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

Coelosmilia fecunda Pourtalès, 1871, by monotypy.

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

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

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.

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 C_1–2 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 S_1, also with straight axial margins bearing slender ribbons. S₃ not exsert, ²/₃ the size of S_2, also bearing ribbons that intermingle with those of S_2. 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.

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.

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.

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

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

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.

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.

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. C_1–2 wider than C_3–4. 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 S_3, 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.

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.

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.

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 S_1–3. Columella trabecular.

Aulocyathus juvenescens von Marenzeller, 1904a, by monotypy.

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).

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

None.

‘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.

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 S_1. S_3–4 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 S_2. If present, S₄ ½ the size of S_3. All septa with vertical and slightly sinuous axial margin, S_3–4 bearing dentate axial margins deeper on fossa. All septal faces with fine granulations. Fossa deep containing a rudimentary columella.

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.

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).

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

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

ORI_DIa1 (2 specimens): no locality data.

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). S_1–2 most exsert septa. S₁ extend almost towards centre of fossa, with vertical to slightly sinuous axial margin. S₂ ¼ less wide than S_1, also has vertical axial margin, and being granular deeper in fossa. S_3–4 progressively less exsert (if at all). S₃ dimorphic in size: those half-systems lacking S₄ only ½ the width of S_2, while those flanked by S₄ attain almost the same width of S_2. If present, S₄ ½ the width of S_3. Axial margins of S_3–4 dentate. Septal faces finely granular. Fossa deep containing a rudimentary columella.

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

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.

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 with thecal margin spines or crests.

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

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

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

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.

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.

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 C_1–2 of their studied specimens in relation to the holotype.

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

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

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.

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.

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.

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 without thecal margin spines or crests.

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

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

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

None.

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.

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.

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

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

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.

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). S_1–2 most exsert, equal in width, and almost reach columella with vertical to slightly sinuous axial margins. S_3–4 progressively narrower and less exsert. S₃ ¾ the width of S_1–2, each bearing a sinuous pali (12 P₃), and consistently 1 mm in width. S₄ slightly smaller than S_3. 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.

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.

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).

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

Types are presumably deposited at the IM.

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.

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 C_1–2 more prominent and double the width of C_3–4. C_3–4 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 S_1–2 and forming a slightly lanceted calicular margin. S₃ ²/₃ the width of S_2, 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.

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).

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 S_4. 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.

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

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

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.

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. C_1–3 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: S_1–3 > S₄ > S₅ (96 septa). S_1–3 highly exsert, with straight to slightly sinuous axial margins. S₄ less exsert than S_1–3, 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.

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.

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.

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

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

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

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: S_1–2 > S₃ > S₄ > S₅ (52 total septa). S_1–2 equal in width and most exsert septa, with straight to slightly sinuous axial margins. S_3–4 progressively less exsert, with sinuous axial margins, S₃ being most sinuous. S₃ ¹/₃ less wide than S_1–2, 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 S_3; whilst in half systems without S₅ S₄ is ¹/₅ less wide than S_3. S₅ slightly less wide than S_4. Septal and palar faces covered with granules arranged in a perpendicular manner. Fossa moderately deep, containing a fascicular columella composed of 12 twisted elements.

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.

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.

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

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

None.

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.

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. C_1–3 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: S_1–2 > S₃ > S₄ > S₅ (96 septa). S_1–2 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 S_1–2. S₄ ~ ¾ the width of S₃, each bearing a broad pali (24 P₄), with vertical inner margin. S₅ ¾ the width of S_4. 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.

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.

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.

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).

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

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.

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 S_1, 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.

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.

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.

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

Syntypes are deposited at the BMNH (Cairns 1994).

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.

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. S_2–3 becoming progressively narrower and less exsert. All septa bear sinuous axial margins, sinuosity being extreme in S_1–2. S₂ bearing a highly sinuous and thick pali (8 P₂). Fossa shallow containing a prominent fascicular columella.

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.

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.

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).

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

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.

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: S_1–2 > S₃ ≥ S₄ (48 septa). S_1–2 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 S_1–2, 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.

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.

All the imaged specimens represented here have: (i) septa hexamerally arranged in four cycles; (ii) S_1–3 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.

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).

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

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.

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: S_1–2 > S₃ > S₄ (48 septa). S_1–2 thick, slightly exsert, and extend to columella deep in fossa with straight to slightly sinuous axial margins. S_2–3 progressively less exsert. S₃ axial margin more sinuous than remaining septa. S₃ ~ ²_/3 the width to S_1–2, 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.

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.

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.

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

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

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.

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.

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: S_1–2 > S₃ > S₄ (48 septa). S_1–2 slightly exsert and extend to columella. S₃ ²/₃ the width of S_1–2, and bear a sinuous pali (12 P₃). S₄ ¼ the width of S_3. All septal have sinuous axial margins, S₃ being the most sinuous. Higher cycle septa (S_3–4) progressively less exsert (if at all). Fossa relatively deep, with a fascicular columella composed of 4–6 ribbon-like elements.

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.

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).

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).

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

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.

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 C_1–2, which are prominent and high. C_1–2 slightly wider than C_3–4. 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 > S_3–4 (64 septa). S₁ most wide, with straight axial margins that meet columella deep in fossa. S₂ slightly less wide than S_1, with straight to slightly sinuous axial margins. S_1–2 most exsert. S_3–4 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.

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.

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, S_1–2 being equally exsert, and also in septal formula: S₁ > S₂ > S_3–4 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 P_3. 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.

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

Crispatotrochus inortatus Tenison-Woods, 1878, by monotypy.

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

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

None.

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

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 C_1––2 more prominent and distinctive. No intercostal striae. Corallum white.

Septa hexamerally arranged in four complete cycles according to the formula: S_1–2 > S₃ > S₄ (48 septa). S_1–2 equal in size and exsertness, with slightly sinuous axial margins, and almost meet columella. Remaining septa (S_3–4) becoming progressively less exsert. S₃ ~ ¾ the width of S_1–2, 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.

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

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 (S_1–2 > 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.

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

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

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

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

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.

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.

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: S_1–2 > S₃ > S₄ > S₅ (96 septa). S_1–2 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 S_1–2, also with vertical and straight axial margin. S₄ least exsert septa and ⁴/₅ the size of S_3, with straight to slightly sinuous axial margin. S₅ more exsert than S₄ and fuses to adjacent S₁–₃ 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.

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.

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.

Southern California, depth unknown (Cairns 1994a).

The type is presumed lost.

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.

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.

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.

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.

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.

Pourtalosmilia dumosa Alcock, 1902c, by original designation.

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).

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

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.

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 C_1–2.

Septa hexamerally arranged in three cycles according to the formula: S₁ > S₂ > S_3. 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.

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.

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.

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.

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

Unknown.

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

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.

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 S_1, 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 P_3. All paliform lobes (P_1–4) 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.

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.

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 C_1–2 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).

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

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

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.

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. C_1–2 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 S_1–2, 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 S_1–2 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.

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.

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.

Off Durban Harbour, South Africa, 99 m.

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.

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.

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. S_3–4 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 S_2. At calicular margin, S₄ fuses to adjacent S_1–2 forming high rectangular lancets. Septal faces granular. No pali. Fossa deep, containing a poorly developed papillose columella composed of ≤ seven sparsely arranged rods.

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

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 S_3–4 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 P_1–4 (in case of H. aequicostatus) or P_{1, 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 S_3–4 axial margins, a feature that is unique in relation to all Heterocyathus.

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

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

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.

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 C_3–4 wider than C_1–2, and progressively diminishing in size towards base. Base smooth. Upper parts of corallum, columella, and S_1–2 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 S_1–2 forming a V-shaped pattern. Pali cylindrical and bear meniane-like ridges. Fossa shallow, containing a papillose columella.

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.

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.

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.

Labyrinthocyathus langae Cairns, 1979, by original designation.