Chondrocladia (Chondrocladia) clavata Ridley & Dendy, 1886

Figures 18 & 19, Tables 10 & 11

Chondrocladia clavata Ridley & Dendy, 1886: 345; Ridley & Dendy, 1887: 100, Pl. XX, Figs. 1, 1a, Pl. XXI, Fig. 11; Burton, 1929: 431; Koltun, 1964: 38; Lévi, 1964: 76, Fig. 26, Pl. IV(d); Bergquist, 1972: 125, Figs. 1–3.

Chondrocladia (Chondrocladia) clavata: Dressler-Allame et al., 2017: 195, Fig. 7

Material examined: Holotype: NHMUK 1887.5.2.169, Kandavu, Fiji, Station 174, 19° 6’ 00” S, 178° 14’ 20” E, 256 m, Coll. H.M.S. Challenger, 2 specimens, 3/viii/1874. Other material: QM G337490 off Jervis Bay, News South Wales, Australia, 35° 19’ 58.8”– 35° 19’ 55.2” S, 151° 15’ 28.9”– 151° 12’ 50.4” E, 2650– 2636 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_ V03, Sample 56-250, 20 specimens, 29/v/2017. QM G337491 same collection details as QM G337490, Sample 56-252, 4 specimens. QM G337500 off Hunter Valley, News South Wales, Australia, 32° 28’ 44.4”– 32° 30’ 25.1” S, 152° 59’ 38.4”– 152° 59’ 27.6” E, 1006–1036 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_ V03, Sample 69-247, 2 specimens, 3/vi/2017. QM G337508 Central Eastern Commonwealth Marine Reserve, News South Wales, Australia, 30° 5’ 51.7”– 30° 7’ 9.5” S, 153° 53’ 55.3”– 153° 52’ 28.2” E, 2429–2518 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_ V03, Sample 86-187, 1 specimen, 5/vi/2017. SAMA S313, Southern Ocean, off Enderby Land, Antarctica, BANZARE Station 41, 65° 48’ S, 53° 16’ E, 193 m, Large Monegasque Trawl, Coll. BANZARE, 24/i/1930. SAMA S3575, same collection details as SAMA S313. NHMUK 1926.10.26.152a, Off Glacier Tongue, about 8 miles north of Hut Point, McMurdo Sound, Antarctica, Terra Nova Station 316, 77° 45’ S, 166° 24’ E, 348–457 m, Agassiz Trawl, Coll. RV Terra Nova, 9/ii/1911. NHMUK 1926.10.26.153a, same collection details as NHMUK 1926.10.26.152a. NHMUK 1926.10.26.154a, 5 miles north of Inaccessible Island, McMurdo Sound, Antarctica, Terra Nova Station 314, 77° 36’ S, 166° 12’ E, 406–441 m, Agassiz Trawl, Coll. RV Terra Nova, 23/i/1911.

Distribution: This species has been recorded from many locations in the Southern hemisphere including: Fiji (Ridley & Dendy 1886, 1887), Antarctica (Burton 1929; Koltun 1964, Dressler-Allame et al. 2017), Madagascar (Lévi 1964), New Zealand (Bergquist 1972, Kelly et al. 2009), and Eastern Australia (present study), bathyal depth.

Description:

Growth form: An erect pedunculate ‘crinorhiza’ form sponge with a subspherical body bearing long tentacular processes, supported by a long thin stem, terminating in long bifurcated root-like anchor processes. The body often has an apical nipple, or tuft of mycalostyles (Figure 18 B). Specimens range from 40–44 mm long, with subspherical body 6–9 mm diameter, tentacular process 9–26 mm long, 0.2–0.6 mm diameter, the stem is a tapering cone 1.5–7 mm in diameter and 3–9 mm in length, basal root processes 16–27 mm long, 0.2–0.5 mm diameter. Antarctic specimen SAMA S313 is a maximum of 56 mm long, with subspherical body 16 mm diameter, tentacular process 11 mm long, 0.3–0.5 mm diameter, the stem is 1mm in diameter and 35 mm in length, and has no basal root processes.

Another Antarctic specimen SAMA S3575 has a total length of 26 mm, with the stem 18 mm in length and 1 mm in diameter, a subspherical body of 18 mm in diameter, also with no basal root processes. The remainder of the tentacular processes are only a maximum of 1.5 mm in length. This species has been well described by previous authors and matches with previous descriptions.

Colour: Light brown with white stalk, light yellow to tan with ectosome removed.

Ectosomal skeleton: The ectosome consists of a crust of isochelae and sigmas (Figures 19 A–D). In the body of the sponge this is penetrated by the smaller mycalostyles from the ectosome producing the hispid surface (Figure 19 B).

Endosomal skeleton: The endosome in the body consists of the radially orientated bundles of large mycalostyles that form the filaments. Subectosomally there is a confused predominantly radial assortment of the smaller mycalostyles penetrating through the ectosome to the surface. The endosome also contains the isochelae and the sigmas (Figure 19 A). The endosome of the filaments consists of the parallel mycalostyles in an extremely well organised bundle (Figure 19 C–D). The endoosome of the stem and root processes also consist of a tightly bound longitudinally orientated mycalostyles.

Megascleres: Styles in two categories, large mycalostyles (916– 2820 x 20 –50 µm), and thinner supporting styles (387– 1190 x 5–19 µm), (see Table 11). The large mycalostyles occur in the stem, the body and the filaments. The smaller styles occur mainly in the body radially orientated and penetrating the ectosome producing the hispid surface of the body, but also occur in the filaments and the stem and roots. These styles could also be described as a continuous range of the one type, as was probably done in the original description.

Microscleres: Tridentate unguiferate anchorate isochelae (20–65 x 2–7 µm) in two sizes, but with intermediate sizes, and sigmas (39–75 x 1.5–3 µm) (see Table 11).

80–120 x 2–5 Axis with tightly bound bundles of styles forming the stem, roots and filaments. Styles 1, 1030–2970 x 19.2–53.4 Styles 2, 372–1490 x 7.0–20.0 undifferentiated undifferentiated Tridentate unguiferate isochelae 49.8–65.8 x 2.4–6.5 35.9–63.1 x 1.2–3.0 absent Off Newcastle, New South Wales, Australia, abyssal

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absent sigmas 38–57 Off N Brazil, N Atlantic, abyssal

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absent sigmas 96–160 (sigmas 96–160) [sigmas 1, 89-(110)-143 sigmas 2, 44-60-75] {sigmas 1, 80–105 x 3 sigmas 2, 25–55 x 2–3} Antarctica, bathyal-abyssal

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10 Axis with longitudinal multispicular bundles of larger mycalostyles and smaller mycalostyles Mycalostyles 1, 2000–2100 x 40–45 Mycalostyles 2, 600–750 x 15–18 undifferentiated unknown Multidentate (up to 7) unguiferate isochelae 1, 65–70 x 6–9 Tridentate unguiferate isochelae 2, 28–32 x 3–4 absent absent Kermadec Trench, bathyal

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undifferentiated undifferentiated Tridentate unguiferate isochelae 57 (43–70 x 4–9) {30–55} [20–65 x 2–7] ((27–60 x–6)) absent sigmas 44 (32–58) {1, 40 2, 75 x 3} [50–75 x 1.5–3] ((35–40)) E Australia, New Zealand, Fiji, RossSea, Madagascar, Antarctica, mesophotic- abyssal

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100–150 Main skeleton of stem and filaments with a core of longitudinal tightly packed mycalostyles, outer surface of stem and top part with subtylostyles mycalostyles 1430– 2104 x 27–52 subtylostyles 488–948 x 10–28 undifferentiated undifferentiated Anchorate isochelae 1, 41–60 Anchorate isochelae 2, 21–32 Anchorate isochelae 3, 15–20 34–49 absent Northwest Guam Seamount, Marianas, NE Pacific, bathyal

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7–22 x 1–1.5 Axial skeleton cored by dense longitudinal bundles of mycalostyles echinated by smaller subtylostyles mycalostyles 1700 x 32 subtylostyles 800–850 x 25–29 undifferentiated unknown Quadrodentate uniguiferate isochelae 45–50 x 2 absent absent Seychelles, bathyal-abyssal

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20–26 x 0.5 Not described Mycalostyles 650–850 x 15–20 Subtylostyles 550 x 13–14 undifferentiated unknown Tridentate unguiferate isochelae 40 x 3 absent sigmas present, size unknown Seychelles, abyssal

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absent (sigmas 52–66 x 2–4) NW of Cap Finisterre (Galice), & Angola Basin, SE Atlantic, abyssal

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500 Axis of stem with parallel bundles of large mycalostyles (1), with a thick cover of rugose tylostyles (2) arranged in a dense irregular feltwork, body and secondary branches with a rugose coating of styles smaller (3), swellings with confused smaller fusiform styles Mycalostyles 1, 700–4750 × 15–75 Rugose tylostyles 2, 300–535 × 5–6 Mycalostyles (3) 510–580 × 17–30 unknown Multidentate (6) unguiferate isochelae 1, 123–140 x 6–7 Multidentate isochelae 2, 20–32 x 2 absent sigmas 45–120 × 2–3 East Pacific Rise, bathyal

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Styles 900–2000 x 20–35 undifferentiated undifferentiated Multidentate (4–5 alae) unguiferate isochelae (size unknown) absent absent Off Kushiro & Miyako, Japan, mesophotic

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undifferentiated Tridentate unguiferate isochelae 1, 38–51 Isochelae 2, 19–26 absent absent Kerguelen- Heard Plateau, subantarctic, esophotic

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Styles 1, 1400–1550 x 28–30 Styles 2, 700–900 x 20–22 undifferentiated unknown Tridentate isochelae 88–90 120–140 absent Off New Caledonia, bathyal

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Styles 1098–1424 x 25–43 undifferentiated unknown Multidentate (4–5 alae) unguiferate isochelae 74–85 absent Sigmas 1, 45-(47)-49 Sigmas 2, 66-(73)-89 Sars Seamount, Drake Passage, S America, mesophotic

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90 x 13 “skeleton is typical, with a layer of peripheral styles” Styles 1, 1400–2500 x 20–30 Styles 2, 700–850 x 12–20 undifferentiated undifferentiated Tridentate unguiferate isochelae 50–58 x 10 absent absent Off New Caledonia, bathyal

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(Mycalostyles 728-(1606)-2815 x 13-(27)-38 Mycalostyles 2, 420-(589)-1240 x 9-(15)-27 Acanthostyles 192-(310)-500) (undifferentiated) unknown (Multidentate (6–7 alae) unguiferate isochelae 1, 41-(65)-78 Multidentate (5–6) isochelae 2, 13-(18)-34 absent sigmas 16-(20)-27 Caribbean, meso- photic-bathyal

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220 Axis of stem with twisted spiral longitudinal bundles of styles, axis of filaments with bundles of larger styles perpendicular to the body, basal disk and outer sheath of stem with acanthostyles Styles 750–2800 x 18–50 Acantho(subtylo) styles 285–455 x 4–5 undifferentiated undifferentiated Multidentate (5 alae) unguiferate isochelae 90–100 x 6 45–55 x 2–3 sigmas 22–25 x 1–5 Sagami Bay, Japan, mesophotic

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Mycalostyles 1, 1000–1700 x 15–40 Mycalostyles 2, 340–1400 x 4–15 undifferentiated Substrongyles/ subtylostrongyles 260–1288 x 12–30 Multidentate (5–6 alae) unguiferate isochelae 1, 68–95 x 5 Isochelae 2 (5 alae), 32–51 x 2 absent trochirhabds 30–50 (3 rings) sigmas 30–50 Chatham Rise seamounts, New Zealand, mesophotic- bathyal

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Styles 1, 1410–1880 x 20–31 Styles 2, 1570–1700 x 9–12 Subtylostyles undifferentiated Tridentate 650–680 x 14–15 unguiferate isochelae 42–44 absent sigmas 1, 67–80 sigmas 2, 44–53 Escanaba Ridge and Monterey Canyon, off California, USA and Marianna NW Pacific, bathyal-abyssal

Molecular data: The 28S sequence of QM G337490 is provided in the Sponge Barcoding Database under accession number SBD#2307 and the molecular difference to other congenerics displayed in Figure 3.

Remarks: These specimens match the previous descriptions of Ch. (Ch.) clavata Ridley & Dendy (1887), and more recently Dressler-Allame et al. (2017) who described some specimens from the Weddell Sea, Antarctica. Examination of the holotype confirmed the size ranges of the isochelae to be similar to the new specimens from this expedition. Our specimens are more similar to the Fijian type specimen in lacking the small mycalostyles (Styles II) described in the Antarctic specimens by Dressler-Allame et al. (2017), and are closer to the medium-small sized mycalostyles described by Lévi, (1964) from Madagascar. Unusually, published records suggest that the species has a wide bathymetric distribution, extending from the shallow mesophotic (e.g. 250m, Ridley & Dendy 1886), bathyal (1060–2650 m, present study), to abyssal depths (e.g. 4820 m, Lévi 1964). Molecular studies may reveal whether these differences are enough to warrant the separation of Ch. (Ch.) clavata into several distinct species. This species also resembles Ch. (Ch.) crinita Ridley & Dendy (1886) in external morphology, but has much smaller isochelae. It also bears some resemblance to Cladorhiza nematophora Lévi, 1964, but lacks the tridentate unguiferate anisochelae, having instead tridentate unguiferate isochelae.