Carnivorous sponges from the Australian Bathyal and Abyssal zones collected during the RV Investigator 2017 Expedition Author Ekins, Merrick Author Erpenbeck, Dirk Author Hooper, John N. A. text Zootaxa 2020 2020-05-12 4774 1 1 159 journal article 22182 10.11646/zootaxa.4774.1.1 269cf599-428d-464d-8f21-994d69110c5c 1175-5326 3825140 B0C4A2F8-F2AB-4147-BB12-63720EEF2516 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. FIGURE 18 . Chondrocladia (Chondrocladia) clavata , QM G337491, A. Tridentate unguiferate isochelae. B. Specimens of QM G337491, the left two specimens with the ectosome removed. C. Large mycalostyle. D. Magnified ends of the mycalostyle illustrated in C. E. Smaller thinner mycalostyle. F. Magnified ends of the mycalostyle illustrated in E. G. Sigma. FIGURE 19 . Chondrocladia (Chondrocladia) clavata . A. Cross section through the body showing the obvious mycalostyle bundles that form the filaments and the disorganised subectosomal smaller mycalostyles. B. Exterior surface showing the echinating smaller mycalostyles and the isochelae on the surface. C. Two filaments showing the isochelae on the ectosome and some of the mycalostyles exposed from the endosome. D. End of filament where it joins the sponge body showing the arrangement of large mycalostyles and the sigmas and isochelae in the ectosome. E. Lower stem composed of tightly packed mycalostyles. F. Root processes showing both types of tightly packed mycalostyles. 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 ). TABLE 10. Comparative morphological and distributional data for all known species of Chondrocladia .
Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) freycinetensus sp. nov. present work Erect peduncu- late ‘crinorhiza’ form with bal- loon-shaped subglobular body, long and very thin stem, with basal rhi- zoid root-like processes 140 x 13 Axis of thick bundles of mycalostyles arising from roots and stem, then radiating outwards through the hispid body, and terminating in spherical swellings. Mycalostyles 567-(1774)-2560 x 10-(29)-65 undifferentiated undifferentiated Tridentate unguiferate isochelae 1, 51-(62)-71 isochelae 2, 22-(30)-38 absent sigmas 1, 50-(68)-86 x 2-(3)-4 sigmas 2, 15-(21)-25 x 1-(1.5)-2 Off Freycinet Peninsula, Tasmania, bathyal
Chondrocladia (Chondrocladia) callistemonex sp. nov. present work Erect stipitate unbranched stem with numerous horizontal filaments at right angles forming 4- 7 distinct columns encircling the stem, short naked stem and rhizoid basal rootless. 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of Chelae (L μm) basal attachment (LxW μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) zygainadentonis sp. nov. present work Erect arbuscular unbranched resembling a ‘Bottlebrush’ with a cylindrical stem with four columns of filaments, no basal attachment collected. 23 x 0.5 Axis with bundles of mycalostyles and subtylostyles longitudinally arranged in the axis of the stem and filaments, becoming radial where filamets inserted into the stem. Mycalostyles 1030-(1261)- 1440 x 18-(24)-36 Subtylostyles 502-(595)-652 x 8-(13)-16 undifferentiated not present Unguiferate chelae with asymmetrical long alae 45-(58)-73 x 4-(6)-9 Tridentate unguiferate isochelae 26-(35)-39 x 3-(3.5)-4 absent sigmas 25-(28)-29 x 1-(2)-3 Off Gladstone, Coral Sea, Queensland, Australia, bathyal
Chondrocladia (Chondrocladia) albatrossi Tendal, 1973 Tendal, 1973: 35 , Figs. 3–4 Erect pedunculate ‘crinorhiza’ form with globular body bearing apical and basal tentacular processes, long thin stem, basal attachment missing 13 x 150 Axis of with 2 categories of styles 1 & 2, in the body 1 category of style 3, in branches a smaller category of style 4 Styles 1, 3200-(3688)- 4000 x 32–44 Styles 2, 1440-(1898)- 2560 x 32–44 Styles 3, 800-(1774)-2240 x 13–38 Styles 4, 960-(1744) x 2080 - 2538 not present Tridentate unguiferate isochelae 1, 120-(152)-170 Quintadentate unguiferate isochelae 2, 63-(82)-113 Quintadentate unguiferate isocheleae 3, 13-(24)-44 absent sigmas 38–57 Off N Brazil, N Atlantic, abyssal
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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) amphactis ( Schmidt, 1880 ) Schmidt, 1880: 83 , Pl.10, Fig. 10 Erect pedunculate ‘crinorhiza’ form with subspherical body pointed at apex and bearing a circular ring of long filaments at centre of body, short broken stem, no basal attachment ? ? Styles 665–1750 x 6–25 undifferentiated undifferentiated Anchorate isochelae 50–55 sigmancistras absent 46 West Indies, mesophotic
Chondrocladia (Chondrocladia) antarctica Hentschel, 1914 Hentschel, 1914: 77 Taf IV Fig. 6, Taf VI Fig. 5; ( Koltun 1964: 37 ); [Goecke & Janussen 2013: 66–67, Fig. 13]; {Dressler- Alame 2017: 193, Fig. 5} Erect pedunculate, rounded body with thick single stalk, long filaments regularly spaced on apex, hispid surface, basal attachment unknown [40–70 x 20–25] Axis very dense aggregation of styles of 2 sizee classes from the stem running into the body, branching into several tracts forming filaments, surrounding axis is a thick bark-like tissue with paratangential styles Styles 2000–2560 x 45–50 (Styles 1, 2000–2560 x 45–50 styles 2, 1100) [styles 1, 1400 mean styles 2, 2400 mean] {styles 1, 1406–4484 x 20–50 styles 2, 500–4750 x 19–42} undifferentiated unknown Tridentate unguiferate anchorate isochelae 40–45 x 14 (isochelae 1, 96–112 isochelae 2, 40–45) [isochelae 1, 80-(95)-109 isochelae 2, 42-(48)-49] {isochelae 1, 60–105 x 3–10 isochelae 2, 20–55 x 2–8} 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras Sigmas (L μm) (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) arenifera Brøndsted, 1929 Brøndsted, 1929: 227 , Fig. 4 Erect stipitate dichotomously branched, partly fused, surface grooved covered by many oblique papillae and hispid surface, basal attachment unknown 30 x 4–5 Axis with thick longitu- dinal primary bundles of styles, trans- verse thinner secondary bundles, and scattered styles throughout the endosome; ectosomal skeleton with tangential styles and a sandy crust Styles 150–180 x 7 undifferentiated unknown Tridentate anchorate isochelae 1, 18–25 Tridentate unguiferate isochelae 2, 8–12 absent sigmas 12–40 Formosa Strait, South China Sea, depth unknown
Chondrocladia (Chondrocladia) asigmata Lévi, 1964 Lévi, 1964: 76 , Fig 25, Pl. 4A, C Erect stipitate arbuscularconcrescens ‘ growth form, stem twisted irregularly branching with small thorn- like lateral processes, basal attachment unknown 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) burtoni Tendal, 1973 Tendal, 1973: 36 , Figs 5–6 Erect stipitate club-shaped stick with regular wart- like structures encircling upper body in spiral rows, body hispid, tapering to thin stem with protruding basal spicules axis 170 x 6 Axis with straight bundles of larger styles, body with smaller styles Styles 1, 1920- (2460)-2880 x 25–38 Styles 2, 480- (1069)-1600 x 13–25 unknown Quadradent ate unguiferate isochelae 40-(44)-48 absent sigmas 44-(85)-113 Off Mauritiana, W Atlantic, abyssal
Chondrocladia (Chondrocladia) clavata Ridley & Dendy, 1886 Ridley & Dendy 1887: 100 (Dressler- Allame et al. 2017: 195) { Levi 1964: 76 , Fig. 26, Pl. 4D} [present study] ((Holotype examined this study)) Erect pedunculate, ‘crinorhiza’ form with suspherical body bearing long tentacular processes, long stem, multiple bifurcated root-like anchor processes 12 x 2 (15 x 15) {10 x 7} [40 x 5] Axis of bundles of mycalostyles breaking into smaller tracts below forming rootlets, and giving off radiating tracts above projecting a long way beyond the body mycalostyles 1, 1000 x 22 mycalostyles 2, absent (1, 520–1976 x 16–38 2, 218–455 x 3–10) {1, 1800–2100 x 35 2, 850–1100 x 15} [1, 633–2820 x 11–50 2, absent] ((1, 1050 x 31 2, 500–1020 x 8–14)) 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Chondrocladia) concrescens ( Schmidt, 1880 ) Schmidt, 1880: 83 , Pl.10, Figs. 8-9; Topsent 1930 ; Koltun 1970 ; (Not Levi 1993 , Not R&D 1887) ( Lopes & Hajdu 2011 ) Erect elongated to globular pedunculate club-shaped, with long lateral and apical processes with terminal (presumably inflatable) swellings and filaments, stem long and thick, basal attachment unknown up to 215 x 60 Axis of stem, body and lateral processes with bundles of styles- mycalostyles of 2 size classes styles 1, 1100–4500 x 27–65 styles 2, 550–880 x10–27 acanthosubtylostyles 220–270 x 1–4 undifferentiated undifferentiated Anchorate isochelae 1, 55–130 Anchorate isochelae 2, 18–50 69–97 (only in Schmidt’s specimen) sigmas 1, 70–165 sigmas 2, 16–55 West Indies, NW Pacific, mesophotic- haddal
Chondrocladia (Chondrocladia) coronata Hestetun, Rapp & Pomponi, 2019 Hestetun, Rapp & Pomponi, 2019: 13 , Fig. 7 Erect, single stem, filaments in radial pattern along whole stem, becoming longer at top of sponge, top stem funnel- shaped with ring-like margin and secondary crown of smaller filaments 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) crinita Ridley & Dendy, 1886 Ridley and Dendy, 1887:101 , P1. XX. Fig. 4; P1. XXI. Figs. 3, 17 ( Koltun 1970: 180 , Pl. 9, Figs. 9–13) Erect pedunculate ‘crinorhiza’ form, with large conical body with a spike- like projection at apex, fringed by long filaments at the base, and short papillae near the centre, basal attachment unknown 12 x 15 Central axis of dense spiculo-fibre forming the apical spike, with radiating bundles of mycalostyles forming axes of fringing filaments styles 2200 x 44 (mycalostyles 1,1600–300 x 38–60 mycalostyles 2, 700–1200 x 16–33) undifferentiated unknown Tridentate unguiferate isochelae 100 (isochelae 1, 100–130 isochelae 2, 50–60) absent sigmas 70 (sigmas 70–165) Indonesia & NW Pacific, bathyalabyssal
Chondrocladia (Chondrocladia) dichotoma Lévi, 1964 Lévi, 1964: 77 , Fig 29, Pl. 4H Erect arbuscular dichotomouss branched carrying secondary branches perpendicul ar to the axis spaced about 3 mm apart, basal attachment unknown 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) fatimae Boury- Esnault & van Beveren, 1982 Boury- Esnault & van Beveren, 1982: 57, Pl. 9, Fig. 34, Fig. 16 D–G Erect pedunculate with globular subspherical body covered with numerous fine filaments supported by thick short stem, with rhizoids as basal attachments 35 x 70, 120 x 60 Axis of body with radial bundles of mycalostyles extending beyond surface, with smaller thinner mycalostyles on surface, and bundles of styles are connected to each other by transverse spicules near the centre, stem and the rhizoids are formed by a large bundles of styles surrounded by a layer of isochelae mycalostyles 1, 1600–3364 x 29–45 mycalostyles 2, 728–1400 x 12–26 undifferentiated Tridentate unguiferate isochelae 1, 53–83 isochelae 2, 22–32 45–59 absent Kerguelen- Heard Plateau, subantarctic, mesophotic
Chondrocladia (Chondrocladia) gracilis Lévi, 1964 Lévi, 1964: 77 , Fig. 27 Erect stipitate pedunculate with small body 4mm high carried by a long stem, basal attachment unknown 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lat- eral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) grandis ( Verrill, 1879 ) Verrill, 1879: 204 (Hestetun, Tomkins- McDonald & Rapp, 2017: 14, Figs. 10–11) (Erect club- shaped, elongated enlarged upper part of body with numerous short or branch-like processes projecting in all directions, thick stem with basal rhizoids) 185–600 x 15–20 (Axis of stem with dense core of spirally twisted bundles of mycalostyles, clearly separated outer tissue layer supported by mycalostyles, processes with axis of overlapping mycalostyles and outer region with longitudinally arranged mycalostyles, outer layer of basal stem and rhizoids contains acanthosubtylostyles) (Mycalostyles 1, undifferentiated 1203-(1785)- 2543 x 14-(28)-51 Mycalostyles 2, 309-(919)-1360 x 7-18-31) (Acanthostyles- subtylostyles, 129-(246)-435 x 1–3) (Multidentate (6 alae) unguiferate isochelae 1, 51.9-(69.1)- 81.5 Multidentate (6–9) isochelae 2, 16.1-(23.1)- 34.0) (Sigmancistras 1, 31.4- (42.4)-59.2 Sigmancistras 2, 20.4- (22.7)-26.7) absent Boreo- Arctic, mesophotic- bathyal
Chondrocladia (Chondrocladia) guiteli Topsent, 1904 Topsent, 1904: 207–209 ( Cristobo, Urgorri & Rios, 2005: 210 , Fig. 5) (Erect pedunculate, globular elongated body with apical or central processes/ filaments, with short stem, basal attachment unknown) (22–39 x 12–25) (Axis of stem cored by polyspicular bundles of styles, body with axial bundles of styles becoming radial in the filaments) (Styles 1500–2100 x 20–22) undifferentiated unknown (Tridentate unguiferous isochelae 1, 175–210 x 5–7 Multidentate (5 alae) isochelae 2, 42–83 x 4–5 Multidentate (8 alae) isochelae 3, 12–22 x 1–2) absent (sigmas 52–66 x 2–4) NW of Cap Finisterre (Galice), & Angola Basin, SE Atlantic, abyssal
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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) koltuni Vacelet, 2006 Vacelet, Erect 2006: 557, pedunculate, Fig. 3 cylindrical stem with enlarged body with 15 radiating branches bearing terminal spherical swellings, and long rhizoid basal anchoring system 400 x 5–7 Axis of stem with longitudinally arranged fusiform mycalostyles (1), less dense in central part and coating of smaller styles (2) making a hispid surface, floats with irregularly radiating bundles of smaller styles (3), rhizoids composed of thick fibers of fusiform styles Mycalostyles 1, 420–5000 × 14–50 Styles- subtylostyles 2, 500– 750 × 6–10 Styles 3, 630–800 × 14–17 undifferentiated Multidentate (5-6alae) unguiferate isochelae 1, 98–105 x 7.5–8 Multidentate isochelae 2, 55–70 x 3.5–4.2 absent sigmas 1, 92–110 sigmas 2, 30–46 Bering I., Kuril Trench, N Pacific, abyssal
Chondrocladia (Chondrocladia) lampadiglobus Vacelet, 2006 Vacelet, Erect 2006: 559, pedunculate, fig. 4, large ovoid 17A–D body with large secondary branches terminating in spherical swellings, cylindrical stem, with rhizoid basal anchoring system 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attach- ment (LxW μm) Chelae (L μm) Sigmancistras Sigmas (L μm) (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) levii Cristobo, Urgorri & Rios, 2005 Cristobo, Urgorri & Rios, 2005: 205 , Fig. 2 Erect pedunculate, tulip or club shaped body with anastomosing central and apical processes, short conical stem, anchored by a hollow filament 55–110 x 9–20 Central axis formed by styles, which are attached to the bottom of branches by transverse bundles of styles, in the endosome which radiate to form conical processes on the surface Styles 1500–2000 x 30–40 undifferentiated undifferentiated Multidentate (5 alae) unguiferate isochelae 1, 190–212 x 6–9 Multidentate (5) isochelae 2, 67–77 x 5–6 Multidentate (9–10) isochelae 3, 16.2–20.0 x 0.6–1.5 absent absent Angola Basin, SE Atlantic, abyssal
Chondrocladia (Chondrocladia) magna Tanita, 1965 Tanita, 1965: 159 , Figs 1–3 Erect columnar arbuscular , with body covered by many thin digitate processes, surface hispid from protruding styles, basal attachment unknown 300 x 50 Axis with tightly packed spirals of longitudinal bundles of styles, with thin spicule bundles radially projecting from axis, axis of digitate processes composed of radially arranged smaller styles 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) multichela Lévi, 1964 Lévi, 1964: 77 , Fig. 28, Pl. 4B Erect stipitate pedunculate, with globular body on a long stem, body with long lateral and apical tentacular processes swollen at their ends, basal attachment unknown 100 x 5 Axis with dense longitudinal bundles of styles- subtylostyles surrounded by a detachable ectosome Subtylostyles 1, undifferentiated 2200–3100 x 45–55 Styles 2, 1500 x 30 unknown multidentate (6 alae) unguiferate isochelae 85–95 x 6 Tridentate unguiferate isochelae 25–33 absent sigmas 70–90 x 1–2 Off Kenya, bathyal
Chondrocladia (Chondrocladia) nani Boury- Esnault & van Beveren, 1982 Boury- Esnault & van Beveren, 1982: 60, Pl. 9, Fig 35, Fig. 16a–c Erect stipitate, spherical body with short conulose filaments on upper part, short thick stem, basal rhizoid attachments 340 x10 Axis of Mycalostyles 1, undifferentiated stem cored 996–1604 by twisted x 19–32 bundles of Mycalostyles 2, mycalostyles 646–899 x 4–22 1 forming longitudinal tracts, axis of body with radial bundles of mycalostyles 1 and secondary transverse spicule tracts, with shorter mycalostyles 2 closer to the surface undifferentiated Tridentate unguiferate isochelae 1, 38–51 Isochelae 2, 19–26 absent absent Kerguelen- Heard Plateau, subantarctic, esophotic
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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) nicolae Cristobo, Urgorri & Rios, 2005 Cristobo, Urgorri & Rios, 2005: 207 , Fig. 4 Erect stipitate, cylindrical stem with secondary branches on one side of the stem 47 x 2 Axis of stem and branches axes and branches formed by thick polyspicular fibres cored by mycalostyles styles Mycalostyles 1180–1550 x 18–29 undifferentiated unknown Tridentate unguiferate isochelae 37–44 x 2–6 absent sigmas 96–105 x 2–3 Angola Basin, SE Atlantic, abyssal
Chondrocladia (Chondrocladia) pulvinata Lévi, 1993 Lévi, 1993: 42 , Fig. 15 C, Pl. 3, Figs. 9–12 Erect pedunculate, globular to pear-shaped body tapering at apex, with short stem, basal attachment unknown 17–20 x 15–25 Axis with dense bundles of longitudinal styles 1 runs through the body ending in a short protruding button, axis of the stem with confused bundles of shorter styles 2 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) robertballardi Cristobo, Rios, Pomponi & Xavier, 2015 Cristobo, Rios, Pomponi & Xavier, 2015: 3 , Figs 2–5 Erect dichotomously branched, stipitate, stem with secondary branches radiating in all directions, each with a subterminal spherical swelling, anchored with a long basal rhizoid 400 x 11 Axis of stem and branches with thick parralel bundles of large mycalostyles, rhizoids composed of similar spicule bundles linked by spongin, swellings with confused fusiform styles, sometimes arranged in bundles Mycalostyles 528–2579 × 6.3–37 undifferentiated undifferentiated Multidentate (6 alae) unguiferate isochelae 52–70 × 3.4–20.3 absent absent Off Galician Peninsular, NE Atlantic, bathyal
Chondrocladia (Chondrocladia) saffronae Goodwin, Berman, Downey & Hendry, 2017 Goodwin, Berman, Downey & Hendry, 2017: 52 , Fig. 9 Erect peduncu- late, thin stalk with flattened body, basal attachment unknown 10 x 1 Stem and base composed of columns of styles, body formed of radiating columns of styles 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attach- ment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) schlatteri Lopes, Bravo & Hajdu, 2011 Lopes, Bravo & Hajdu, 2011: 431 , Figs 14–15 Erect pedunculate, spherical body on thick stem, basal rhizoid attachment 222 x 13–21 Axis of stem with thick fibres cored by large longitudinaly arranged styles 1, with coating of smaller styles 2 arranged irregularly, axis of body irregularly or longitudinally arranged larger styles 1 with ectosome of body covered by acantho(subtylo) styles, also in basal rhizoids Styles/mycalostyles 1 (body, stalk, rhizoids) 1825–4600 x 45–65 Styles/mycalostyles 2 (ectosome) 525–1600 x 28–35 Strongyles 50–390 x 8–68 Acantho(subtylo)st yles 205–359 x 5–10 undifferentiated undifferentiated Multidentate (5 alae) unguiferate isochelae 1, 65–85 Isochelae 2, 28–60 absent absent Off SE Diego Ramírez Archipelago, bathyal
Chondrocladia (Chondrocladia) scolionema Lévi, 1993 Lévi, 1993: 42 , Fig. 15 B, Pl. 4, Fig 3 Erect stipitate crinorhiza growth form, with long thin stem, body subspherical tapering to apical point, supporting ~45 long lateral filaments and other shorter filaments towards the apex, basal attachment unknown 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) vaceleti Cristobo, Urgorri & Rios, 2005 Cristobo, Urgorri & Rios, 2005: 207 , Fig. 3 Erect pedunculate ‘crinorhiza’ form, elongated conical body with thick branches/ filaments in the equatorial region, basal attachment unknown 25 x 12 Axis of stem with parallel bundles of styles, axis of branches with polyspicular fibres cored by styles Mycalostyles 950–2250 x 10–35 undifferentiated undifferentiated Tridentate (rounded alae) unguiferate isochelae 1, 60–100 x 6–9 Tridentate (pointed alae) isochelae 2, 32–52 x 3–4 absent sigmas 65–86 x 2–3 Angola Basin, SE Atlantic, abyssal
Chondrocladia (Chondrocladia) verticillata Topsent, 1920 Topsent, 1920: 12 (Pomponi & Rapp, 2016: 528, Figs. 5–7) (Erect stipitate, single cylindrical stem carrying lateral projections terminating in swellings, basal attachment unknown) (250 x 2–4) (Axis of stem made up of fibers of mycalostyles in a slightly spiraling pattern, outer layer of lower stem detachable containing acanthostyles, skeleton of branch processes composed of tightly packed mycalostyles) (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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Chondrocladia) virgata Thomson, 1873 Thomson, 1873: 187 (Hestetun, Tomkins- McDonald & Rapp, 2017: 19, Figs 12–13) (Erect stipitate, short lower stem, elongated dichotomou sly branching main upper part with large number of processes projecting in all directions, with inflatable spherical structures halfway along their length, with branched basal root) (200– 500) (Axis of stem with thick spirally twisted bundles of mycalostyles in the lower stem, more parallel in the upper stem and branches, outer layer clearly separated with more confusedly arranged mycalostyles, basal stem and roots also with acanthostyles) (Mycalostyles 1, 1200-(1646)-2321 x 17-(27.5)-36 Mycalostyles 2, 505-(800)-1277 x 14-(22)-33 undifferentiated (Acanthostyles 249-(307)-395 x 1–3) (Multidentate (6–8 alae) unguiferate isochelae 51.1-(62.1)- 71.4) (31.2- (40.4)-45.8) (absent) Ibero-Mo- roccan Gulf to the Ca- nary Islands, NE Atlantic, mesophotic- bathyal
Chondrocladia (Chondrocladia) yatsui Topsent, 1930 Topsent, 1930: 421 Erect pedunculate elongate globular body, long filaments with expanded terminations on lower portion of body, short spine-like projects on apex of body, with basal disk attachment embedded with sediment 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of Chelae basal attachment (L μm) (LxW μm) Sigmancistras Sigmas (L μm) (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Meliiderma) latrunculioides Lopes, Bravo & Hajdu, 2011 Lopes, Bravo & Hajdu, 2011: 436 , Fig. 16 Encrusting subspherical body with an enlarged basal attachment 7 x 4–5 Ectosome a dense crust of trochirhabds, endosome with irregular or vaguely mycalostyles 450–1300 x 15–30 strongyles 1, 307–800 x 25–53 undifferentiated undifferentiated Multidentate (5–6 alae) unguiferate isochelae 53–73 absent trochirhabds 35–50 x 8–15 (3 rings) Off Diego Ramírez Archipelago, Southern S America, bathyal
longitudinal tracts of mycalostyles and strongyles strongyles 2, 91–165 x 38–55
Chondrocladia (Meliiderma) occulta ( Lehnert, Stone & Heimler, 2006 ) Lehnert, Stone & Heimler, 2006: 30 , Fig. 17 Subglobular stipitate ? Unknown Mycalostyles 1200–2100 x 30–38 undifferentiated undifferentiated Multidentate (5 alae) unguiferate isochelae 75–98 absent trochirhabds 42–54 (2 rings) Central Aleutian Islands, Alaska, bathyal
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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Meliiderma) rogersi Hestetun, Rapp & Xavier, 2017 Hestetun, Erect stipitate, Rapp & pedunculate, Xavier, 2017: body spherical- 180, Fig. 11 subspherical, cylindrical stem, basal attachment unknown 20 x 4 Axis of stem composed of bundles of mycalostyles/ subtylostyles entering centre of body which is composed by a radial skeleton ending in protrusions and looser tissue in between Mycalostyles- subtylostyles 466-(912)-1050 x 13-(21)-33 undifferentiated undifferentiated Multidentate (5 alae) unguiferate isochelae 1, 63-(70)-78 isochelae 2, 42-(48)-60 16-(18)-21 trochirhabds 40-(44)-48 (2 rings) SW Indian Ocean Ridge, seamount, bathyal
Chondrocladia (Meliiderma) stipitata Ridley & Dendy, 1886 Ridley & Dendy, 1886: 344 , 1987:00:0 102–103 ( Hestetun, Rapp & Xavier, 2017: 181 , Fig. 12) Erect stipitate, pedunculate spherical body with slightly hispd surface, cylindrical stem with a disk- shaped basal attachment 36 x 2 Axis of stem solid with bundles of mycalostyles 2 entering centre of body, body with radial bundles of mycalostyles 1 and also confusedly arranged in between spicule tracts Mycalostyles 2, 695-(944)-1209 x 20-(28)-35) Mycalostyles 1, 1226-(1526)-1880 x 22-(31)-38 undifferentiated Multidentate (5 alae) unguiferate isochelae 85-(93)-103 absent subtrochirhabds (spear-like microstyles) 66-(78)-107 sigmas 53-(57)-64 Subantarctic islands, bathyal
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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral filaments or body (LxW μm) Spicules of basal attachment (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other micro- scleres Locality/ depth range
Chondrocladia (Meliiderma) tasmaniensis Vacelet, Kelly & Schlacher- Hoenlinger, 2009 Vacelet, Kelly & Schlacher- Hoenlinger, 2009: 63, Fig. 4 Erect stipitate, pedunculate, body subspherical, cylindrical stem attached by an enlarged basal disk 12–30 x 0.9–1.5 Axis of stem with tightly packed longitudinal tracts of mycalostyles and substrongyles covered by a dense sheath of closely packed trochirhabds, body with radiating bundles of mycalostyles Mycalostyles 1, 470–950 x 11–26 Mycalostyles 2, 740–950 x 15–28 Mycalostyles 3, 350–660 x 10–21 Multidentate (5 alae) unguiferate isochelae 67–90 x 5.5–5.6 absent trochirhabds 32–50 (1 ring) Tasmanian seamounts, Australia, bathyal
Chondrocladia (Meliiderma) turbiformis Vacelet, Kelly & Schlacher- Hoenlinger, 2009 Vacelet, Kelly & Schlacher- Hoenlinger, 2009: 59, Figs 2–3 Erect stipitate, pedunculate, body subspherical, cylindrical stem, no basal attachment 10–30 x 1.2–1.9 Axis of stem tightly packed longitudinal bundles of slightly sinuous mycalostyles and substrongyles, body with radiating bundles of mycalostyles, extending asan axis in the filaments 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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Species Source Morphology Total height x width (mm) Skeleton Spicules of main axis (LxW μm) Spicules of lateral Spicules of filaments or body basal attachment (LxW μm) (LxW μm) Chelae (L μm) Sigmancistras (L μm) Sigmas (L μm) and other microscleres Locality/ depth range
Chondrocladia (Symmetrocladia) lyra Lee, Reiswig, Austin & Lundsten, 2012 Lee, Reiswig, Austin & Lundsten, 2012: 262 Figs 2–12 Hetetun, Rapp & Pomponi, 2019: 15, Fig. 8 Erect, harp- shaped, with main stolons branching in 1 or more planes, and vertical branches with medial and terminal swellings bearing filaments, attached with basal rhizoids Axis of basal rhizoids, stolons and vertical branches contain a massive bundles of longitudinally oriented styles occupying most of their diameter towards the base, and less distally, filaments supported by single axial bundles of subtylostyles with their points oriented distally 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 .
TABLE 11. Comparisons of spicule measurements (µm) for some of the new specimens of Chondrocladia (Chondrocladia) clavata along with other recorded measurements.
Specimens country Depth (m) Unguiferate isochelae Large Mycalo- styles Small Mycalostyles Sigmas S Sigma C
QM G337490 Australia 2636 24-(54)-64 x 2-(5)-6, n=54 916-(1535)-2820 x 21-(30)-47, n=55 387-(763)-981 x 5-(14)-20, n=45 53-(66)-77 x 1-(2)-3, n=16 -
QM G337491 Australia 2636 30-(57)-65 x 3-(5)-7, n=49 886-(1534)-2720 x 20-(31)-46, n=40 461-(791)-975 x 9-(15)-20, n=33 39-(60)-70 x 2-(2)-3, n=11 -
QM G337500 Australia 1036 20-(44)-58 x 2-(4)-6, n=82 1070-(1767)-2260 x 21-(30)-47, n=55 513-(804)-989 x 10-(16)-20, n=21 38-(55)-66 x 1-(2)-3, n=15 -
QM G337508 Australia 2518 23-(48)-62 x 2-(4)-6, n=56 1110-(1750)-2280 x 20-(32)-50, n=36 520-(873)-1190 x 9-(15)-19, n=24 51-(64)-74 x 1-(2)-3, n=13 -
SAMA S313 Antarctica 193 38.2-(51.4)-64.8 x 2-(4)-6, n=50 858-(1438)-2000 x 15-(25)-35, n=56 486-(892)-1590 x 4-(12)-17, n=21 - 42.3-45.0)-48.2 x 2-(2)-3, n=2
SAMA S3575 Antarctica 193 37.1-(48.0)-58.7 x 3-(4)-5, n=54 971-(1390)-1710 x 16-(26)-34, n=40 138-(732)-996 x 2-(14)-21, n=24 - 40.6 x 3, n=1
Ridley & Dendy, 1887 Fiji 256 57 1000 x 22 - 44 -
Holotype : NHMUK 1887.5.2.169 (this study) Fiji 256 27.4-(50.0)-59.3 x 2-(4)-6, n =80 1050 x 31 , n=1 500-(712)-1020 x 8-(11)-14, n=11 35-(39)-40, n=4 -
Burton, 1929 Antarctica 450 24, 48 1260-2000 x 24 - 45 -
Lévi, 1964 Madagascar 4820 30-55 1800-2100 x 35 850-1100 x 15 40 75
40-50 x 3
Koltun, 1964 Antarctica 434 24-57 1000-1260 x 22-24 present 44-45 -
Bergquist, 1972 New Zea- land 990 39-(69)-80 x 5 508-(1170)-1560 x 17-(23)-28 300-(327)-340 x 4.5 60-(78)-89 -
Dressler- Allame et al., 2017 Antarctica 840 42-70 x 4.0-8.9 520- 1975 x 16 -38 218-455 x 3.5-10.2 45-56 32-58
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.