Lycopodina calyx ( Hentschel, 1914 )

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 : 123-141

Lycopodina calyx ( Hentschel, 1914)

Fig. 26 , Tables 14 & 15

Asbestopluma calyx Hentschel, 1914: 66 , Pl. IV, Fig. 4 , PL. V, Fig. 11 ; Burton 1932: 293;

Burton 1934: 24; Koltun 1964: 32, Pl. V, Figs 8–12 .

Asbestopluma (Asbestopluma) cf. calyx: Lopes & Hajdu 2014: 333 , Fig. 3 .

Lycopodina calyx: Goodwin et al. 2017: 54 , Fig. 11 ; Dressler-Allame et al. 2017: 200, Fig. 13 .

Material examined: QM G337513 off Central New South Wales, Tasman Sea, Station 88, New South Wales, Australia, 30° 15’ 50.4”– 30° 17’ 12.1” S, 153° 52’ 12”– 153° 49’ 48.7” E, 4481– 4401 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_ V03 , Sample 88-141, 2 specimens, 6/vi/2017 . QM G337463 same collection details as QM G337513 , Sample 88-141.2 . QM G337476 , off Gippsland , Tasman Sea, Station 33, Victoria, Australia, 38° 31’ 15.6”– 38° 29’ 52.8” S, 150° 12’ 46.8”– 150° 12’ 25.2” E, 4107– 4064 m, Brenke epibenthic Sledge, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_ V03 , Sample 33-156, 24/v/2017 . SAMA S314 , off Kaiser Wilhelm Land , Antarctica, BANZARE Station 100, 65° 48’ S, 89° 49’ E, 393 m, large rectangular dredge, Coll. BANZARE, 3/v/1931 .

Distribution: This species is a Southern Ocean species known predominately from the mesophotic, bathyal and abyssal depths of Antarctica ( Hentschel 1914, Burton 1932, 1934, Koltun 1964, Dressler-Allame et al. 2017), Drakes Passage ( Goodwin et al. 2017), Campos Basin, SE Brazil, SW Atlantic Ocean ( Lopes & Hajdu 2014), and now off Southern and central Eastern Australia, Tasman Sea, at abyssal depth.

Description:

Growth form: A delicate erect cup-shaped pedunculate sponge on a thin stem, with a slightly expanded basal attachment ( Figure 26 A , K–M). Stems measure 7–30 mm long, 0.5–1 mm diameter; sponge bodies are elongated, 3–7 mm long, 3–4 mm in diameter. Apex of the body is a shallow convex cup.

Colour: Beige on deck and in ethanol.

Ectosomal skeleton: The ectosome on the exterior surface of the cup-shaped body is composed of a palisade of radially projecting mycalostyles 3, formed into vertical columns tangential to the surface all orientated towards the cup aperture ( Figure 26 A , I–J). These are overlaying the two other sizes of mycalostyle in the endosome in parallel. There are also anisochelae incorporated into the ectosome. The ectosome on the interior surface is covered with anisochelae ( Figure 26 J ). The ectosome of the stem is also covered with the anisochelae and perforated by the smallest mycalostyles 3, forming a hispid surface.

Endosomal skeleton: The endosome of the body is composed of ribs of longitudinally projecting large mycalostyles 1, formed into vertical columns tangential to the surface all orientated towards the cup aperture, together forming a palisade. This palisade is overlaid by the smaller mycalostyles 2, which in turn are overlaid by the smallest mycalostyles 3, in the ectosome, all in parallel. The endosome of the stem is a series of longitudinally orientated mycalostyles 1.

Megascleres: There are three size classes of mycalostyles. The largest mycalostyle 1 with tapering points, slightly subtylote or tylote bases, and with their largest diameter in the midsection of the spicule (511-(667)-911 x 9.6-(16.7)- 30.1 µm, n=29) ( Figure 26 C–D).

Mycalostyle 1 provides all the support longitudinally aligned in the stem and forms the structural component of the axis of the calyx body. The medium sized mycalostyle 2, also with tapering points, slightly subtylote or tylote bases, and with their largest diameter in the midsection of the spicule (371-(464)-566 x 6.0-(8.7)-11.6 μm, n=12) ( Figure 26 E–F).

Mycalostyle 2 only occurs in the calyx surrounding the mycalostyle 1 and projecting all the way to the lip of the calyx. The smallest mycalostyle 3 occurs in the calyx as a protective sheath over mycalostyles 2 and at right angles to the longitudinally mycalostyles 1 in the stem, producing the hispid stem (128-(174)-234 x 2.6-(4.1)-6.1 μm, n=15) ( Figure 26 G–H). ( Table 15 ).

Microscleres: Anisochelae have three larger upper palmate alae with the two lateral alae fully attached to the fimbriae, and three smaller basal arcuate alae, partially or nearly fully fused together, and with serrations on the upper edges of the alae ( Table 15 ) ( Figure 26 B ).

Molecular data: It was not possible to get unambiguous molecular data from the present material

Remarks: These specimens from the Tasman Sea conform to the description and measurements of the predominately Antarctic species, L. calyx , as described by Hentschel (1914). The spicule measurements also fall within the range of those published by other authors ( Koltun 1964, Lopes & Hajdu 2014, Dressler-Allame et al. 2017, Goodwin et al. 2017). The general morphology of these specimens differ from those of Goodwin et al. (2017) in that the external surface of the cup is composed entirely of subtylostyles ( Figure 26 A , I–J), and it is only the inside of the cup that contains all the anisochelae ( Figure 26 J ). This species currently has a Southern circum-polar distribution and may possibly be found to extend into lower latitudes as deep-ocean sampling continues.

Figures

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 4. Abyssocladia annae sp. nov. A. Abyssochelae. B. Subtylostyle. C. Magnified ends of the subtylostyle depicted in B. D. Sigmancistra. E. Holotype QM G337611. F. SEM showing the abundant abyssochelae and sigmancistras in the ectosome and the underlying subtylostyles in the endosome.

FIGURE 8. Cladorhiza australis nov. sp. Holotype QM G337456.A. Stem of the sponge, showing the carpet of ‘cleistochelate’ anisochelae, and where they have been abraded off the underlying sheath of the ectosome. B. Close-up of the ‘cleistochelate’ anisochelae in A. C. Stem of the sponge with the ectosome peeled off the underlying mycalostyle bundle of the endosome. Note the mycalostyle contained in the ectosome. D. Two filaments, the uppermost filament has part of the ectosomal layer containing the ‘unguiferate’ anisochelae removed showing the mycalostyles in the endosome.

FIGURE 9. Cladorhiza poritea sp. nov. A. Tridentate ‘unguiferate’ anisochelae. B. Asymmetrical pseudoamphiaster. C. Variation in the alae of the pseudoamphiasters. D. Large mycalostyle. E. Magnified ends of the mycalostyle illustrated in D. F. Medium styles. G. Magnified ends of the style illustrated in F. H. Small styles. I. Magnified ends of the style illustrated in H. J. Section through the body and stem of the sponge with one of the filaments on the right. K. Paratype QM G337437. L. Holotype QM G337401. M. Paratype QM G337544.

FIGURE 10. Cladorhiza poritea sp. nov. A. Body of the sponge showing the upper and lower ectosomal surface with the endosomal radiating mycalostyles in between, there are two of the filaments which have been inadvertently folded back over the sponge body. B. Magnified body surface showing the abundant pseudoamphiasters, and the underlying anisochelae. C. Stem showing the encrusting pseudoamphiasters in the ectosome. D. L.S. stem interior with the pseudoamphiaster crust and the styles forming an outer sheath, and an internal cored of styles. E. External layer of the filaments with the anisochelae exterior to the pseudoamphiasters. F. L.S. filament interior showing a sheath of styles similar to the stem and a hollow core.

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 12. A. Cladorhiza investigator sp. nov. Holotype QM G337458. B. Large anisochelae. C. Small anisochelae. D. Large Mycalostyle. E. Magnified ends of the mycalostyle illustrated in D. F. Medium Mycalostyle. G. Magnified ends of the mycalostyle illustrated in F. H. Subtylostyles. I. Magnified ends of the tylostyle illustrated in H. J. Sigma. K. Sigmancistra.

FIGURE 13. Cladorhiza investigator sp. nov. Holotype QM G337458. A. External view of the filaments showing the tightly packed anisochelae in the ectosome. B. The ectosome of the filaments exposed revealing the mycalostyles in the endosome. C. The external surface of the stem showing the ectosomal plates formed by the mycalostyles and the almost alternate juxtaposition of the filaments. D. Internal T.S. of the stem showing the mycalostyles buttressing the filaments after they have emerged from the confused core. In addition it shows the longitudinal tracks of mycalostyles. E. Cross section of the stem, showing the nine radial filaments intersecting in the core of the stem, and the thin wedges of anisochelae and sigmas between these arms. F. Branching roots of the sponge, showing an almost absent ectosome, but the tight bundles of mycalostyles and three thin subtylostyles in the upper branch.

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 14. Comparative morphological and distributional data for all known species of Lycopodina.

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)	other microscleres (LxW μm)	Forceps (LxW μm)	Locality/ depth range
Lycopodina nikitawimandi sp. nov.	present work	Erect stipitate, with multiple filaments covering the upper 80% of the stem, bulbous holdfast with protruding spicules	28 x 1–2	Axis of stem with longitudinal bundles of larger mycalostyles 1 only found in the stem, and smaller mycalostyles 2 occuring throughout the sponge including stem, filaments and basal holdfast	Long mycalostyles 1, 1040–910 x 12–37 Small mycalostyles 2, 209–992 x 3–19	small mycalostyles 2	small mycalostyles 2	palmate anisochelae 9–18 x 2–6	absent	absent	Central E Coast of Australia to Tasmania, bathyalabyssal
Lycopodina cassida sp. nov.	present work	Erect, pedunculate, of ‘crinorhiza’ form, with short stem supporting radial upper section of filaments crowned by a pointed apex, basal attachment unknown	3 x 2	Axis of stem and filaments with longitudinal bundles of subtlyostyles	Subtylostyles 374-(536)-1220 x 4-(7)-15	undifferentiated	unknown	larger palmate anisochelae 16-(19)-22 x 4–11 smaller anisochelae 9–15 x 3–5	absent	absent	Off Fraser Island, Queensland, Australia, bathyal
Lycopodina brochidodroma sp. nov.	present work	Erect, central axis with 4 long columns of lateral filaments on opposite sides, alternating every second row, basal attachment unknown	20 x 1	Axis with longitudinal bundles of mycalostyles and secondary subtylostyles, and subtylostyles also occur as supporting structures where the filaments attach to the main axis	Mycalostyles 621-(1364)-1816 x 7-(19)-33 Subtylostyles 202-(285)-328 x 4-(5)-8	undifferentiated	unknown	palmate anisochelae (11-(13)-15 x 2-(3)-5	absent	absent	Off Moreton Island, Queensland, Australia, abyssal

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TABLE 15. A comparison of spicule measurements (µm) between the different records of Lycopodina calyx.

Specimens	Anisochelae	Mycalostyles 1 (axis of stem and body)	Mycalostyles 2 (axis of body)	Mycalostyles 3 (dermis of stem and body)
Hentschel, 1914	14–21 x 7–8	456–592 x 8–9	296–392 x 10	200–256 x 6
Koltun, 1964	14–21	456–600 x 8–9	290–390 x 10	200–256 x 6
Dressler-Allame et 14-(22)-33 al., 2017		600-(733)-850 x 15-(26)-32	400-(500)-650 x 9-(13)-30	150-(226)-340 x 6-(9)-12
Goodwin et al., 2017	20–25		326-(496)-588 x 9-(12)-14	278-(379)-533 x 7-(10)-14
Lopes & Hajdu, 2014	13-(16)-25	287-(517)-729 x 8-(9)-15	226-(390)-657 x 8-(9)-13 185-(560)-821 x 3-(7)-10	92-(186)-257 x 8-(8)-10
QM G337513	16.3-(19.8)-22.6 x 8.5-(10.2)-12, n=55	511-(667)-911 x 9.6-(16.7)-30.1, n=29	371-(464)-566 x 6.0-(8.7)-11.6, n=12	128-(174)-234 x 2.6-(4.1)-6.1, n=15