Chondrocladia (Chondrocladia) clavata Ridley & Dendy, 1886

Ekins, Merrick, Erpenbeck, Dirk & Hooper, John N. A., 2020, Carnivorous sponges from the Australian Bathyal and Abyssal zones collected during the RV Investigator 2017 Expedition, Zootaxa 4774 (1), pp. 1-159 : 89-113

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 ).

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.

Figures

FIGURE 1. Distribution of the carnivorous sponges collected during this expedition.

FIGURE 3. Maximum-likelihood phylogram of the DNA sequences of this study in comparison with other, published cornivorous sponges. Sequences from taxa of this study are in bold. Number at the branches are RAxML rapid bootstrap values. Numbers following the taxa are either QM registration numbers (QM G3xxxxx) or NCBI Genbank accession numbers. Scale bare are substitutions per site. Please note that the reconstruction does not represent a phylogeny as due to the variability of the 28SrRNA gene C-Region barcoding marker no unambiguous alignment is possible and clipping ambiguous sites would remove diagnostic characters for taxon distinction. The tree therefore indicates molecular differences rather than a robust phylogeny.

FIGURE 7. Cladorhiza australis nov. sp. Holotype QM G337456. A. Larger multidentate (3–4 upper alae) ‘cleistochelate’ anisochelae. B. ‘Cleistochelate’ anisochelae showing four upper alae. C. Smaller tridentate ‘unguiferate’ anisochelae. D. Sigma. E. Large mycalostyle. F. Magnified ends of the mycalostyle illustrated in E. G. Medium-sized mycalostyle. H. Magnified ends of the mycalostyle illustrated in G. I. Small mycalostyle. J. Magnified ends of the small mycalostyle illustrated in I. K. preserved holotype. L. preserved roots of the holotype.

FIGURE 11. A. Type of Axoniderma mirabile, BMNH 1887.5.2.141. B. Pseudoamphiaster and an unguiferate anisochelae. C. Pseudoamphiaster. D. Pseudoamphiaster and an unguiferate anisochelae. E. Sigma. F. Section through a filament showing the encrusting pseudoamphiasters in the ectosome and two sizes of styles in the endosome. G. Section through the end of the filament showing the external layer of pseudoamphiasters and the anisochelae in the endosome. H. Section through the body of the sponge showing the intersection of the stem and the filament rays, with encrusting pseudoamphiasters. I Section through the body with ectosome of pseudoamphiasters and anisochelae in the endosome.

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.

FIGURE 26. Lycopodina calyx (Hentschel, 1914). A. SEM of the sponge body QM G337513 B. Palmate anisochelae. C. Mycalostyle 1. D. Magnified ends of the mycalostyle 1 illustrated in C. E. Mycalostyle 2. F. Magnified ends of the mycalostyle 2 illustrated in E. G. Mycalostyle 3. H. Magnified ends of the mycalostyle 3 illustrated in G. I. SEM of the exterior of the sponge body (calyx), showing the large mycalostyles underneath the protecting medium mycalostyles. J. SEM of the inside of the cup. K. Preserved specimen QM G337463. L. Preserved specimen QM G337513. M. Preserved specimen QM G337476.

Tables

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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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