia micranthus. –Eguchi 1965a: 294, fig. 1. –Eguchi 1968: C66, pl. C24, figs 2, 3.

Dendrophyllia minuscula. –van der Horst 1922: 51–52, pl. 8, fig. 30. –Utinomi 1965: 257. –Boshoff 1981: 42. –Tribble and Randall 1986: 159.

Dendrophyllia subcornigera cylindrica Eguchi, 1968: C64–C65, pl. C32, figs 1, 2.

Dendrophyllia subcornigera. –Wells 1984: 215–216, fig. 5.

Dendrophyllia sp. –Zibrowius and Grygier 1985: 123, 126, figs 22, 23.

Dendrophyllia sp. cf. D. ijimai. –Cairns and Zibrowius 1997: 191–192, fig. 29E.

Type locality

Presumably off Japan (Cairns 1994).

Type material

Presumably lost (Cairns 1994).

Material examined

SAMC_A073008 (4 specimens): Eastern margin, 33 km from Port Dunford/38 km off Mlalazi Estuary, 29°05'30.11"S, 32°09'06.11"E; 95 m. SAMC_A090121 (1 specimen): Eastern margin, off 33 km from Port Dunford/37 km off Mlalazi Estuary, 29°08'59.99"S, 32°05'24.00"E; 85 m. USNM 91843 (1 specimen): Eastern margin, 39 km from Cape Vidal/29 km off Mgobezeleni Estuary, 27°47'21.59"S, 32°39'03.60"E; 62–84 m. USNM 91844 (1 specimen): Eastern margin, 28 km from Shaka’s Rock/19 km off Mdlotane Estuary, 29°26'59.99"S, 31°31'11.99"E; 68–70 m.

Description

Colony composed of one elongate, straight to slightly curved axial corallite, from which secondary corallites bud. Secondary corallites robust and bud in all directions, reaching ≤ 130 mm in H. Tertiary corallites small (< 3 mm in H). Corallites circular to slightly elliptical (GCD:LCD = 1.0–1.1), with lanceted calicular margins Costae well defined, slightly ridged, and highly granular. Intercostal furrows deep and porous. Corallum white.

Septa hexamerally arranged in four cycles, S5 occasionally present in some half-systems, in a strongly developed Pourtalès plan according to the formula: S1 ≥ S2 > S4 > S3 > S5 (≤ 60 septa). S1 independent and with straight axial margins. S2 as wide to only slightly smaller than S1, and have slightly sinuous axial margins. Both S1 and S2 extend to columella. S3 narrowest, also with slightly sinuous axial margins. S4 dimorphic in size, with laciniate axial margins: in half-systems without S5, S4 1/5 smaller than S2; however, in half-systems with S5, S4 half the size of S4. S4 arranged in Pourtalès plan: curving towards common S3, and fusing before extending to columella as one septum. However, in half-systems with S5, the S5 is arranged in Pourtalès plan: merging in front of flanked S4, before meandering towards S3 and joining S4 neighbouring S2. Septal faces finely granular. Fossa shallow to moderately deep, with a non-discrete spongy columella.

Distribution

Regional: Eastern margin of South Africa, from off Shaka’s Rock extending towards Cape Vidal; 62–95 m. Elsewhere: Japan (Yabe and Eguchi 1934; Cairns 1994); Philippines; Indonesia (Cairns and Zibrowius 1997); Australia (Cairns 2004a); New Zealand; Red Sea (Scheer and Pillai 1983); Zanzibar (Cairns and Keller 1993); 10–366 m.

Remarks

Dendrophyllia ijimai is the only Dendrophyllia species in the region that has arborescent colonies bearing large axial corallites that give off shorter corallites budding in an irregular form (Cairns 1994). This growth form makes it easily distinguishable from the other South African congeners. Only one other western Pacific species is known to exhibit such a growth form (D. cribrosa Milne-Edwards & Haime, 1851) and D. ijimai may be distinguished by its non-anastomotic branches (and exsert corallites) which are not flushed as in the case of D. cribrosa (see Cairns 1994). As noted by Cairns and Keller (1993), this species may be mistaken with Tubastraea micranthus, but differs in having its septa arranged in a well-developed Pourtalès plan.

Dendrophyllia sp. 1

Fig. 9A, B

Material examined

DSCS–INV 29 (1 fragment): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS–INV 30 (1 fragment Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS–INV 31 (1 fragment): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS–INV 33 (1 fragment): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS–INV 35 (1 fragment): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS–INV 44 (1 fragment): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS_INV 527 (1 specimen): Southern margin, 65 km from Cape St. Francis/70 km off Slang Estuary, 34°47'05.01"S, 24°45'42.30"E; 392–418 m. SAMC_A090158 (1 fragment): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m.

Imagery data

SAM_H1441 (in part: 3 fragments): Southern margin, 3 km from Mosselbaai/11 km off Hartenbos Estuary, 34°10'59.99"S, 22°10'00.00"E; 216 m.

Description

Colonies uniplanar with sympodial budding and dendroid, formed by extra-tentacular budding. Calices occur in lateral plane of branching. Diameter of largest basal branch examined (SAMC_A090158) 7.0 mm. Calice circular, with slightly serrate margin, reaching a maximum CD of 5.5 mm. Costae prominent, but occasionally inconspicuous in some calices. Costal granulation consisting of small spines better developed on terminal branches. Intercostal ridges narrow. Corallum white.

Septa hexamerally arranged in three complete cycles according to the formula: S1 > S2 > S3 (≤ 24 septa). S1 widest and extend to columella with straight and vertical to slightly concave axial margins. S2 ~ 1/2 width of S1, also with straight axial margins, sometimes bearing a palus. S1–2 sometimes slightly exsert. S3 smallest septa and have laciniate axial margins. All septa joining columella deep in fossa. Poorly developed Pourtalès plan only visible deeper in fossa. Fossa deep, containing a tall and thick lamellar columella.

Distribution

Regional: Southern margin of South Africa, off Agulhas extending towards Gouritsmond; 212–445 m.

Remarks

Specimens examined closely resemble the Atlantic Dendrophyllia alternata Pourtalès, 1880

but disagree with its diagnosis in costae being poorly developed, having three cycles of septa, S1–2 sometimes being slightly exsert, and its columella being a solid and elongated rod. However, the uniplanar, dichotomous branching, sympodial budding, and spines on corallum resemble D. alternata.

Figure 9. 

A, B Dendrophyllia sp. 1 (SAMC_A090158, off Knysna, 333 m A close-up calicular view B lateral view C, G Ednapsammia columnapriva sp. nov. C, D (SAM_H1441, off Mossel Bay, 212 m) C calicular view D lateral view E-G (SAMC_A090159, off Gouritsmond, 333 m) E calicular view F lateral view G calicular view H Enallopsammia pusilla (DSCS-INV238, off Gouritsmond, 333 m): lateral view. I, J Enallopsammia rostrata (SAMC_A073270, off Port St Johns, 200 m) I close up calicular view J lateral view K, L Endopachys grayi (DEFF_NANSEN-INV 32, off Shaka’s Rock, 185 m) K calicular view L lateral view M, N Endopsammia philippensis (DIIIb1, off Durban, 442 m) M calicular view N lateral view O, P Heteropsammia cochlea (EIe1, locality data unknown) O calicular view P lateral view. Scale bars: 10 mm (A–C, E–I, K–P); 100 mm (D, J).

Ednapsammia Filander, gen. nov.

Diagnosis

Colony dendroid with sympodial branching, all achieved by extra-tentacular budding (frequently from theca of a parent corallite at an acute angle). Thin epitheca present. Septa normally arranged and granular. Columella absent. Endothecal dissepiments present.

Type species

Ednapsammia columnapriva Filander, 2020, by original designation.

Etymology

The genus name Ednapsammia is to honour the late Dr Edna Molewa, who was instrumental for the declaration of the offshore Phakisa Marine Protected Areas in South Africa. Gender: feminine.

Remarks

Ednapsammia gen. nov. closely resembles Dendrophyllia in having a dendroid colony formed from sympodially arranged branches and well-developed costae. However, it can be differentiated in lacking a Pourtalès plan and columella. Ednapsammia gen. nov. is also morphologically similar to Atlantia López & Capel, 2020, a genus that has dendroid to phaceloid colonies and normally arranged septa, but differs in the presence of a thin epitheca, porosity being prevalent near calicular margin, and lacking a columella.

Ednapsammia columnapriva Filander, sp. nov.

Fig. 9C–G

Type locality

Off Knysna, South Africa (RV ‘Algoa’ stn. DCS13: 35°07'11.34"S, 23°02'41.91"E); 333 m.

Material examined

Holotype – SAMC_A090159 (1 specimen): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. Paratypes–DSCS–INV 226 (1 specimen), DSCS–INV 227 (1 specimen), DSCS–INV 229 (1 specimen), DSCS–INV 231 (1 specimen), DSCS–INV 232 (1 specimen), DSCS–INV 235 (1 specimen), DSCS–INV 238 (1 specimen), SAMC_A090149 (1 specimen): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m.

Imagery data

SAM_H1441: Southern margin, 3 km from Mosselbaai/11 km off Hartenbos Estuary, 34°10'59.99"S, 22°10'00.00"E; 216 m.

Etymology

The species name columnapriva (derived from Latin columna meaning pillar and privus meaning deprived of) alludes to the lack of columella.

Description

Corallum dendroid formed by extra-tentacular budding from base and from theca of parent corallite. Holotype consists of 19 corallites, and 54.7 mm in H. Calice circular to elliptical (GCD:LCD = 1.0–1.1), calicular margin finely laciniate. Thin epitheca, predominantly porous near calicular margin. Costae thick, equal in width, granular, extending to base, and separated by deep intercostal striae. Corallum white.

Septa hexamerally arranged in three cycles, last cycle being incomplete, according to the formula: S1 > S2 > S3 (≤ 21 septa). S1 extend to fossa, sometimes almost meeting opposite septa. S2 ~ 1/3 smaller than S1. Third septal cycle incomplete, usually with only one S3 per half-system. S3 1/3 the width of S2. Primary and secondary septa cycles (S1–2) with straight and vertical axial margins, whilst S3 bears slightly laciniate axial margins. Fossa deep, columella absent.

Distribution

Regional: Southern margin of South Africa, extending from off Mosselbaai towards Gouritsmond; 216–333 m.

Remarks

Examined specimens are easily distinguished from other colonial dendrophylliids by their septa being hexameral and arranged normally (no Pourtalès plan), in three incomplete cycles. Despite that the imaged specimen (SAM_H1441 [in part: 1 specimen]) was not traceable in the Iziko Museums collection. This record nonetheless confirms that such a form was historically collected off the southern margin of South Africa.

Enallopsammia Michelotti, 1871

Diagnosis

Colonial. Arborescent colonies formed by extra-tentacular budding. Corallites often, but not always, unifacially arranged. Coenosteum dense, synapticulotheca porous only near calices and on distal branches. Septa arranged normally. Columella small.

Type species

Coenopsammia scillae Seguenza, 1864, by monotypy

Enallopsammia pusilla (Alcock, 1902)

Fig. 9H

Dendrophyllia (Coenopsammia) pusilla Alcock, 1902a: 113. –Alcock 1902c: 44, pl. 5, figs 38, 38A.

Dendrophyllia (Coenopsammia) profunda. –Alcock 1902c: 43.

Coenopsammia profunda. –von Marenzeller 1904a: 313–314, pl. 18, fig. 24.

Enallopsammia marenzelleri . –Zibrowius 1973: 49–51, pl. 1, figs 1–7. –Zibrowius 1980: 204–205.

Enallopsammi a sp. cf. E. marenzelleri. –Cairns 1982: 57–58, pl. 18, figs 5–6. –Cairns 1995: 128–129, pl. 44, figs G, H.

Enallopsammia pusilla. –Cairns and Zibrowius 1997: 194, fig. 29F. –Cairns 2004a: 316, fig. 12F, G.

Type locality

Off Sulu Archipelago, Philippines (HMS ‘Siboga’ stn. 95: 5°43.5'00"N, 119°40'00"E); 522 m (Alcock 1902a).

Type material

The holotype and the paratypes are deposited at the ZMA (Cairns and Zibrowius 1997).

Material examined

DSCS_INV 34 (1 specimen): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS_INV 40 (1 specimen): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS_INV 42 (1 specimen): Southern margin,140 km from Agulhas/144 km off Ratels Estuary, 36°02'29.58"S, 19°41'24.61"E; 445–463 m. DSCS_INV 223 (1 specimen): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. DSCS_INV 225 (2 specimens): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. DSCS–INV 238 (1 specimen): Southern margin, 116 km from Gouritsmond/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. SAMC_A090153 (1 specimen): 116 km from Knysna/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m.

Description

Irregularly shaped colonies formed by extra-tentacular budding, and firmly attached to substrate by an encrusting base. Buds projecting ≤ 3 mm above branch coenosteum. Corallites circular to slightly elliptical (GCD:LCD = 1.0–1.1), calicular margin serrated. Costae well developed on entire branch, slightly convex, and covered with small granules. Intercostal striae deep, thin, and porous. Corallum white.

Septa hexamerally arranged in three complete cycles according to the formula: S1–2 > S3 (24 septa in two size classes). S1 independent, narrow, extending closer to columella deep in fossa with smooth axial margins. S2 equal in width to S1, and bearing smooth upper axial margins that become finely serrated deep in fossa. S3 slightly less wide than S1–2, sometimes appearing rudimentary, and have dentate axial margins. S3 fuses to adjacent S2 near columella. Fossa deep, containing a spongy columella.

Distribution

Regional: Southern margin of South Africa, extending from Agulhas towards Knysna; 333–463 m. Elsewhere: Philippines; Indonesia; South China Sea (Alcock 1902a; Cairns and Zibrowius 1997); New Zealand (Cairns 1995); Meteor Seamounts; Azores; Nicobar Islands (Zibrowius 1980; Cairns 1982); Bay of Bengal (von Marenzeller 1904a); Australia (Cairns 2004a); 371–805 m.

Remarks

The examined specimens of Enallopsammia pusilla represent a new record for the region, the species differing from its congeners in having three complete cycles, corallites forming on all branch faces, and well-developed costae throughout the corallum (Zibrowius 1973).

Enallopsammia rostrata (Pourtalès, 1878)

Fig. 9I, J

Amphihelia rostrata Pourtalès, 1878: 204, pl. 1, figs 4, 5. –Agassiz 1888: 152, fig. 473. –Gourret 1906: 122, pl. 12, fig. 11A, B.

Stereopsammia rostrata . –Pourtalès 1880: 97, 110–111.

Dendrophyllia (Coenopsammia) amphelioides Alcock, 1902a: 43–44, pl. 5.

Anisopsammia amphelioides . –Vaughan 1907: 156–157, pl. 47, figs 1, 2.

Coenopsammia amphelioides var. cucullata Vaughan, 1907: 157, pl. 48, figs 1–4.

Anisopsammia rostrata . –Gravier 1915: 3. –Gravier 1920: 102, pl. 12, figs 181–185.

Enallopsammia rostrata. –Squires 1959a: 40. –Laborel 1970: 156. –Zibrowius 1973: 44–45, pl. 2, figs 14–15. –Cairns 1978: 9. –Cairns 1979: 186–188, pl. 37, figs 2–3, 6. –Zibrowius 1980: 201–203, pl. 105, figs A–K, pl. 106, figs A–C. –Cairns 1982: 57, pl. 18, figs 1–4. –Cairns 1984: 27–28. –Zibrowius and Grygier 1985: 131, figs 48–50. –Zibrowius 1985: 314, 319, 322, 323. –Zibrowius and Gili 1990: 39–42, pl. 6, figs A–F, pl. 7, figs A–F. –Cairns 1991: 26, pl. 12, fig. B. –Dawson 1992: 45. –Cairns and Parker 1992: 52–53, pl. 18, figs E–I. –Cairns and Keller 1993: 281–282. –Cairns 1994: 92–93, pl. 39, figs D–F. –Cairns 1995: 127–128, pl. 44, figs C–F. –Cairns and Zibrowius 1997: 195. –Cairns 1999a: 134–135. –Cairns et al. 1999: 27. –Romano and Cairns 2000: 1049. –Cairns 2004a: 267, 316. –Le Goff–Vitry et al. 2004: 170, 176. –Cairns 2006: 48. –Kitahara 2007: 504, 505, 511, 513, 516, fig. 5G. –Pires 2007: 269. –Kitahara et al. 2010b. –Kitahara and Cairns 2021: 326–328, figs 173D–G, 174.

Enallopsammia amphelioides . –Zibrowius 1973: 45–46. –Cairns 1979: 187, pl. 40, figs 4, 5. –Zibrowius 1980: 203–204, pl. 106, figs D–I. –Grygier and Newman 1985: 6, fig. 2A–D.

Type locality

Off the Straits of Florida (SSS ‘Blake’ stn. 2: 23°14'00"N, 82°25'00"W); 1472 m (Pourtalès 1878).

Type material

Two syntypes are deposited at the MCZ (Cairns 1979; Zibrowius 1980).

Material examined

SAMC_A073270 (1 specimen): Eastern margin, 39 km from Port St. Johns/13 km off Mkweni Estuary, 31°30'06.11"S, 29°55'12.00"E; 200 m. DSCS_INV 158 (1 specimen): Southern margin, 200 km from Gouritsmond/204 km off Goukou Estuary, 36°09'28.13"S, 21°59'53.81"E; 226–236 m. DSCS_INV 160 (1 specimen): Southern margin, 200 km from Gouritsmond/204 km off Goukou Estuary, 36°09'28.13"S, 21°59'53.81"E; 226–236 m. DSCS_INV 238 (1 specimen): Southern margin, 114 km from Knysna/off Bulolo Estuary, 35°06'11.27"S, 23°02'41.91"E; 333m. DSCS_INV 238 (1 specimen): 116 km from Knysna/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. DSCS_INV 224 (1 specimen): 116 km from Knysna/off Goukamma Estuary, 35°07'11.34"S, 23°02'41.91"E; 333 m. DEFF_SVMEC_INV 261 (1 specimen): Southern margin, 88 km from Oesterbaai/85 km off Tsitsikamma Estuary, 34°52'21.97"S, 24°12'51.01"E; 341–367 m.

Description

Uniplanar colonies formed by extra-tentacular budding, and firmly attached to substrate by an encrusting base. Corallites confined to one face of corallum, arranged uniserially, and projecting ≤ 4.1 mm above branch. Corallites circular to slightly elliptical (GCD:LCD = 1.0–1.1); calicular margin serrate. Costae prominent at calicular margin, often forming a hood that partially covers the calice; becoming poorly-defined in direction to base, slightly convex, and covered with small granules. Intercostal striae deep, thin, and porous. Corallum white.

Septa hexamerally arranged in three complete cycles according to the formula: S1 > S3 > S3 (24 septa). S1 most exsert, narrow, and extending closer to columella deep in fossa with concave and slight serrate axial margins. S2 ¾ the width of S1, and bear serrated axial margins. S3 slightly less wide than S1–2, sometimes appearing rudimentary, and have dentate axial margins. S3 sometimes fusing to adjacent S2 near columella. Fossa deep, containing a rudimentary or trabecular columella.

Distribution

Regional: Southern to eastern margin of South Africa, extending from Knysna towards Port St. Johns; 200–367 m. Elsewhere: Cosmopolitan, except for eastern Pacific and continental Antarctica; 110–2165 m.

Remarks

The examined specimens extend the known distribution of Enallopsammia rostrata in the southwest Indian Ocean (Cairns and Keller 1993) further south from the Madagascar Plateau, thus representing a new record for South Africa. Species is easily distinguished by its uniplanar development with calices in only one side of the colony, (unlike the bushy and irregularly arranged calice of both E. profunda (Pourtalès, 1868) and E. pusilla), and costae being prominent at the calicular margin (instead of being prominent throughout corallum as in the case of both E. profunda and E. pusilla). Although previous specimens of E. rostrata have been reported to display septocostal rostrum (the enlargement of one CS1) (Cairns 1982; Cairns and Zibrowius 1997), this feature is absent in the South African examined representatives (all of which are small colonies).

Endopachys Lonsdale, 1845

Diagnosis

Corallum solitary and free, resulting from transverse division or budding from corallum margin. Corallum straight. Shape of corallum variable, including cuneiform, compressed-cylindrical, and flabellate. Some species with six or twelve robust ridges or flanges aligned to C1 and C2. Epitheca absent. Base of corallum covered with spines. Towards calices, spines usually aligned into narrow costae. Pourtalès plan present. ≤ five cycles of septa. P3 or P4 usually present. Columella elongate, discrete, and spongy.

Type species

Endopachys alatum Lonsdale, 1845, by subsequent designation (Wells 1975).

Endopachys grayi Milne-Edwards & Haime, 1848

Fig. 9K, L

Endopachys grayi Milne-Edwards & Haime, 1848b: 82–83, pl. 1, figs 2, 2A. –Semper 1872: 267. –van der Horst 1922: 68, 74. –van der Horst 1926: 51. –van der Horst 1927: 6–7, pl. 2, fig. 12. –Faustino 1927: 240–241. –Gardiner and Waugh 1939: 241. –Yabe and Eguchi 1942b: 139. –Squires 1961: 17. –Pillai 1972: 213. –Boshoff 1981: 42 (in part). –Cairns 1984: 27, pl. 5, fig. E. –Zibrowius and Grygier 1985: 137. –Veron 1986: 610. –Cairns 1989b: 34. –Cairns 1991a: 24–25, pl. 10, figs I–J, pl. 11, figs A, B. –Cairns and Keller 1993: 276. –Cairns 1994: 84–85, pl. 36, figs E, H pl. 37, fig. I. –Cairns 1995: 121–122, pl. 41, figs C–H. –Cairns and Zibrowius 1997: 185–186. –Cairns 1998: 362, 365. –Cairns 1999: 132, fig. 22F. –Cairns et al. 1999: 27. –Cairns 2001: 25, pl. 7, fig. G. –Cairns 2004a. 276, 316. –Cairns 2006: 49. –Kitahara and Cairns 2021: 329–330, 332, figs 173H–J, 175, 176A–B.

Endopachys weberi Alcock, 1902a: 109–110.

Endopachys oahense Vaughan, 1907: 147–148, pl. 44, fig. 3.

Endopachys japonicum Yabe & Eguchi, 1932e: 388, 399. –Yabe and Eguchi 1932b: 443. –Yabe and Eguchi 1932a: 14–17, pl. 2, figs 1–6. –Eguchi 1934a: 268. –Yabe and Eguchi 1942b: 139. –Eguchi 1965: 293. –Eguchi and Miyawaki 1975: 59.

Endopachys vaughani Durham, 1947: 39–40, pl. 11, figs 6–8, 10, 11.

Endopachys sp. –van der Horst 1922: 68, pl. 8,fig. 4.

Type locality

Unknown (Cairns 1994; Cairns and Zibrowius 1997).

Type material

Presumably lost (Cairns 1994).

Material examined

ORI_EId1(2 specimens): Eastern margin, other locality data unknown. SAMC_A073064 (2 specimens): Eastern margin, 4 km from Cape Vidal/off Groot Brak Estuary, 28°07'30.00"S, 32°36'24.11"E; 75–80 m. SAMC_A073069 (1 specimen): Eastern margin, 48 km from Cape Vidal/21 km off Mgobezeleni Estuary, 27°42'53.99"S, 32°40'54.11"E; 160 m. SAMC_A073213 (1 specimen): Eastern margin, 29 km from Durban/14 km off Mbokodweni Estuary, 30°06'24.12"S, 31°00'47.88"E; 160–170 m. SAMC_A073266 (5 specimens): Southern margin, 2 km from Stilbaai/1 km off Goukou Estuary, 34°22'55.26"S, 21°25'25.49"E; 88 m. SAMC_A090114 (1 specimen): Eastern margin, 16 km from Port St. Johns/13 km off Bulolo Estuary, 31°45'00.00"S, 29°26'59.99"E; 70 m. SAMC_A090115 (9 specimens): Eastern margin, 29 km from Durban/22 km off Mdloti Estuary, 29°46'00.00"S, 31°16'59.99"E; 110–130 m. SAM_H1420 (5 specimens): Eastern margin, 39 km from Mtunzini/8 km off Zinkwasi Estuary, 29°13'00.00"S, 31°30'00.00"E; 66–77 m. SAM_H1427 (5 specimens): Eastern margin, 2 km from Durban/8 km off Umgeni Estuary, 29°52'00.00"S, 31°00'00.00"E; 99 m. SAM_H1476 (3 specimens): Eastern margin2 km from Durban/8 km off Umgeni Estuary, 29°52'00.00"S, 31°00'00.00"E; 99 m. SAM_H3120 (28 specimens): Eastern margin, 39 km from Mtunzini/8 km off Zinkwasi Estuary, 29°13'00.00"S, 31°30'00.00"E; 73 m. SAM_H3121 (2 specimens): Eastern margin, 39 km from Mtunzini/8 km off Zinkwasi Estuary, 29°13'00.00"S, 31°30'00.00"E; 73 m. SAM_H3122 (3 specimens): Eastern margin, 6 km from Durban/9 km off Umgeni Estuary, 29°52'59.99"S, 31°03'05.00"E; 86 m. SAM_H3123 (2 specimens): Eastern margin, 9 km from Shaka’s Rock/2 km off Tongati Estuary, 29°34'00.00"S, 31°10'59.99"E; 66 m. SAM_H4593 (2 specimens): Eastern margin, 26 km from Port St. Johns/off Bulolo Estuary, 29°34'47.99"S, 31°41'59.99"E; 138 m. DEFF_NANSEN–INV 32 (8 specimens): Eastern margin, 30 km from Shaka’s Rock/31 km off Tongati Estuary, 29°43'11.99"S, 31°25'47.99"E; 185 m. USNM 91812 (3 specimens): Eastern margin, 29 km from Shaka’s Rock/24 km off Mdloti Estuary, 29°45'54.00"S, 31°18'11.88"E; 105 m. USNM 91813 (1 specimen): Eastern margin, 44 km south of Ponta Do Ouro/28 km off Kosi Bay Estuary, 27°14'35.88"S, 32°48'47.87"E; 74 m.

Imagery data

PF 10983 (4 specimens): Eastern margin, 19 km from Shaka’s Rock/3 km off Mdloti Estuary, 29°38'59.99"S, 31°07'59.99"E; 71–73 m.

Description

Corallum (anthocyathus) variable in shape, including cuneiform, compressed-cylindrical, and flabellate. Corallum free, compressed and usually with a rounded base on GCD plane. Largest specimen examined (DEFF_NANSEN–INV 32) 37.2 × 19.7 mm in CD (excluding crests), and 25.8 mm in H. Thecal edges project outward to form slightly porous, and straight to slightly sinuous thecal crests. Upper thecal crest meets in acute angle, and often support one bud obliquely oriented (sometimes supporting ≤ four buds). Crest wider at lower half of corallum. Costae equal in width and flat to slightly convex, becoming progressively less developed towards base. Intercostal striae thin, narrow, and porous. Corallum white.

Septa hexamerally arranged in five complete cycles according to the formula: S1–2 > S3 > S5 > S4 (96 septa). S1–2 thick, most exsert, porous at upper distal margin, and extend towards columella with straight, vertical, and finely dentate axial margins. S3 less exsert and extend ¾ the distance of S1–2 with dentate axial margins. S4 variable in development: if not flanked by a pair of S5, both S4 in a half-system bend towards each other meeting before S3 in a characteristic Pourtalès plan, but, if flanked, S4 have highly laciniate axial margins. S5 dimorphic in development: those adjacent to S1 more exsert and wider than those adjacent to S2. Each Pourtalès plan terminates in a palus-like structure. All septal faces granular. Fossa deep, containing septum in the elongate and spongy columella.

Distribution

Regional: Eastern margin of South Africa, extending from off Port St. Johns towards Kosi Bay Estuary (44 km south of Ponta Do Ouro: Mozambique); 66–170 m. Elsewhere: Mozambique; Tanzania (Cairns 1989b); Zanzibar; Mauritius (van der Horst 1926); Saya de Malha; Arabian Sea (Gardiner and Waugh 1939) Cairns and Keller 1993); Philippines and Indonesia (Cairns and Zibrowius 1997; Kitahara and Cairns 2021), Malaysia; Australia (Cairns 2004); New Caledonia (Zibrowius and Grygier 1985); Wallis and Futuna; Vanuatu (Cairns 1999a); Hawaii; United States; Japan (Cairns 1984; Zibrowius and Grygier 1985; Cairns 1994); 37–550 m.

Remarks

Endopachys grayi is easily recognised by its corallum shape and presence of lateral thecal crest. Overall, Tropidocyathus lessoni (Michelin, 1842) resembles E. grayi, but belongs to a different family (Turbinoliidae) and is easily distinguished by having a solid corallum (not porous as dendrophylliids). Within the genus, only two species, E. grayi and Endopachys bulbosa Cairns & Zibrowius, 1997, are known and they differ in: (i) distribution (E. grayi = Indo-Pacific vs. E. bulbosa = South Pacific), (ii) basal thickness (E. grayi = 3 mm vs. E. bulbosa = 4 mm), (iii) thecal face angle in relation to height (E. grayi = face angle being low initially and broadens with height vs. E. bulbosa = face angle being high initially and decreases with height), (iv) septal exsertness (E. grayi = S1–2 being ≤ 3.0 mm vs. E. bulbosa = S1–2 being ≤ 5.0 mm), (v) costae (E. grayi = poorly-defined vs. E. bulbosa = well-defined costae), and (vi) presence or absence of pali (E. grayi = pali present vs. E. bulbosa = pali absent) (Cairns and Zibrowius 1997). The DEFF_NANSEN–INV 32 specimen appears to be the largest E. grayi recorded to date. It is 37.2 × 19.7 mm in CD (excluding crests), 25.8 mm in H, and has a pair of S6, totalling 98 septa.

Endopsammia Milne-Edwards & Haime, 1848

Diagnosis

Corallum solitary, conical to subcylindrical, and firmly attached. Epitheca thin, covering most of the synapticulotheca. Underlying epitheca weakly costate, covered with low granules. Septa arranged in normal insertion pattern in ≤ four cycles. Axial margins of all septa coarsely dentate to laciniate. Columella spongy, non-discrete. Tabular endothecal dissepiments present in elongate coralla.

Type species

Endopsammia philippensis Milne-Edwards & Haime, 1848, by monotypy.

Endopsammia philippensis Milne-Edwards & Haime, 1848

Fig. 9M, N

Endopsammia philippensis Milne-Edwards & Haime, 1848b: 91, pl. 1, figs 5, 5A. –Faustino 1927: 243–244, pl. 77, figs 5, 6. –Pillai and Scheer 1976: 71–72. –Cairns and Zibrowius 1997: 188, fig. 28C–E. –Cairns 2001: 23, pls 5H, I . –Cairns 2004a: 316.

Balanophyllia regularis. –van der Horst 1922: 63. –van der Horst 1926: 50, pl. 3, figs 10, 11.

Endopsammia philippinensis. –Wells 1964: 118, pl. 2, figs 12, 13. –Cairns 1991: 26. –Cairns and Keller 1993: 221.

Certotrochus brunneus. –Boshoff 1981: 36.

Type locality

Philippines, depth unknown (Milne-Edwards and Haime 1848b).

Type material

The holotype is deposited at the NHMUK or MNHN (Cairns and Zibrowius 1997; Cairns 2001, 2004a).

Material examined

ORI_DIIIb1 (2 specimens): Eastern margin, 33 km from Durban/31 km off Beachwood Mangroves, 29°55'00.00"S, 31°19'59.99"E; 442 m.

Imagery data

SAM_H1576 (1 specimen): Eastern margin, 35 km from Port Edward/10 km off Mtentu Estuary, 31°18'00.00"S, 29°58'00.00"E; depth unknown.

Description

Corallum conical to sub-cylindrical, relatively small, attached to substrate by a thin encrusting base. Calice circular and calicular margin lanceted. Largest specimen examined (ORI_DIIIb) 7.80 × 7.70 mm in CD, 8.20 mm in H. Epitheca thin and extending ¾ of lower corallum. Theca near calicular margin porous and non-costate. Corallum white.

Septa hexamerally arranged in four cycles, last cycle being incomplete, according to the formula: S1 ≥ S2 > S3 > S4 (≤ 30 septa). S1 slightly exsert and reaches columella. Higher cycle septa (S2–4) progressively less exsert (if at all). S2 equal or ¾ the width of S1. S3 being 1/4 the width of S2, sometimes rudimentary in some half-systems without S4. S4, if present, rudimentary. All septa bear dentate to slightly laciniate axial margins. Septa faces bearing spines. Fossa shallow, containing a spongy or rudimentary columella.

Distribution

Regional: Eastern margin of South Africa, from north of Port Edward extending towards Durban; 442 m. Elsewhere: Philippines (Milne-Edwards and Haime 1848b); Indonesia (Cairns and Zibrowius 1997); Indian Ocean islands (Maldives, Zanzibar, Seychelles, Chagos) (van der Horst 1926; Pillai and Scheer 1976); tropical Pacific Ocean; Loyalty Islands; Queensland; Papua New Guinea (Wells 1964; Cairns and Zibrowius 1997; Cairns 2004a); 0–73 m.

Remarks

Among congeners (Endopsammia regularis (Gardiner, 1899) and E. pourtalesi (Durham & Barnard, 1952)), E. philippensis closely resembles E. regularis in having septa hexamerally arranged in four incomplete cycles, in corallum being solitary (relatively squat), and bearing an epitheca that extends ¾ of the corallum height. However, they differ as E. regularis has exsert septa. Although E. pourtalesi also has septa arranged hexamerally in four incomplete cycles, this species is distinguished from E. philippensis by its quasi-colonial coralla with slender corallites, presence of endothecal dissepiments, and bearing non-exsert septa (Cairns 1991). Endopsammia philippensis has previously been documented in the southwest Indian Ocean (van der Horst 1922, 1026; Cairns and Keller 1993), therefore the examined Boshoff (1981) material (ORI_DIIIb1) represents a species range extension further south of Zanzibar, and a new record for South Africa.

Heteropsammia Milne-Edwards & Haime, 1848

Diagnosis

Corallum solitary or colonial, latter condition achieved by intratentacular budding and resulting in ≤ 40 contiguous corallites. Adult corallum free and mobile, globular in shape. Coralla usually attached to small gastropod shells, these subsequently overgrown. Each specimen apparently in obligate symbiosis with a sipunculid worm, which lives in base of corallum. Epitheca absent. Synapticulotheca covered with finely serrate ridges, usually one to three ridges per corresponding septum (not considered to be conventional costae). Pourtalès plan present. Paliform lobes may be present. Columella spongy, not discrete. Endotheca absent.

Type species

Heteropsammia michelinii Milne-Edwards & Haime, 1848, by monotypy.

Heteropsammia cochlea (Spengler, 1781)

Figs 9O, P, 10A, B

Madrepora cochlea Spengler, 1781: 240–248, figs A–D.

Psammoseris cylicioides Tenison-Woods, 1879: 10–11. –Tenison-Woods 1880: 297–299.

Heteropsammia michelini . –Kent 1893: 106, 177. –Wells 1964: 108, 120.

Heteropsammia cochlea. –van der Horst 1922: 66–67. –van der Horst 1926: 51. –Veron and Pichon 1980: 416–420, figs 727, 729. –Zibrowius and Grygier 1985: 129, figs 43–44. –Veron 1986: 576–577. –Hoeksema and Best 1991: 234–237, figs 24–28 (in part). –Cairns 1998: 406–408. –Cairns 1999a: 132–133. –Cairns et al. 1999: 27. –Veron 2000: 407. –Cairns 2001: 19–20, pl. 2, figs H–J, pl. 3, figs A–E –Cairns 2004a: 316. –Kitahara and Cairns 2021: 334, 336, figs 176C,178A–C.

Heterocyathus aequicostatus. –Boshoff 1981: 37.

Heteropsammia aphrodes. –Boshoff 1981: 42.

Heteropsammia cochleata. –Cairns 2009: 25.

Type locality

Off Tranquebar, southeastern India, depth unknown (Spengler 1781).

Type material

The type specimen is presumably lost (Cairns 2004a).

Material examined

ORI_DIIIe1_3 (1 specimen), ORI_EIe1 (10 specimens): Eastern margin, locality data unknown. SAMC_A073006 (1 specimen): Locality data unknown; 17 m. SAMC_A073051 (6 specimens): Eastern margin, , 41 km south of Ponta Do Ouro/26 km off Kosi Bay Estuary, 27°13'00.11"S, 32°49'41.87"E; 72 m. SAMC_A073061 (8 specimens): Eastern margin, 25 km from Cape Vidal/23 km off St Lucia Estuary, 27°54'42.11"S, 32°36'42.11"E; 42–50 m. SAMC_A073065 (1 specimen): Eastern margin, 67 km from CAPE VIDA/6 km off Mgobezeleni Estuary, 27°32'48.12"S, 32°42'00.00"E; 50 m. SAMC_A073072 (3 specimens): Eastern margin, 36 km from Cape Vidal/33 km Mgobezeleni Estuary, 27°48'54.00"S, 32°38'24.00"E; 52 m. SAMC_A073084 (3 specimens): Locality data unknown. SAMC_A073086 (1 specimen): Eastern margin, 329 km from Port Edward/330 km off Mtentu Estuary, 32°55'18.12"S, 32°55'18.12"E; 49 m. SAMC_A073089 (1 specimen): Eastern margin, 67 km south of Ponta Do Ouro/14 km off Mgobezeleni Estuary, 27°26'12.11"S, 32°44'12.11"E; 55–60 m. SAMC_A073090 (2 specimens): Eastern margin, 69 km from Cape Vidal/5 km off Mgobezeleni Estuary, 27°31'36.12"S, 32°41'48.11"E; 40 m. SAMC_A073091 (2 specimens), SAMC_A073093 (1 specimen): Locality data unknown. SAMC_A073095 (3 specimens): Eastern margin, 66 km south of Ponta Do Ouro/15 km off Mgobezeleni Estuary, 27°25'59.87"S, 32°44'30.12"E; 55–100 m. SAMC_A073098 (1 specimen): Locality data unknown. SAMC_A073100 (5 specimens): Eastern margin, 42 km south of Ponta Do Ouro/27 km off Kosi Bay Estuary, 27°13'30.00"S, 32°49'30.00"E; 78 m. SAMC_A073112 (2 specimens): Eastern margin, 35 km from Cape Vidal/32 km off St Lucia Estuary, 27°49'41.87"S, 32°38'12.11"E; 54 m. SAMC_A073115 (5 specimens): Eastern margin, 35 km from Cape Vidal/32 km off St Lucia Estuary, 27°49'41.87"S, 32°38'12.11"E; 47–50 m. SAMC_A073118 (12 specimens): Eastern margin, 25 km off Cape Vidal/23 km off St Lucia Estuary, 27°54'42.11"S, 32°36'42.11"E; 42–50 m; SAMC_A073124 (1 specimen): Eastern margin, 66 km south of Ponta Do Ouro/15 km off Mgobezeleni Estuary, 27°25'54.11"S, 32°44'17.88"E; 46–66 m. SAMC_A073142 (1 specimen): Eastern margin, 19 km south of Ponta Do Ouro/12 km off Kosi-Kumpungwini (Sifungwe) Estuary, 27°01'05.87"S, 32°55'12.00"E; 78 m. SAMC_A073143 (1 specimen): Eastern margin, 38 km south of Ponta Do Ouro/24 km off Kosi Bay Estuary, 27°11'24.00"S, 32°51'00.00"E; 100 m. SAMC_A073156 (1 specimen): Eastern margin, 39 km from Cape Vidal/29 km off Mgobezeleni Estuary, 27°47'23.99"S, 32°38'53.87"E; 65–70 m. SAMC_A073170 (31 specimen): Eastern margin, 39 km from Cape Vidal/29 km off Mgobezeleni Estuary, 27°47'23.99"S, 32°38'53.87"E; 65–70 m. SAMC_A073192 (1 specimen): Eastern margin, 69 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°31'48.00"S, 32°42'47.99"E; 70 m. SAMC_A073193 (1 specimen): Eastern margin, 67 km from Cape Vidal/6 km off Mgobezeleni Estuary, 27°32'30.11"S, 32°42'00.00"E; 48–58 m. SAMC_A073205 (3 specimens): Eastern margin, 65 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°33'47.88"S, 32°42'15.12"E; 64 m. SAMC_A073214 (30 specimens): Eastern margin, 37 km from Cape Vidal/32 km off Mgobezeleni Estuary, 27°48'47.88"S, 32°38'53.87"E; 50 m. SAMC_A073218 (1 specimen): Eastern margin, 42 km south of Ponta Do Ouro/27 km off Kosi Bay Estuary, 27°13'30.00"S, 32°49'30.00"E; 74 m. SAMC_A090117 (1 specimen): Eastern margin, 66 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°33'11.88"S, 32°42'47.87"E; 85 m. SAMC_A090118 (2 specimens): Eastern margin, 59 km from Cape Vidal/10 km off Mgobezeleni Estuary, 27°37'00.00"S, 32°40'54.00"E; depth unknown. SAM_H814 (21 specimen): Eastern margin, 19 km from Shaka’s Rock/3 km off Mdloti Estuary, 29°38'59.99"S, 31°07'59.99"E; depth unknown. USNM 90242 (1 specimen, sub-sample of SAMC_A073086): Eastern margin, 329 km from Port Edward/330 km off Mtentu Estuary, 32°55'18.12"S, 32°55'18.12"E; 49 m. USNM 90426 (1 specimen): Eastern margin, , 41 km south of Ponta Do Ouro/26 km off Kosi Bay Estuary, 27°13'00.11"S, 32°49'41.87"E; 72 m. USNM 90434 (5 specimens, sub-samples of SAMC_A073205): Eastern margin, 65 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°33'47.88"S, 32°42'15.12"E; 64 m. USNM 90435 (1 specimen): Eastern margin, 67 km from Cape Vidal/6 km off Mgobezeleni Estuary, 27°32'42.00"S, 32°42'29.88"E; 63 m. USNM 90439 (11 specimen): Eastern margin, 64 km from Cape Vidal/7 km off Mgobezeleni Estuary, 27°34'12.00"S, 32°42'06.12"E; 62–64 m.

Description

Corallum solitary, encapsulating a gastropod shell, with one large efferent pore projecting downward from base of corallum, and several smaller pores on lower theca. Calice elliptical (GCD:LCD = 1.3–1.5); calicular margin lanceted. Largest specimen examined (SAMC_A073170) 11.4 × 9.1 mm in CD, and 13.1 mm in H. Upper theca highly porous and usually composed of discontinuous interconnected granular rows. Lower theca granular. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S1–2 > S4 > S3 (48 septa). S1–2 highly exsert, equally wide, with straight axial margins that fuse to columellar elements. S3 least exsert septa and bear serrate axial margins. S4 fuse to neighbouring septa near calicular margin forming well-developed and porous triangular lancets. In each half-system, a pair of S4 fuses before flanked S3 and reach columella as one septum. S4 upper axial margins highly concave becoming almost vertical after fusing to adjacent S4 near columella. Fossa of moderate depth, containing a spongy columella.

Distribution

Regional: Eastern margin of South Africa, extending from off Port Edward towards Kosi-Kumpungwini (Sifungwe) Estuary (19 km south of Ponta Do Ouro: Mozambique); 17–100 m. Elsewhere: Widespread throughout the tropical Indo-Pacific (Veron and Pichon 1980; Hoeksema and Best 1991; Cairns 1999a); 6–622 m.

Remarks

Heteropsammia cochlea is a symbiotic facultative species, known to be zooxanthellate or azooxanthellate in shallow waters and those found in deeper waters being azooxanthellate (Hoeksema and Mathews 2015). The variation in corallum type has resulted in a long standing discussion regarding the number of species represented within Heteropsammia. As such, two valid species are recogniszed (H. cochlea and H. eupsammides (Gray, 1849)) on the basis of the number of calices (monostomous or polystomous) (Hoeksema and Best 1991; Hoeksema 1993). Whilst Cairns (2001) on the other hand, cautiously acknowledges three (H. cochlea, H. eupsammides (Gray, 1849), and H. moretonensis Wells, 1964) in his generic revision of dendrophylliids. Irrespective of this, two species are represented in South Africa, and the differences between the two are elaborated in the account of H. eupsammides below. Zibrowius and Grygier (1985) have reported H. cochlea from the Great Barrier Reef and Somalia to host endoparasites of the ascothoracid crustacean Petrarca okadai, an observation not evident in the South African specimens examined. Additionally, specimens examined include zooxanthellate representatives (> 40 m) (Hoeksema and Best 2015) and should therefore not be considered in biodiversity assessments focusing on azooxanthellate forms.

Heteropsammia eupsammides (Gray, 1849)

Fig. 10C, D.

Heterocyathus eupsammides Gray, 1849: 77, pl. 2, figs 5–7.

Heteropsammia geminata Verrill, 1870: 370–371, fig. 1. –van der Horst 1922: 67. –Faustino 1927: 239–240, pl. 76, figs 11–13.

Heteropsammia multilobata Moseley, 1881: 196–197, pl. 12, figs 1–3.

Heteropsammia michelini. –van der Horst 1926: 51, pl. 3, figs 14–20. –Gardiner and Waugh 1938: 241.

Heteropsammia cochlea . –Veron and Pichon 1980: 416–420.

Heteropsammia eupsammides . –Hoeksema and Best 1991: 237–240. –Kitahara and Cairns 2021: 336–338, figs 178D–I, 179.

Type locality

China Sea (Verrill 1870).

Type material

Unknown.

Material examined

SAMC_A090129 (1 specimen): Eastern margin, 69 km from Cape Vidal/5 km off Mgobezeleni Estuary, 27°31'36.12"S, 32°41'48.11"E; 40 m.

Imagery data

ZMK (in part: 2 specimens): Locality data unknown.

Description

Corallum solitary, encapsulating a gastropod shell, with one large efferent pore projecting downward from base of corallum, and several smaller efferent pores on lower theca. Calice irregular to elliptical (GCD:LCD = 1.5), calicular margin lanceted. Only specimen examined (SAMC_A090129) 12.9 × 8.7 mm in CD and 8.9 mm in H. Upper theca highly porous and usually composed of discontinuous but interconnected granular rows. Lower theca becoming more granular. Corallum white.

Septa hexamerally arranged in six incomplete cycles according to the formula: S1–2 > S3 > S5 > S4 (107 septa). S1–2 highly exsert, equally wide, with lower axial margins fusing to columellar elements. Higher cycle septa (S1–2) progressively less exsert. S3 1/2 the width of S1–2. S4 dimorphic in development: those in half-systems lacking S6 ~ 1/5 the width of S3; but those in half-systems with S6 1/3 the width S3. S5 also dimorphic, being double the width of S4 in half-systems lacking S6 and merging in front of S4 before meeting columella as one septum. However, in half-systems with S6, those S5 neighbouring S3 are double the width of S4, but those flanked by S6 are rudimentary. S6 1/3 the width of S4 and fuse before flanked S5, reaching columella as one septum. All septa with straight and slightly serrate axial margins. Fossa of moderate depth, containing a spongy columella.

Distribution

Regional: Eastern margins of South Africa, off Cape Vidal; 40 m. Elsewhere: China Sea (Verrill 1870); Indonesia (van der Horst 1922; Hoeksema and Best 1991); Philippines (Moseley 1881); New Caledonia (Kitahara and Cairns 2021); Australia (Veron and Pichon 1980); Maldives; Seychelles (van der Horst 1926); and Zanzibar (van der Horst 1926; Gardiner and Waugh 1938); 38–281 m.

Remarks

Heteropsammia eupsammides is one of the apozooxanthellate species, like H. cochlea, it exhibits a symbiotic relationship with sipunculan worms. These two species may be distinguished from one another by whether they are monostomous or polystomous (Cairns 2001: Heteropsammia eupsammides is polystomous [i.e., has > 2 calices formed by intra-stomodeal budding] (Hoeksema and Best 1991). The examined specimen of H. eupsammides appears to be in its early stages of calyx separation and therefore can be mistaken with H. cochlea. However, can be distinguished by the number of cycles (six incomplete cycles in H. eupsammides as compared with four complete ones in H. cochlea) and a higher number of septa (H. eupsammides :107 vs. H. cochlea :48), which is a resuly of varying stomata (smaller in H. eupsammides). Apart from the number of septa cycles, the septa profile varies between the two species, whereby H. eupsammides bears S3 > S5 > S4, with straight to slightly serrated axial margins, and lacking porous triangular lancets (as compared with H. cochlea which has S4 > S3, with consistent straight axial margins, and bearing triangular lancets). This record represents a range southern range extension from Zanzibar.

Figure 10. 

A, B Heteropsammia cochlea (ORI_EIe1, locality data unknown) A calicular view B lateral view C, D Heteropsammia eupsammides (ZMK, locality data unknown) C calicular view D calicular view E–H Pourtalopsammia togata E, F (SAM_H2829, off East London, 146–238 m) E calicular view F lateral view G, H (SAM_H2831, off Cintsa, 159 m) G calicular view H lateral view I, J Rhizopsammia annae (SAM_H1497, off the Agulhas, depth unknown) I calicular view J lateral view K, L Rhizopsammia compacta (SAMC_A073041, off Kosi Bay Estuary, 69 m) K calicular view L lateral view M, N Rhizopsammia verrilli (SAM_H1502, off Richards Bay, 165 m) M calicular view N lateral view O Tubastraea coccinea (ORI_EIb4, Isipingo; depth unknown) P Tubastraea sp. cf. diaphana (SAM_H5103, Cape Vidal, 59 m): Calicular view. Scale bars: 10 mm.

Pourtalopsammia Cairns, 2001

Diagnosis

Corallum solitary, conical to subcylindrical (sometimes scolecoid), and attached. Epitheca well developed, covering basal synapticulotheca; coenosteum distal to epitheca covered with thin, hispid ridges. Septa arranged in normal insertion pattern (not Pourtalès plan); three or four cycles of septa; axial margins of S1 highly sinuous. Columella absent. Endothecal dissepiments absent.

Type species

Balanophyllia togata van der Horst, 1927, by monotypy.

Pourtalopsammia togata (van der Horst, 1927)

Fig. 10E–H

Balanophyllia togata van der Horst, 1927: 5–6.

Thecopsammia togata . –Wells 1935: 531.

Trochopsammia togata. –Cairns and Keller 1993: 275–276. –Cairns 2000: 22, pl. 4, figs F–I, pl. 5, fig. A.

Pourtalopsammia togata. –Cairns 2001: 22.

Type locality

Off Buffalo River mouth, South Africa; 567 m (van der Horst 1927).

Type material

The type material is presumably at the BMNH (Cairns 2001).

Material examined

DEFF_AI2–INV 135 (1 specimen): Eastern margin, 37 km from Cintsa/21 km off Cwili Estuary, 32°49'59.99"S, 28°30'00.00"E; 228 m. DSCS–INV 124 (1 specimen): Southern margin, 192 km from Agulhas/198 km off De Mond-Heuningnes Estuary, 36°15'34.13"S, 21°11'46.61"E; 513 m. DSCS–INV 364 (1 specimen): Southern margin, 68 km from Cape St. Francis/70 km off Slang Estuary, 34°47'35.77"S, 24°38'35.69"E; 520 m. DSCS–INV 424 (1 specimen): Southern margin, 65 km from Cape St. Francis/78 km off Slang Estuary, 34°43'40.13"S, 25°08'53.47"E; 622 m. DSCS–INV 425 (1 specimen): Southern margin, 65 km from Cape St. Francis/78 km off Slang Estuary, 34°43'40.13"S, 25°08'53.47"E; 622 m. DSCS–INV 477 (2 specimens): Southern margin, 58 km from Port Alfred/35 km off Mgwalana Estuary, 33°39'10.19"S, 27°29'57.58"E; 304 m. DSCS–INV 516 (17 specimens): Southern margin, 58 km from Port Alfred/35 km off Mgwalana Estuary, 33°39'10.19"S, 27°29'57.58"E; 304 m. DSCS–INV 569 (8 specimens): Southern margin, 65 km from Cape St. Francis/70 km off Slang Estuary, 34°47'05.01"S, 24°45'42.30"E; 304 m. SAMC_A073015 (3 specimens): Southern margin, 32 km from Mazeppa Bay/19 km MenduEstuary, 32°25'00.11"S, 28°58'18.11"E; 330–340 m. SAMC_A073042 (1 specimen): Eastern margin, 53 km from Shaka’s Rock/46 km off Zinkwasi Estuary, 29°32'53.88"S, 31°47'12.11"E; 200 m. SAMC_A073105 (3 specimens): Eastern margin, 36 km from Cape Vidal/32 km off Mgobezeleni Estuary, 27°48'54.00"S, 32°38'24.00"E; 52 m. SAMC_A073157 (2 specimens): Eastern margin, 10 km from Port Edward/24 km off Bilanhlolo Estuary, 31°05'48.11"S, 30°18'47.88"E; 140 m. SAMC_A073158 (2 specimens): Eastern margin, 10 km from Port Edward/24 km off Bilanhlolo Estua